Fungus Flora of Tropical Africa
Volume 2
MONOGRAPH OF LACTARIUS IN TROPICAL AFRICA
by Annemieke Verbeken & Ruben Walleyn
Plates / Planches
Acknowledgements
Introduction
Material and Methods
Characters used for the identification of Lactarius species
1 Macroscopical characters
1.1 Basidiocarps
1.2 Pileus
1.3 Lamellae
1.4 Stipe
1.5 Context
1.6 Latex
2 Microscopical characters
2.1 Basidiospores
2.2 Basidia
2.3 Pseudocystidia
2.4 True cystidia
2.5 Marginal cells
2.6 Hymenophoral trama
2.7 Structures of pilei- and stipitipellis
Distributional patterns
Abbreviations
Lactarius Pers.
Brief history of Lactarius collections from tropical Africa
Practical key to sections of the genus Lactarius in tropical Africa
Sect. Lactariopsis (Henn.) Singer
Key to tropical African species
Species descriptions
1. Lactarius annulatoangustifolius (Beeli) Buyck
2. Lactarius zenkeri (Henn.) Singer
3. Lactarius pelliculatus (Beeli) Buyck
4. Lactarius heimii Verbeken
5. Lactarius velutissimus Verbeken
Sect. Chamaeleontini Verbeken
Key to tropical African species
Species descriptions
6. Lactarius indusiatus Verbeken
7. Lactarius chamaeleontinus R. Heim
8. Lactarius madagascariensis Verbeken & Buyck
9. Lactarius sesemotani (Beeli) Buyck
10. Lactarius laevigatus Verbeken
11. Lactarius pruinatus Verbeken & Buyck
12. Lactarius uapacae Verbeken & Stubbe
13. Lactarius emergens Verbeken
14. Lactarius brachystegiae Verbeken & C. Sharp
Sect. Russulopsidei Verbeken
Key to tropical African species
Species descriptions
15. Lactarius ruvubuensis Verbeken
16. Lactarius longipes Verbeken
17. Lactarius cyanovirescens Verbeken
18. Lactarius urens Verbeken
19. Lactarius rufomarginatus Verbeken & Van Rooij
20. Lactarius pseudotorminosus R. Heim
21. Lactarius claricolor R. Heim
22. Lactarius corbula R. Heim & Gooss.‑Font.
Sect. Edules Verbeken
Key to tropical African species
23. Lactarius edulis Verbeken & Buyck
24. Lactarius nodosicystidiosus Verbeken & Buyck
25. Lactarius latifolius Gooss.‑Font. & R. Heim
26. Lactarius aureifolius Verbeken
27. Lactarius densifolius Verbeken & Karhula
28. Lactarius inversus Gooss.‑Font. & R. Heim
29. Lactarius phlebophyllus R. Heim
30. Lactarius roseolus Verbeken
Sect. Aurantiifolii Verbeken
Species description
31. Lactarius aurantiifolius Verbeken
Sect. Rubroviolascentini (Singer) Verbeken
Key to tropical African species
Species descriptions
32. Lactarius rubroviolascens R. Heim
33. Lactarius denigricans Verbeken & Karhula
Sect. Pseudogymnocarpi Verbeken
Key to tropical African species
Species descriptions
34. Lactarius gymnocarpoides Verbeken
35. Lactarius pseudogymnocarpus Verbeken
36. Lactarius longisporus Verbeken
37. Lactarius pumilus Verbeken
38. Lactarius medusae Verbeken
39. Lactarius carmineus Verbeken & Walleyn
40. Lactarius luteopus Verbeken
Sect. Rugati Verbeken
Key to tropical African species
Species descriptions
41. Lactarius volemoides Karhula
42. Lactarius pseudovolemus R. Heim
43. Lactarius kivuensis Verbeken
44. Lactarius rubiginosus Verbeken
45. Lactarius xerampelinus Karhula & Verbeken
Sect. Polysphaerophori Singer
Key to tropical African species
Species descriptions
46. Lactarius gymnocarpus R. Heim ex Singer
47. Lactarius flammans Verbeken
48. Lactarius albocinctus Verbeken
49. Lactarius tanzanicus Karhula & Verbeken
50. Lactarius goossensiae Beeli
51. Lactarius foetens Verbeken & Van Rooij
52. Lactarius brunnescens Verbeken
Sect. Phlebonemi R. Heim ex Verbeken
Key to tropical African species
53. Lactarius phlebonemus R. Heim & Gooss.‑Font.
54. Lactarius angustus R. Heim & Gooss.‑Font.
55. Lactarius arsenei R. Heim
56. Lactarius pisciodorus R. Heim
57. Lactarius nonpiscis Verbeken
Sect. Chromospermi Verbeken
Species description
58. Lactarius chromospermus Pegler
Sect. Piperites (Fr. ex J. Kickx f.) Burl. s.l.
Key to tropical African species
Species descriptions
59. Lactarius barbatus Verbeken
60. Lactarius acrissimus Verbeken & Van Rooij
61. Lactarius afroscrobiculatus Verbeken & Van Rooij
Sect. Amari Verbeken
Key to tropical African species
62. Lactarius subamarus Verbeken
63. Lactarius amarus R. Heim
Sect. Russularia Fr. ex Burl.
Species description
64. Lactarius hepaticus Plowr.
Sect. Nigrescentes Verbeken
Key to tropical African species
Species descriptions
65. Lactarius melanogalus R. Heim ex R. Heim
66. Lactarius baliophaeus Pegler
67. Lactarius orientalis (Verbeken) Verbeken
68. Lactarius griseogalus R. Heim
Sect. Plinthogali (Burl.) Singer s.l.
Key to tropical African species
Species descriptions
69. Lactarius angiocarpus Verbeken & U. Eberh.
70. Lactarius dolichocaulis (Pegler) Verbeken & U. Eberh.
71. Lactarius pulchrispermus Verbeken
72. Lactarius miniatescens Verbeken & Van Rooij
73. Lactarius sulcatulus Verbeken
74. Lactarius adhaerens R. Heim
75. Lactarius desideratus Verbeken & Stubbe
76. Lactarius kalospermus (Beeli) Verbeken & Walleyn
77. Lactarius congolensis Beeli
78. Lactarius melanodermus R. Heim & Gooss.‑Font.
79. Lactarius nudus R. Heim
80. Lactarius rumongensis Verbeken
81. Lactarius sulcatus Verbeken & Walleyn
82. Lactarius saponaceus Verbeken
83. Lactarius pusillisporus Verbeken
84. Lactarius kabansus Pegler & Piearce
85. Lactarius sciaphilus Verbeken & C. Sharp
86. Lactarius tenellus Verbeken & Walleyn
87. Lactarius acutus R. Heim
88. Lactarius pseudolignyotus R. Heim
Sect. Pseudofuliginosi Verbeken
Key to tropical African species
Species descriptions
89. Lactarius atro-olivinus Verbeken & Walleyn
90. Lactarius undulatus Verbeken
90.1 Lactarius undulatus var. undulatus
90.2 Lactarius undulatus var. rasilis Verbeken
Unclassified taxa
91. Lactarius badius Verbeken
92. Lactarius caperatus R. Heim & Gooss.‑Font.
93. Lactarius hispidulus R. Heim
94. Lactarius russuliformis (Beeli) Verbeken
Insufficiently known taxa
1. Lactarius fulgens R. Heim
2. Lactarius fulgens var. africanus R. Heim
3. Lactarius striatus R. Heim
Excluded taxa
1. Lactarius lignyotus ss. Dade
2. Lactarius pellucidus Gooss.‑Font. & R. Heim
3. Lactarius piperatus ss. Pegler & Piearce and Piearce
4. Lactarius trivialis ss. To & Ob
5. Lactarius uvidus ss. Bresadola. & Saccardo and De Wildeman & Durand
6. Lactarius vellereus ss. Morris & Williamson
7. Lactarius velutinus Bres.
8. Lentinus clitocyboides Henn.
Translation of the keys into French
References
Taxonomic index
In this book 96 Lactarius species of tropical Africa, including Madagascar, are described, as well as 2 accepted varieties. Except for Lactarius hepaticus, an introduced species which is only found in exotic Pinus plantations, all taxa in the region are endemic for the area.
A recently published molecular phylogeny of the Russulaceae (Buyck & al. 2008) leads to the conclusion that the genus Lactarius as currently defined is not monophyletic and falls apart in two different genera, while one American species, Lactarius furcatus, is transferred together with the Russula species previously assigned to R. subsection Ochricompactae Bills & O.K. Mill. in the seperate new genus Multifurca Buyck & Hofstetter. As the nomenclatural consequences of these results are still pending, the genus Lactarius is presented here in a more traditional concept. However, the sequestrate lactarioid genera Arcangeliella Cavara, Zelleromyces Singer & A.H. Sm. and Gastrolactarius J.M. Vidal are considered synonyms of Lactarius as been shown by molecular research (Nuytinck & al. 2003, Eberhardt & Verbeken 2004, Lebel & Tonkin 2007).
Material and Methods
The results of this monograph are based on studies of the author’s collections from Burundi, Cameroon, Malawi and Zimbabwe and herbarium specimens from BR, E, FH, GENT, H, K, LG, LY, PC (including an important number of collections previously cited as ‘herbarium B. Buyck’), WAG and material sent by D. Arora, A. Bâ, C. Sharp and D. Thoen. In total about 1400 collections were studied, about 1220 were identified and cited in this monograph. Nearly all literature records of Lactarius have been checked when documented with voucher material, except for the species cited by Onguene (2000) which are therefore not considered in this book.
In the macroscopical descriptions the colours are described according to Kornerup & Wanscher (1978).
Microscopic features were studied from dried material, mainly in Congo-red in L4 (Clémençon 1972), sometimes in NH4OH. Old or bad material was eventually pretreated in KOH 10% solution. Spore ornamentation is described and illustrated as observed in Melzer's reagent. Line-drawings were made using a drawing tube at original magnifications 6700× or 6000× for spores, 3200× or 3000× for individual elements and 1100× or 1000× for sections and surface views. Stippling indicates refractive content in cystidia and lactifers or intracellular pigmentation in the elements of pilei- and stipitipellis. Basidia length excludes sterigmata length. Spores were measured in side view in Melzer's reagent, excluding the height of the ornamentation (Fig. 1f, g).
Measurements are given as (MINa)[AVa-2*SD]–AVa–AVb–[AVb+2*SD](MAXb), with AVa = lowest mean value for the measured collections, AVb = greatest mean value and SD = standard deviation. Q corresponds with spore length/width ratio and is given as (MINQa)Qa–Qb(MAXQb) with Qa and Qb being the lowest, respectively the highest, mean ratio for the measured collections.
The nomenclature of infrageneric taxa follows Redeuilh & al. (2001).
Characters used for the identification of Lactarius species
1 Macroscopical characters
The vast majority of African Lactarius species are agaricoid with a distinct stipe and pileus and a hymenophore consisting of well developed lamellae. Size and consistency are very variable: species with large and firm basidiocarps occur, as well as tiny and brittle species. Besides the agaricoid type, a few sequestrate Lactarii also are known in tropical Africa: a distinctly gasteroid species, Lactarius angiocarpus, and one epigeous species with a distinct pileus and stipe but with a sinuate and glebulose, somewhat radiating to alveolate labyrinthuloid hymenophore.
Most agaricoid Lactarius species have a gymnocarpic development, but some species of L. subgenus Lactariopsis have a partial veil covering the young hymenium. This velum is called a secondary velum by Reijnders (1963) as it originates after the development of the hymenium started and in fact is a fusion of the parallel covering layers of the stipe and the pileus (mixangiocarpic development; Heim 1937, 1938b, Reijnders 1963).
The basidiocarps are characterized by the exudation of latex, in some specimens very scarce or presumably absent, in others extremely abundant.
1.2 Pileus
Shape. Most species have a planoconvex to infundibuliform pileus, the shape varies strongly with the developmental phase. In contrast with temperate Lactarii, few umbonate species occur. A papilla is sometimes present (Lactarius griseogalus). In most species, the margin is first incurved, later straight or deflexed. Some species have a conspicuously crenate or crenulate margin (Lactarius undulatus, L. sulcatus).
Size. The diameter of the pileus may vary from 1 (or less, e.g. Lactarius roseolus and L. desideratus) to 15 cm (L. edulis, L. brunnescens, L. latifolius). In general, rain forest species are usually smaller and more brittle than species growing in woodlands.
Colour. There is a large variation in colours, but the main colours can be described as rather dull, especially when compared to many members of the genus Russula: brown, orange-brown, reddish brown, dark brown to greyish or buff. The colour of the pileus is an important character at species level, and sometimes it might also give some indications on sectional or subgeneric level. However, one has to keep in mind that colours are liable to changes, depending on the age of the basidiocarps and the weather conditions. The typical golden yellow to orange pileus margin in young basidiomes of Lactarius edulis disappears soon as the basidiocarp matures. Some species have peculiar additional tinges in dried condition; e.g. L. edulis, which shows no red colours in the field, becomes reddish in exsiccatum. Hygrophaneous species are less common in the tropics than in temperate regions, but do occur (e.g. Lactarius rumongensis).
Surface. The surface aspect of the pileus is often related to the microscopic structure of the pileipellis and, therefore, has to be considered as an important taxonomic character. Some common types of surface aspects in temperate species, such as a zonate pileus or a hairy cap margin, are very rare or almost lacking in tropical African species. Only Lactarius aurantiifolius and L. subamarus have a zonate cap, and only L. barbatus has a hairy cap margin. Another character common for temperate species and lacking in the studied area is a glutinous or slimy pileus. Species with a viscid cap are Lactarius barbatus, L. brachystegiae while L. chromospermus is greasy when humid only.
Rugose, rimulose and finely or even strongly wrinkled pileus surfaces are very common in tropical African Lactarius-species. They occur especially in Lactarius subg. Lactifluus, but also in L. subg. Plinthogalus, where the pileus is usually more velvety. In some species there are even remarkable veins which inspired Heim to define a separate subgenus Venolactarius. Another common type is a dry pileus which becomes cracked and areolate when the basidiomes grow older (especially found in L. section Edules). Dry and finely velvety, slightly pruinose pilei seem to be the rule, while smooth and glabrous, rather shiny pilei occur sporadically. A peculiar type is observed in Lactarius section Lactariopsis where a secondary velum occurs. In some representatives this veil is remarkably thick and felty. Stellate rupturing towards the margin shows a smooth and glabrous layer, which is striate or translucently striate at the margin.
Strongly pruinose pilei are present in Lactarius aurantiifolius (whitish pruinose) and in L. medusae (velutinate aspect reminding the temperate species L. vellereus).
1.3 Lamellae
Attachment. Attachment of the lamellae is taxonomically not important, as in almost every species the lamellae are adnate to subdecurrent, sometimes becoming decurrent when the basidiocarp is older.
Colour. The colour of the lamellae varies mostly from white, whitish, cream-colour to pale yellowish. Some African species have however exceptionally bright coloured lamellae: bright yellow to greenish yellow in Lactarius luteopus, dark yellow to golden yellow in L. aureifolius, bright orange in L. aurantiifolius. Lactarius pulchrispermus has pale brown lamellae. The colour of the mature lamellae of Lactarius chromospermus exceeds the traditional limits of the genus significantly: they are chocolate-brown by the dark-coloured spores which makes the basidiocarps almost appear like an Agaricus species.
Lamellar edge. The lamellar edge is mostly entire and concolorous, but dark grey or brown in some species of Lactarius sect. Plinthogali (conspicuous in L. saponaceus), white and fimbriate in L. aurantiifolius. Lactarius medusae has a remarkable yellowish orange lamellar edge. The colour of the lamellar edge is a good help to recognize some species but should be considered with care as it seems not constant in all species.
Spacing and branching. The density of lamellae varies from very crowded to very distant. It is an important character on species-level (e.g. Lactarius densifolius vs. L. latifolius in L. sect. Edules) and sometimes on sectional level (e.g. dense lamellae in L. section Phlebonemi, distant lamellae in L. section Polysphaerophori). Branching of the lamellae occurs often near to the apex of the stipe and this kind of irregular branching is not taxonomically important. Frequently branching lamellae are observed in Lactarius phlebophyllus. Anastomosing lamellae or venation between the lamellae may be important on species level. Lamellulae are present in all species. They often occur in a regular pattern: often three lamellulae between two lamellae, the middle lamellula being the longest one.
1.4 Stipe
Attachment. In some species, such as Lactarius pulchrispermus, the stipe is often eccentric, but in most species the stipe is regular and centrally inserted. No representatives of L. section Panuoidei Singer, which have a lateral or no stipe, are known in tropical Africa.
Shape. The shape of the stipe is mostly cylindrical, sometimes slightly clavate or tapering downwards. The shape does not seem taxonomically important, and not even a stable species character; however there are some species with a remarkably long stipe (Lactarius longipes), a very short and thick stipe (L. edulis) or an almost pointed stipe which tapers downwards (L. aurantiifolius).
Colour and surface. The stipe is in most cases concolorous with the pileus or slightly paler; an exception is Lactarius luteopus with a vividly yellow to greenish yellow stipe. The surface of the stipe is comparable to the pileus-surface: glabrous, polished, felty or sometimes with veins. Glutinous stipes are lacking in the African tropics; fibrillose or squamulose stipes are rare. At the base of the stipe a subiculum, which is mostly tomentose and paler than the stipe, may be observed. A scrobiculate stipe is only present in Lactarius afroscrobiculatus.
Annulus. In Lactarius sect. Lactariopsis, species with an annulus occur. The annulus is membranaceous, thin, whitish and not mobile. L. indusiatus has a very fugitive annulus. Annulate species are only known from Central and South America and Africa.
1.5 Context
Colour and colour-changes. The colour of the context varies from whitish to cream-colour or pale yellow. Just beneath the pellis the context may have the same colour (or slightly paler) as the pellis, but this colour has no taxonomic importance. A colour-change of the context is often related to the colour or colour-change of the latex (see further). Some remarkable and unique colour and colour changes occur: blackening context in Lactarius section Nigrescentes or context changing to red, violet or black in L. sect. Rubroviolascentini. Species with a slightly pinkening or reddening context occur, as well as species with a strongly browning context.
Consistency. In many species the consistency is firm and solid, but not elastic, rather crumbling. The context in the stipe is solid, fistulous or hollow. A regular chambered stipe, as known in numerous Russula species, occurs in Lactarius aurantiifolius.
Smell and taste. Smell and taste are less constant characters than in temperate regions. Distinct smells as Pentamogium bugs, spices, Pelargonium, apple, coconut ... are up to now not found in the African tropics. There are some species with a distinct smell of fish (Lactarius pisciodorus, L. nonpiscis), but the range of smells is limited to fish-like, sperm-like or sweetish and fruity. The same is true for the taste of context and latex. In the genus it is very important to distinguish between the taste of the latex and the taste of the context, where this is not given attention, data on the taste of the context are ambiguous. Numerous species described by Heim (1955a) are based on collections from Mme Goossens-Fontana. The data on the taste have to be considered with great suspicion. The use of quinine probably influenced her taste capacity and therefore, a lot of species are described to be acrid or very acrid when they are possibly just mild (Heinemann, pers. comm.). I observed in the field that the taste in some African Lactarius species differs considerably from specimen to specimen (L. saponaceus may taste acrid, astringent or soap-like). Other species, like Lactarius urens and L. aureifolius were always burning acrid and their acridity exceeds everything tasted in other continents.
1.6 Latex
Presence and abundance. The major macroscopic character which distinguishes Lactarius from Russula is the presence of latex. Fresh Lactarius basidiomes, under normal conditions, exude latex when the basidiocarp is broken or bruised. In the tropics, the latex is often scarce or even lacking in certain specimens, because of dry conditions or an older state of the basidiome (a phenomenon also known from temperate regions). As several species are known from very few collections, it is possible that for some species the latex has not yet been observed. The occasional virtual absence of latex, together with other aberrant macroscopic characters (presence of an annulus, lignicolous habit) may confuse the identification of this genus in the field.
Colour and colour change. Colour and colour change of the latex are characters traditionally used to classify the species within the genus. Nowadays, it is accepted that a classification based on such characters is very artificial, but still it remains an important taxonomic character.
Usually, the latex is originally white or watery, remaining so or changing colour. Some species have latex which is coloured from the beginning; bright orange, red, green or yellow latex is not observed in tropical African species. White latex, changing to yellow, is common in European and North American species but not known in tropical Africa. The latex of Lactarius undulatus var. rasilis dries greyish to yellowish, but never turns bright yellow. The most important colour change occurs in species such as L. baliophaeus and L. orientalis; the latex is watery and translucid, becoming bright red and finally greyish or whey-like. In other species of this section the latex turns from whey-like to greyish and black. In Lactarius rubroviolascens, the latex is transparent or pale reddish grey and changes to reddish and black. In several species of Lactarius section Phlebonemi the latex is brownish and stains the context brown. The colour of the context may change under influence of the latex; this may also happen when apparently unchanging latex dries up on the context: in L. cyanovirescens the dried latex and the surrounding context become bluish green.
In some temperate species a colour change of the latex is observed when bringing the latex on a white tissue or paper. I tried this out several times during my fieldwork in Africa, but only observed a reaction in L. desideratus, recently described from Cameroon. In this species the milk turns distinctly yellow on white tissue. When observing latex colours it is important to observe the fresh mushroom over some hours.
Taste. Often latex is scarce, and its taste difficult to observe separately from the context. In Lactarius urens and L. aureifolius however, the latex is so abundant that it is easy to observe that the latex is as hot and burning as the context. The latex of these species is even so acrid that it causes a burning and irritating feeling in contact with the skin, which lasts for hours.
2 Microscopical characters
2.1 Basidiospores
Spore deposit. Spore deposit colours in Lactarius vary from white to dark buff, but the African L. chromospermus outlines these limits by far with its dark chocolate brown spore deposit (which becomes paler brown after drying). Apart from L. chromospermus, the darkest spore print in tropical African species is observed in L. pulchrispermus. Obtaining spore prints in the tropics appears not easy (basidiocarps rotten quickly etc.) and data are far from complete. Spore deposit colours noted by Mme Goossens-Fontana should be interpreted with care since she usually took a piece of a lamella and rubbed it on a white paper, thus obtaining the so-called spore colour. When she took a real spore deposit, she imitated the colour by a water-colour and threw the spore deposit back into nature (Heinemann, pers. comm.).
Size and shape. The spores are bilaterally symmetric along the longitudinal axis, the adaxial side is applanate or slightly convex, sometimes even slightly depressed, while the abaxial side is strongly convex. Only length (distance of the longest axis) and width (distance of the shortest axis when observed in profile) are measured. The spore ornamentation is not included in these dimensions (Fig. 1f, g & h).
The overall shape of the spores is determined by the length:width ratio, the quotient (Q). Globose spores are defined as having Q ranging from 1.00 to 1.05, subglobose as Q = 1.06 to 1.12, ellipsoid 1.13 to 1.39 and elongate spores have a Q bigger than 1.40, this according to Buyck (1991) but without making the distinction between broadly and clearly ellipsoid spores.
The size and the shape of the spores are remarkably constant throughout Russulaceae. The range according to Hesler & Smith (1979) is 5–12 × 5–9 mm, with very few species having spores outside this range. Among the African taxa, the longest spores are found in Lactarius carmineus (11–13.5 × 8–10 µm) and L. longisporus (8.5–12.5 mm), the shortest ones in L. pusillisporus (5.3–7.1 mm) and L. fulgens var. fulgens (4.7–5.8 µm, but material in very bad condition). The broadest spores are these from Lactarius subamarus (7.7–9.3 mm) and the narrowest ones from L. badius (width 4.0–5.6 mm) and L. fulgens var. fulgens (3.8–5 µm, but material in very bad condition). No strictly bisporic species are known in tropical Africa.
The spore shape varies from subglobose to ellipsoid, rarely elongate. In the African taxa, the average Q varies from 1.03–1.05 (globose) in L. pulchrispermus to 1.6 and even more (elongate) in L. longisporus. Most species have a Q value around 1.15 to 1.30 (ellipsoid).
Ornamentation. The amyloid spore ornamentation is a typical feature for the Russulales and a very important character in the delimitation of species and often also characteristic for higher taxonomical levels. The patterns of the amyloid ornamentation range from isolated low warts to a completely winged and heavy reticulum. Describing the highly differentiated spore ornamentation in the different species has never been easy. Josserand (1941) stated that "the ornamentation of the different species shows such a variety of decorative combinations that one may wonder if it is not the inflexibility of a wrong mind wanting to define them rigorously. It's like demarcating a moving thing." Romagnesi (1967) agrees that words are insufficient to describe an ornamentation.
Given the difficulties in giving an accurate and unambiguous description of the spore-ornamentation, drawings of Russulaceae spores are indispensable. The use of Scanning Electron Microscopy contributes to the study of the spore ornamentation ["the additional magnification and resolution possible with the SEM leaves no doubt about the characters of the spore surface previously suggested by the light microscope" (Homola & Kimball, 1975)], without derogation to the value of drawings made by light microscope, which are more practical when it comes to identifications and observation of the amyloidity. To describe this complex character following characters are used:
• Maximal height of spore ornamentation – only the maximal height is taken into account as there are always lower ornamentations in between the higher ones. The lowest ornamentation in the African taxa is found in species like Lactarius rubroviolascens (unmeasurable with a light microscope, lower than 0.1 mm), the highest in L. nudus (up to 3 mm).
• Shape of the spore ornamentation – in the case of isolated ornamentation units it is easy to determine whether they are acute and sharp or rounded, rather truncate or cylindrical or even hemispherical. When the ornamentation is composed of ridges, the shape is usually considered as sharp, but in some species the ridges have distinctly rounded edges.
• Type of the spore ornamentation – this considers the shape of the basis of the most important units of the ornamentation; these may be linear, irregular or rounded (or in case of almost smooth spores: not recognizable).
• Connections of the spore ornamentation – the connections may be crest-like, meaning that they are as high as the basic units, and as a result the basic units are mostly not recognizable any more. The connections may be of a lower height and are then observed as fine lines connecting the basic units. The connections may be lacking, then the basic units are isolated.
• Reticulation of the spore ornamentation – the overall view of the ornamenation can be described as completely reticulate, partly reticulate or not at all reticulate. Besides this we also distinguish zebroid spores.
In some species, aberrantly ornamented spores are observed. These spores may have the same size as normal spores, but the ornamentation is aggregated into droplet-like, globose warts, which are irregular in size and number. Some of these warts may even become loosened from the spores and float free in the mount as amyloid globules, which was also observed by Hesler & Smith (1979). Heim (1955a: fig. 3.15) illustrated such a spore for Lactarius melanogalus. I observed them in numerous African (e.g. Lactarius melanogalus, L. corbula) and several European and Asian species. The phenomenon occurs occasionally and does not seem to be associated with any species in particular. Malençon (1931) used these abnormal spores to support his hypothesis about the destructive origin of the spore ornamentation (e.g. Malençon, 1931: Pl. 4, fig. 3.6).
Plage. At the adaxial side, above the hilar appendix or apiculus, there is a differentiated zone called the suprahilar plage, where the normal ornamentation is usually absent or very reduced. The amyloid reaction of this plage is an important taxonomical feature. The plage can be non-amyloid (naked or almost naked, Fig. 1a), centrally amyloid (with a small amyloid or faintly amyloid spot in the centre, this type often occurs in lowly ornamented spores, Fig. 1b), distally amyloid (only amyloid in the part closest to the ornamentation, often occurring in highly ornamented spores, Fig. 1c, 1d) and totally amyloid (plage is completely amyloid, a very common type in Russula but rare in Lactarius, Fig. 1e).
2.2 Basidia
Though as in most Agaricales s.l., the size and the shape of the basidia are no important taxonomical features, there seems to be a relation between the volume of the spores and the volume of the basidia. In the large-spored species (Lactarius barbatus, L. undulatus var. undulatus), long and broad basidia occur. The smallest basidia among the African species (30–45 mm) are found in L. pusillisporus and L. badius, both species with remarkably small spores (Fig. 2). There could also be a relation between the ornamentation-type of the spore and the basidium type. Very long and slender basidia often produce lowly ornamented spores, while most of the strongly winged or zebroid spores of several members of L. sect. Plinthogali are formed by broad and almost clavate basidia with rather slender sterigmata.
The length of the basidia is measured without sterigmata and the width is measured at the largest place. According to Hesler & Smith (1979), the basidia in Lactarius should be at most 60 mm long and 14 mm wide. In the African species however, there are several species with basidia up to 80 mm long. The measurements are indicated as a certain range with extreme values in between brackets, no average values are given. The basidia are observed in the middle part of the lamellae, not very close to the margin (where they are significantly smaller). The longest basidia are found in Lactarius edulis (60–80 mm), the broadest in L. undulatus var. undulatus (11–14 mm), the shortest in L. badius (32–40 mm) and the narrowest in L. aureifolius and L. badius (6–8 mm).
As a rule, basidia of agaricoid Lactarius species have four sterigmata. Up to now, only the European L. acerrimus is known to have exclusively 2-spored basidia. Two-spored basidia are sometimes observed in other species but the percentage of these 2-spored basidia is usually very small. I observed 2-spored basidia in Lactarius inversus (abundant), L. longipes, L. acutus, L. adhaerens, L. annulatoangustifolius and some other species (Fig. 3a, 3b). In Lactarius inversus also 1-spored basidia are present (Fig. 3c), and occasionally even 5-spored basidia were observed (L. luteopus and L. kabansus, Fig. 3d, 3e). Sequestrate taxa are often 2-spored, but both African species are 4-spored.
The sterigmata may be curved or straight, but this character varies from basidium to basidium and is not constant for a species, not even for a basidiome. Especially in 2-spored basidia, the sterigmata may be deformed and extremely long (though they still produce a spore). Sometimes deformed basidia resemble deformed pleuromacrocystidia, which could be interpreted as transitional stages. This is especially observed in Lactarius chromospermus (Fig. 3f). The longest sterigmata are found in Lactarius urens (8–10 mm).
2.3 Pseudocystidia
Pseudocystidia are extremities of laticiferous or differentiated hyphae, ascending in the hymenium where they may be emergent or not. In Lactarius basidiocarps they are always present, though sometimes very scarce or hard to observe, and form in fact the best distinction with the genus Russula, where they are always absent.
The content is comparable with the content of the hyphae of the laticiferous system: refringent, dense, oleiferic or needle-like to granular, yellowish in water or KOH, greyish in sulfovanillin.
The shape varies from cylindrical to fusiform and more spectacular shapes (such as corkscrew-like or branching) occur more commonly in African species than in species of other continents; the top is rounded or tapering and sometimes even forked. The presence of these pseudocystidia is a generic character and thus not taxonomically important within the genus, but the shape, the size and the emergence are important characters at infrageneric level. They may occur both on the face (pleuropseudocystidia) (Fig. 4) and on the edge (cheilopseudocystidia) (Fig. 5) of the lamellae. Extremely emergent and broad pleuropseudocystidia occur in L. subg. Lactariopsis. The broadest pseudocystidia are observed in Lactarius pelliculatus and L. zenkeri.
2.4 True cystidia
True cystidia sometimes have the same content as the laticiferous system (then called macrocystidia) but they are never connected with a lactifer, so a delimitating septum is always present. The presence, abundance, type and size are important characters to recognize a species. One third of the African taxa has true cystidia, belonging to the following types:
• Lamprocystidia are cystidia with thickened walls (Fig. 6a, 6b). They are usually emergent, cylindrical and rounded on top. They originate in the subhymenium (exceptionally in the trama, as in the temperate species Lactarius volemus). Emergent lamprocystidia are typical for L. subg. Lactifluus. The largest cystidia belong to this type: 110–140 ´ 10–15 mm in Lactarius rubroviolascens and 80–130 ´ 8–12 mm in L. gymnocarpoides. Because of their large size, lamprocystidia may sometimes be observed with a handlens, also in the dried material. Some species (Lactarius aurantiifolius, L. baliophaeus, L. melanogalus) have slightly thick-walled cystidia, embedded in the hymenium, with the same shape as the basidioles.
• Macrocystidia with a needle-like content (Fig. 7), slightly greyish in sulfovanillin, occur in Lactarius pulchrispermus, L. nudus and L. chromospermus. They are fusiform and tapering or moniliform on top. In Lactarius chromospermus and L. pulchrispermus the cystidia have slightly thickened walls and could be called lampromacrocystidia. These cystidia are sometimes septate. The typical capitate, thin-walled macrocystidia which are so common in Russula and in temperate Lactarius species do not occur in tropical African Lactarius species. Rounded, cylindrical macrocystidia with slightly thickened walls occur in L. urens. I also use the term macrocystidia for these cystidia which have exactly the same size and shape as the pseudocystidia in the same species, but without granular, oleiferic or needle-like content (observations on the content are here based on dried material). These cystidia often arise very deep in the subhymenium or in the trama. They are present in Lactarius badius, L. densifolius and L. ruvubuensis.
• Leptocystidia are thin-walled cystidia without remarkable content (Fig. 6c, 6d, 6e) and thus only deviating by their shape. They are tapering at the top and often even have a rostrate apex, which makes them easy to confuse with monosterigmatic basidia. One can consider them to be cystidia if they are regularly observed and if they never bear a spore or spore primordium. They occur in Lactarius indusiatus and L. acutus.
2.5 Marginal cells
Pseudocystidia or true cystidia may occur at the edge of a lamella and are then called cheilopseudocystidia and true cheilocystidia; they resemble the pleurocystidia present in a same species, but are usually smaller. Their presence on the lamellar edge does not implicate that the edge is sterile, on the contrary, it usually means that the hymenium, as it occurs at the lamella-sides, prolongs at the edge and thus contains cystidia among basidioles and basidia. However, in numerous Lactarius species the edge is sterile and consists of sterile elements which occupy the entire edge. These elements differ from the pleurocystidia and are in fact "hairs" sensu Romagnesi. They are called marginal cells here. They are mostly thin-walled and hyaline, but may also contain a remarkable dark content (in species with a dark lamellar edge, e.g. L. congolensis). The shape varies from clavate, fusiform to irregular. The cheilocystidia are sometimes branching or septate. In a few cases they are long, cylindrical and narrow. Marginal cells are often comparable (concerning shape, thickness of the wall and pigmentation) to the elements of the stipiti- or pileipellis occurring in the same species (e.g. L. aurantiifolius, L. saponaceus) (Fig. 8).
2.6 Hymenophoral trama
The hymenophoral trama is bordered by the usually well developed subhymenium, mainly composed of subisodiametric cells. Lactifers are especially present in the hymenophoral trama of Lactarius, but also occur in the lamellae of Russula (Fig. 9f).
Where Russula is traditionally distinguished from Lactarius by the presence of rosettes (clusters of sphaerocytes) in the hymenophoral trama, this difference seems hardly maintainable. Not only do all Lactarius species contain single globose cells in the trama, but some species also show distinct rosettes. This is true for the European Lactarius volemus, (Fig. 10a, 10b) but also for several African species which have a hymenophoral trama that looks exactly like an average Russula-trama (sections Polysphaerophori, Russulopsidei, Lactariopsidei). On the other hand, Buyck (1989b) discovered that tropical Russulae without sphaerocytes in the hymenophoral trama exist: in Russula oleifera var. levecqui, the trama of the lamellae is entirely filamentous, without any sphaerocyte, which confirms again that the absence or presence of sphaerocytes in the hymenophoral trama is not useful as distinction between the two genera.
A hymenophoral trama composed of sphaerocytes is here called cellular (Fig. 11b), a trama composed of hyaline hyphae (with some occasional globose cells) filamentous (Fig. 11a). A mixed hymenophoral trama is observed in some species. The lamellar trama is observed on mature basidiomes and the sections of the lamellae are made halfway the pileus. The structure of the hymenophoral trama is considered to be taxonomically very important and is a constant character in many sections.
2.7 Structures of pilei- and stipitipellis
Especially the structure of the pileipellis is a very important phylogenetic character and it is also reflected in the macroscopical aspect of the basidiocarp. By looking at the pileus surface (both macro- and microscopical), one can tell in most cases which subgenus the species belongs to. These macroscopical features of the pileus (viscid, velvety, tomentose, squamulose) were since long used as important characters to delimitate certain groups, but it lasts until 1980 before the microscopic characters of the pileipellis play a major role in the identification key, as well as in the classification system.
It is true that the structure of a pileipellis and stipitipellis may be variable within a species according to the developing stage and according to the place on the basidiome, but the variation is limited; a cutis may become an ixocutis or a trichoderm but not a cellular structure. Another frequently observed phenomenon is that thick-walled ascending elements are much denser when a basidiome is young but may become apparently less frequent when the basidiocarps are older. In this study pilei- and stipitipellis structures are observed on mature, but not too old basidiomes; drawings were made halfway the pileus radius or halfway the stipe height.
The pellis typology and terminology used in this study is explained here. To characterize a pellis-structure four basic characters have to be observed:
• Presence of thick-walled elements – In tropical species the presence of thick-walled and mostly hair-shaped elements is much more common than in temperate species. Elements are considered to be thick-walled when the walls measure 1 mm or more. The presence of thick-walled elements is reflected in the name of the structure by adding the prefix “lampro”, for example in lamprotrichoderm and lampropalisade. In most cases the thick-walled elements form an entire upper layer, but there are some particular cases where they only occur scattered in a pileipellis structure consisting of thin-walled elements, I call this latter type "mixed"; it is observed in L. sect. Chamaeleontini.
• Presence of isodiametric cells – A distinct layer of isodiametric cells may be present, isodiametric cells may also occur scattered in the pellis-structure or they may be totally absent. The scattered occurrence of isodiametric cells is often due to the inflation of parts of hyphae. A distinct layer of isodiametric cells is mostly basal, but in rare cases the whole pileipellis consists of isodiametric cells.
• Orientation of the terminal elements – The upper layer of the pellis consists of cylindrical terminal elements, which may also be undifferentiated hyphal endings, these may be arranged periclinic, anticlinic or obliquely in relation to the underlying surface.
• Presence of a slime-layer – Especially in the case of a cutis or trichoderm the hyphae may secrete slime or the walls become gelatinized which results in a visible slime-layer embedding the upper layer. As in tropical Africa the group with slimy, viscid or glutinous pileus is restricted, the presence of a typical covering slime layer is rare. In the tropical species some other slime-layers can be observed: in Lactarius sulcatulus there is a very thin and well-delimited slime-layer covering the pileipellis which is a hymeniderm. It is not clear how this slime originated and as such a structure is only observed in one species, no special name is given to this type. In another species, L. undulatus, the underlayer has a slimy appearance.
The combination of these four characters leads to the following types, between which intermediate forms may of course occur:
– Pellis without inflated elements or sphaerocytes, composed entirely of filamentous elements
a. cutis: the upper layer consists of mostly hyaline and thin-walled hyphae which are pericline, recumbent and parallel or slightly intermixed. Sometimes the only difference with the underlying pileustrama is a more dense and compact structure of the pellis. Differentiated structures are mostly lacking, although pileocystidia with a special shape or content may occur (Fig. 12a, 14a).
b. ixocutis: the hyphae of the cutis may secrete slime or the walls of the hyphae gelatinize, in dried material it is impossible to detect which of the two processes occurred, maybe they even occurred together; this results in a slime layer wherein parallel, very thin and hyaline hyphae are visible (Fig. 12b, 14b).
c. trichoderm: the pellis consists of mostly hyaline and thin-walled hyphae, but at least part of them are ascending and then anticline or oblique; in the most typical case all terminal elements are anticline and form a dense turf of "hairs" (Fig. 12c, 14c).
d. ixotrichoderm: same structure as a trichoderm but the hyphae are gelatinized or secrete slime, which results in being embedded in a slime-layer (Fig. 12d).
e. lamprotrichoderm: the ascending, anticline terminal elements are clearly differentiated, mostly by being thick-walled (Fig. 12e, 14d).
– Pellis with a distinct layer of inflated elements or sphaerocytes
f. irregular epithelium: cellular structure, pseudoparenchymatic structure, the upper layer is completely composed of isodiametric cells, which are irregularly ordered (Fig. 12f).
g. regular epithelium: subcellular structure, pseudoparenchymatic structure, the upper layer is completely composed of isodiametric (or almost isodiametric) cells, these cells are not irregularly ordered, but more or less arranged in anticline rows (Fig. 12g, 14e).
h. palisade: consists of an upperlayer of anticline, elongated, hair-shaped elements and an underlying dense and compact layer of isodiametric cells (Fig. 12h, 15a).
i. lampropalisade: a palisade with thick-walled terminal elements (Fig. 13a, 15b).
j. hymeniderm: special case of palisade where the terminal elements are not elongate and septate (as is often the case in a palisade) but clavate; the underlying layer of isodiametric cells is often rather thin, the whole structure reminding thus somewhat a hymenium (Fig. 13b, 15c).
– Pellis with isodiametric cells or sphaerocytes, but these never forming a distinct layer
k. trichopalisade: between a trichoderm and a real palisade; this may look like a trichoderm where parts of the ascending, anticline hyphae are inflated or almost rounded; it may also be a structure where some of the terminal elements are ascending from isodiametric cells, others are ascending from hyphae (Fig. 13c, 15d).
l. lamprotrichopalisade: a trichopalisade where the terminal elements have thickened walls (Fig. 13d, 15e).
• Pigmentation – Extracellular pigmentation usually needs to be observed on fresh material, this type of incrustrations has only been reported for Lactarius arsenei and L. latifolius (Heim 1938b, 1955a). The intracellular pigmentation is visible over a longer period, even in older exsiccata. Many of the African species, especially of L. subg. Plinthogalus, have intracellular pigmentation in the pileipellis: the elements of the upper layer have a remarkable brownish content.
• Dermatocystidia – Dermatocystidia usually occur in a cutis or ixocutis and are very common in Russula species, but never occurring in temperate Lactarius species. In African Lactarius however, dermatocystidia occur in almost all members of L. sect. Russulopsis and in L. chromospermus (Fig. 9a, 9b, 9c).
Distributional patterns
In this study material from 19 different countries is included (Benin, Burundi, Cameroon, Central African Republic, Congo, Gabon, Ghana, Guinée, Ivory Coast, Kenya, Liberia, Madagascar, malawi, Nigeria, Senegal, Tanzania, Togo, Zambia and Zimbabwe) but of course the exploration rate per area is very variable, with some areas which hardly have been prospected. This is illustrated by the fact that about 25% of the actually known species are only known from the type locality. We realize that the real number of Lactarius species is higher and that more new species certainly will be discovered. An overview of the collections sites is given in figure 16.
Though conclusions about the species distribution can only be provisional, we notice that most of the species have been collected either exclusively in woodlands (miombo, Uapaca woodland, Sudanian woodland, …) or in more humid forest types (rainforest, riparian forest, gallery forest, swamp forest, …) suggesting a high ecological selection. Other species occur in various forest types and seem widespread in tropical Africa: L. annulatoangustifolius (Fig. 17), L. chamaeleontinus, L. rubroviolascens.
Some species (L. heimii (Fig. 18), L. brunnescens, L. chromospermus, L. urens) are restricted to the Zambezian region (as defined by White, 1983) and others occur in both the Zambezian and Sudanan region (L. longisporus, L. emergens, L. luteopus (Fig. 19)), though the latter is poorer in ectotrophs and lacks important mycorrhizal tree genera as Brachystegia and Julbernardia.
Both the Guineo Congolian region (L. gymnocarpus (Fig. 20), L. russuliformis) and Madagascar (L. madagascariensis (Fig. 21), L. phlebophyllus) might be considered as centre of endemic ectotrophs, but they also have species in common (L. annulatoangustifolius, L. pelliculatus (Fig. 22)).
The existence of e.g. Uapaca gallery forests in the woodland regions makes it possible for Guineo-congolian ectomycorrhizal fungi to invade these regions (L. melanogalus (Fig. 23), L. acutus (Fig. 24)).
Abbreviations
The following abbreviations are used: Q = length/wide ratio of the spores, n = number of spores measured, L = number of entire lamellae, l = number of lamellulae. In the illustrations H stands for the elements of hymenium, H mc for marginal cells, H(p) for hymenium (profile view), H plcy for pleurocystidia, H plpscy for pleuropseudocystidia, P pe for the elements of pileipellis, P pe (p) for pileipellis (profile view), P pe (s) for pileipellis (surface view), S pe for the elements of stipitipellis and sp for basidiospores. The scale bar always represents 10 µm.
Numbering of the figures in the plates refers to the numbering of the taxa in the text.
Herbaria acronyms follow Holmgren & Holmgren (1998), colour codes are according to Kornerup & Wanscher (1978), authors of fungal names are abbreviated according to Kirk & Ansell (1992).
Lactarius Pers.
Lactarius Pers., Tent. Disp. Meth. Fung.: 63 (1797), as “Lactaria,” conserved orthography (Taxon 37: 456, 1988). – Lectotype: Lactarius piperatus (L.: Fr.) Pers. (Earle 1909: 409).
Agaricus section Lactifluus Pers., Syn. meth. fung.: 429 (1801); Lactifluus (Pers.) Roussel, Fl. Calvados, 2ème éd.: 66 (1806), lectotype: Lactifluus piperatus (L.: Fr.) Kuntze (Earle 1909: 409); Agaricus tribus Galorrheus Fr.: Fr., Syst. mycol. 1: 61 (1821); Galorrheus (Fr.: Fr.) Fr., Stirpes Agri Femsion. 3: 56 (1825), lectotype: Galorrheus piperatus (L.: Fr.) Fr. (Donk 1949); Lactariella Schröter in Cohn, Kryptog.-Fl. Schlesien, Pilze 3(1): 544 (1889), lectotype: Lactariella lignyota (Fr.) Schröter (Singer & Smith 1946); Arcangeliella Cavara, Nuovo Giorn. Bot. Ital., ser. 2, 7(2): 126 (1900), type: Arcangeliella borziana Cavara; Lactariopsis Henn., Bot. Jahrb. Syst. 30: 51 “1901” (1902), type: Lactariopsis zenkeri Henn.; Gloeocybe Earle, Bull. New York Bot. Gard. 5: 409 (1909), type: Gloeocybe insulsa (Fr.: Fr.) Earle; Pleurogala Redhead & Norvell, Mycotaxon 48: 377 (1993), type: Pleurogala panuoides (Singer) Redhead & Norvell; Zelleromyces Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 18 (1960), type: Zelleromyces cinnabarinus Singer & A.H. Sm.; Gastrolactarius R. Heim ex J.M. Vidal, Rev. Catalana Micol. 46: 76 ‘2004’ (2005), type: Gastrolactarius densus (R. Heim) J.M. Vidal.
brief history of Lactarius collections from Tropical Africa
Before our studies the knowledge of the genus Lactarius Pers. in tropical Africa was almost completely based on the monographs of Heim for Madagascar (1938b) and Congo and former French colonies in Western Africa (Heim 1955a, 1955b). Before Heim some other authors described several species receiving little attention until recently (Verbeken 1996a). Eichelbaum (1906) and Walker (1931) mentioned some vernacular names of Lactarius-like fungi without giving full descriptions, whereas Bresadola (in De Wildeman 1914) described a new species, Lactarius velutinus, which turned out to be a Russula (Buyck 1988). Beeli (1928) also described two Lactarius species and Hennings (1902) created Lactariopsis in which he described the annulate Lactariopsis zenkeri, later placed in Lactarius by Singer (1942). The presence in the tropics of veiled Lactarius species, species fructifying on rotten wood and species with a very restricted presence of milk, together with his lack of field experience in the tropics, apparently confused Beeli (1927a, 1927b, 1928, 1933, 1936) as he described Lactarius specimens in the genera Armillaria (1), Omphalia (1), Clitocybe (1), Russula (2) and Laccaria (1) (Verbeken 1996a).
More recently, scattered descriptions of African Lactarius species, including some new species can be found in Berthet & Boidin (1966), Heim (1966, 1967), Morris (1984, 1987, 1990), Pegler (1969, 1977, 1982) and Pegler & Piearce (1980). Our studies of tropical African milkcaps started in 1993, initially based on the collections of BR (mainly collected by M. Goossens-Fontana, M.C. Schmitz-Levecq, J. Rammeloo, D. Thoen, B. Buyck, J. Schreurs), Bart Buyck (from Zambia and Burundi, now at PC), Härkönen and coll. (Tanzania, H), and personal collecting in Burundi (preserved at BR). More recently, collections have been studied of André de Kesel (Benin, preserved at BR), Bart Buyck (Madagascar, preserved at PC), Roy Watling and Peter Roberts (Cameroon, E & K), Jérôme Degreef (Gabon, BR), A. Verbeken & F. Noe (Malawi, GENT), A. Verbeken & C. Sharp (Zimbabwe, GENT). African milkcaps received also attention in several modern studies on edible fungi from Burundi (Buyck 1994), Tanzania (Härkönen & al. 1995, 2003) and Benin (De Kesel & al. 2002, Yorou & De Kesel 2002, Yorou & al. 2002). The number of accepted species has raised from 35 in 1989 to 96 in 2008 (Buyck & Verbeken 1995; Buyck & al. 2007; Eberhardt & Verbeken (2004); Karhula & al. 1998; Van Rooij & al. 2003; Verbeken 1995a, 1995b, 1996a, 1996b, 1996c, 1996d, 1998a, 1998b, 2001a; Verbeken & Sharp 2003; Verbeken & Walleyn 2000; Verbeken & al. 2000, 2008). The real number of tropical African Lactarius species is estimated to be at least 125 (Verbeken & Buyck 2001).
PRACTICAL key to sections of the genus Lactarius in tropical Africa
1. Carpophores sequestrate (gasteroid or secotioid)............................. sect. Plinthogali s.l.
Carpophores with normally developed stipe and lamellae .............................. 2
2. Species associated with exotic Pinus trees; cap reddish brown........... sect. Russularia
Species associated with indigenous trees (mainly Caesalpinioideae, Uapaca, Papilionoideae, Dipterocarpaceae) ..... 3
3. Stipe with a distinct, more or less fugaceous annulus ....................... sect. Lactariopsis
Stipe without an annulus; velar remnants only present at pileus margin or absent................... 4
4. Pileus margin with velar remnants .................................................. sect. Chamaeleontini
Velar remnants absent ................................................................. 5
5. Context blackening ........................................................................ 6
Context not blackening (browning, unchanging etc.) ............................. 7
6. Hymenium with lamprocystida; spores ornamented with low warts or ridges ........ sect. Rubroviolascentini
Hymenium without lamprocystidia; spores with winged ornamentation, at least 1 µm high .................. sect. Nigrescentes
7. Spore print and mature gills chocolate brown; pileus viscid ...................... sect. Chromospermi
Spore print white to yellowish or pale pinkish, at most pale brown; pileus dry or viscid ........... 8
8. Pileus viscid with strongly hairy margin OR stipe scrobiculate ......... sect. Piperites s.l.
Pileus viscid or dry, cap margin not hairy, stipe never scrobiculate .............................................. 9
9. Pileus pruinose, orange and concentrically zonate when moist; lamellar edge fimbriate, appendiculate, with transparent dropplets when fresh; stipe tapering downwards and clearly chambered .................................. sect. Aurantiifolii
Not with this combination of characters ...................................... 10
10. Hymenium with conspicuous lamprocystidia ........................ sect. Pseudogymnocarpi
Hymenium without lamprocystidia ................................................ 11
11. Pileipellis a palisade with an upper layer of anticline, thick-walled elements (lampropalisade); context (e.g. gills) often browning when bruised ..................................................................... 12
Pileipellis structure different, in case of a palisade without thick-walled elements .................. 13
12. Spore ornamentation typically composed of almost isolated, distinct rounded warts and some very fine connective lines between them ....................................... sect. Phlebonemi
Spore ornamentation more or less reticulate ............................. sect. Polysphaerophori
13. Spore ornamentation zebroid ..................................................................... sect. Plinthogali
Spore ornamentation different ................................................... 14
14. Pileus dull coloured (brown, greyish, coffee with milk, buff), without orange or reddish tinges; spores often winged or reticulate, sometimes (sub)globose ................................................. 15
Pileus differently coloured, often more vivid, with orange or reddish tinges ............................ 17
15. Spores small, 6–7 × 4–5.5 µm, ellipsoid to elongate; pileipellis a trichoderm ......... L. badius
Spores larger or subglobose; pileipellis a palisade or trichopalisade ......................................... 16
16. Pileipellis a palisade or hymeniderm, dry, often velvety or finely tomentose to slightly squamulose; context sometimes changing pinkish when bruised ............. sect. Plinthogali
Pileipellis a trichoderm, smooth, sometimes shiny or subviscid; context not changing pinkish when bruised ........ sect. Pseudofuliginosi
17. Carpophores rather small; pileus ochraceous, concentrically zonate when fresh; stipe fistulous; pileipellis a cutis to trichoderm, composed of long interwoven hyphae ........... sect. Amari
Not with this combination of characters .................................. 18
18. Pileipellis a palisade, with an upper layer of anticline elements; basidia long and slender; spores ellipsoid to elongate, verrucose or more or less reticulate, plage often with a central amyloid spot; pileus dry, orange, yellowish, reddish, reddish brown (sub)globose .......... sect. Rugati
Pileipellis structure different .......................... 19
19. Pileipellis an ixocutis (to locally ixotrichoderm) and cream-coloured...... sect. Piperites
Pileipellis dry or viscid but not an ixocutis, if an ixotrichoderm then pileus orange ................ 20
20. Pileipellis most often with scattered thick-walled hairs; pileus shiny and smooth, dry to viscid, margin often striate or sulcate; pleuropseudocystidia emergent to very emergent, mostly broad and fusiform .................... sect. Chamaeleontini
Pileipellis dry, lacking thick-walled hairs ......................................................................................... 21
21. Pileus < 35 mm diam., ochraceous red, brownish red, tomentose to finely scaly ........ L. hispidulus
Not with this combination of characters ........................................................................................... 22
22. Basidiomes reminding of a small, reddish or pinkish Russula; spore ornamentation composed of slender spines up to 1.5–2 µm high, regularly connected by fine connective lines ..... L. russuliformis
Not with this combination of characters ................................... 23
23. Pileus vividly ochraceous orange; lamellae dense; spores with rounded, mostly isolated warts; pleuromacrocystidia abundant, slightly emergent and with dense needle-like content .............................. L. caperatus
Not with this combination of characters ................................................................. 24
24. Pileipellis dry, a trichoderm or composed of chains of subcylindrical to subsphaerical cells (cap often cracked); pileomacrocystidia absent ............................................... sect. Edules
Pileus dry to slightly viscid; pileipellis cutis-like with distinct macrocystidia; pileus often with reddish colours; pseudocystidia often branched and diverticulate ........ sect. Russulopsidei
Sect. Lactariopsis (Henn.) Singer
Lactarius sect. Lactariopsis (Henn.) Singer, Ann. Mycol. 40: 111 (1942) (as Lactariopsideae).
Lactariopsis Henn., Bot. Jahrb. Syst. 30: 51 “1901” (1902).
sect. Lactariopsidei “Singer ex Singer”, Sydowia 15: 83 (1962, illegit.).
− Type species: Lactarius zenkeri Henn.
Key to tropical African species
1. Thick-walled hair-shaped elements in the pileipellis scarce, pleurocystidia present ... sect. Chamaeleontini, 6. L. indusiatus
Pileipellis entirely composed of thick-walled and hair-shaped elements, pleurocystidia absent ........... 2
2. Spore ornamentation well developed, up to 1 mm high ..................... 1. L. annulatoangustifolius
Spore ornamentation faint, mostly up to 0.2 mm high ........................................................................ 3
3. Spores ellipsoid to elongate, Q = (1.15)1.3–1.5(1.65), spores up to 11.5 mm long; spore-ornamentation an almost complete reticulum ................ 4. L. heimii
Spores ellipsoid, sometimes subglobose, Q = (1.09)1.19–1.32(1.45), spores usually shorter than 9(10) mm; spore-ornamentation an incomplete reticulum, always with distinct warts ............. 4
4. Pileipellis remarkably thick, felty, yellowish white, persistent, stellately rupturing; spores 8.4–9.2 mm long; only known from Zambezian miombo woodland .......................... 5. L. velutissimus
Pileipellis thinner, submembranaceous .......................................... 5
5. Basidiomes fragile, yellowish brown to pale ochraceous; spores small, 7.1–7.8 ´ 5.5–6.3 mm, with extremely low ornamentation; marginal cells tortuous and branched ................. 2. L. zenkeri
Basidiomes firmer, pale yellow ochre to pale yellowish; spores larger, 8.0–8.6 ´ 6.4–7.0 mm, ornamentation 0.2 mm high; marginal cells narrowly utriform to conical ...... 3. L. pelliculatus
Species descriptions
1. Lactarius annulatoangustifolius (Beeli) Buyck
Buyck, Bull. Jard. Bot. Nat. Belg. 59: 241 (1989). – Russula annulatoangustifolia Beeli, Bull. Jard. Bot. Etat Bxl 14: 87 (1936). – Type: Democratic Republic of Congo, Binga, 8 Nov. 1928, M. Goossens‑Fontana 834 (BR 4976–29, holotype).
Lactarius pandani R. Heim, Candollea 7: 376 (1938). – Type: Madagascar, Betsimisaraka Prov., South of Soanierana Gagnoa, 9 Dec. 1934, R. Heim G89bis (PC, holotype).
Lactarius pandani f. aurantiacus R. Heim (invalid), Bull. Jard. Bot. Etat Bxl 25: 18 (1955).
Misappl.: Lactarius zenkeri ss. Singer (1946).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 37-43 & Pl. 1a (1938); Heim, Bull. Jard. Bot. Etat Bxl 25: 17–20 & Pl. 1, fig. 1a–e (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 13, fig. 4 (1955); Verbeken, Edinburgh J. Bot. 53: 72–77 (1996a); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 143–145 & Pl. 69–76 (1996d); Karhula & al., Karstenia 38: fig. 12 (1998).
Pileus (10–)30–50(–75) mm diam., fleshy and fairly thick when young, fragile and thin when older, convex with papilla, then applanate and slightly depressed; margin thin, incurved, then straight; pellis thick, gelatinized but not glutinous, with characteristic radial to sinuous veins when young, yellow to brownish orange, darker in the centre, paler yellow-brown with faint pink tinge ('tan') at the margin, rupturing and leaving whitish fine flocks, showing an underlying striate layer, dehiscent at the margin. Secondary velum closed in the young basidiomes (development velangiocarpic), then forming a submembranaceous annulus, whitish to cream, not mobile, ring-shaped. Lamellae adnate to decurrent, prolonging into veins on the stipe, unequal with lamellulae of different lengths, narrow (1–3 mm), crowded, yellow to cream, then darker with pink tinge, more dirty when bruised; edge entire, concolorous. Stipe (25–)35–50 × (3–)6–12 mm, cylindrical, tapering downwards, slightly sinuous or curved, wrinkled when young, then smooth, with gelatinized veins in the upper part, spongy solid, then fistulous, subconcolorous with the pileus, paler cream at the apex, yellowish at the base; with whitish and abundant mycelium at the base. Context white to cream, first solid then granulose and fragile in pileus, fibrillose in stipe; taste (very) acrid; smell disagreeable, acrid. Latex abundant when young, then scarce, white; taste acrid. Spore-deposit white. – Pl. 1
Chemical reactions. Context after some minutes blue-green with gaiac, unchanging with gaiacol and pyramidon, immediately dark brown with pyrogallol, immediately purple with phenol, unchanging with Fe2SO4, &-naphtol, NH3 and anniline, reddish with sulfovanilline and brown with sulfoformol (according to Heim 1955a).
Basidiospores ellipsoid, 7.6–8.7–9.5–10.1(–10.7) × 6.3–7.0–7.6–8.1(–8.7) mm (Q = 1.14–1.24–1.28–1.42; n = 100); ornamentation amyloid, composed of conical to irregular warts and short ridges, < 1 mm high, often connected by fine and lower lines, forming an incomplete reticulum; surface rugose between the ornamentation; plage inamyloid. Basidia (30–)35–50 × 9–13 mm, cylindrical to narrowly clavate, 4-spored, rarely 2-spored. Pleurocystidia absent. Pleuropseudocystidia rather abundant, 12–20 mm diam., emergent up to 50 mm, clavate to tortuous, tapering upwards, capitate, mucronate to rostrate; wall slightly thickened; content needle-like to granular. Lamellar edge sterile; marginal cells 12–30 × 5–10 mm, irregularly fusiform to utriform, often septate, thin-walled. Hymenophoral trama composed of sphaerocytes in the upper part, subregular and composed of narrow hyaline hyphae towards the edge of lamellae; laticiferous hyphae present, narrow. Pileipellis a lampropalisade to lamprotrichopalisade; elements of suprapellis 30–110 × 4–8 mm, long, slender, hair-shaped and tapering upwards, very thick-walled (1–2 mm), sometimes septate and branched; subpellis composed of narrow cells, 5–20 mm, slightly thick-walled to thin-walled, with some elements of the upper layer penetrating the subpellis. Stipitipellis a lampropalisade to lamprotrichopalisade, as pileipellis, although at the base of the stipe consisting of narrower and longer thick-walled, unbranched hyphae. Annulus composed of thin-walled, hyaline hyphae, which are densely interwoven, often septate, sometimes branched, 2–5 mm diam. Clamp-connections absent. – Fig. 25
Ecology. Found terrestrial or on rotten wood but connected to white mycelium and mycorrhizal roots) in drier Guineo‑Congolian rainforest with Gilbertiodendron dewevrei, Malagasy lowland rainforest, dry evergreen forest (muhulu) and wetter Zambezian miombo woodland. In Madagascar found under Uapaca louvelii and U. densifolia.
Distribution. Widespread in tropical Africa. Known from Cameroon, Democratic Republic of Congo, Gabon, Guinea, Liberia, Madagascar, Zambia and Zimbabwe.
Revised specimens
Cameroon. South Western Prov., near Mundema, Korup National Park, transect P, Jan. 1989, R. Watling 21466 (E); Ibid., Jan. 1989, R. Watling 21472 (E); Ibid., March 1989, R. Watling 24187 (E); Ibid., Jan. 1988, I. Alexander 16b (E); Ibid., trail from Mana Bridge to transect P, 12 April 1997, P. Roberts K1136 (K(M) 50411); environment of Douala, 12 Aug. 1959, Berthet 314 (LY, cited as L. pandani f. aurantiacus in Berthet & Boidin 1966).
Democratic Republic of Congo. Equateur Prov., Binga, Nov. 1928, M. Goossens-Fontana 834 (BR 4976–29, holotype); Ibid., Aug. 1929, M. Goossens-Fontana 877 (BR 4969–22); Ibid., Sept. 1940, M. Goossens-Fontana 2038 (BR 12318–96); Ibid., Oct. 1945, M. Goossens-Fontana 3061 (BR 4941–91); Tshopo Prov., 5 km NNE of Batiabongena, in humus, April 1984, B. Buyck 1344 (BR 6088–74); Ibid., May 1984, B. Buyck 1656 (BR 6094–80); Yanero, May 1984, B. Buyck 1730 (BR 6097–83); Shaba Prov., Muhulu de la Luiswishi, Dec. 1971, D. Thoen 5137 (BR 66283–32).
Gabon. Research Station of Ipassa-Makokou, 8 March 2005, B. Toirambe 1 (BR 164547–35); Ibid., 13 March 2005, S. Dibaluka Mpulusu 17 (BR 164552–40).
Guinea. Forêt humide de Ziama, arboretum des stagiaires, 12 April 2000, A. Bâ 360 (GENT).
Liberia. Near Nengbe, April 1939, G.W. Harley 57 (FH).
Madagascar. Betsimisaraka Prov., S of Soanierana Gagnoa, Dec. 1934, R. Heim G89 (PC); Ibid., R. Heim G89bis (PC, type L. pandani); Ibid.?, R. Heim s.n. (PC); Fianarantsoa, Ranomafana National Park, 1 Feb. 2000, B. Buyck & al. 00-1031, 00-1111, 00-1112 & 00-1153 (all PC); Ambotantihely, N of Ankazobe, sentier botanique, 15 Feb. 2000, B. Buyck & al. 00-1518 (PC); Toamasina Prov., station forestière Andasibe-Analamazaotra, 23 Jan. 1999, B. Buyck & G. Eyssartier 99-324 & 99-332 (all PC).
Zambia. Western Prov., Mwinilunga, road to Solwezi km 50, Jan. 1991, B. Buyck 3516, 3567-3569 & 3595 (all PC).
Zimbabwe. Manicaland Prov., Eastern Highlands, Mguzu, leg. Gansemans 5 Feb. 1999, A. De Kesel 2739 (BR 115761–40).
Notes. The macroscopic description is based on Heim (1938b, 1955), Singer (1946) and fieldnotes of Thoen. The microscopic description is based on Goossens-Fontana 2038, Heim G89 and Thoen 5137. The water-colours of Goossens-Fontana 834, 877 & 3061 show conspicuous (dried out?) whitish basidiomes although no microscopic differences could be found. Moreover, the exsiccata of these collections have the same colour as the other studied specimens. As a consequence we consider them conspecific. New field observations are necessary to find out whether this concerns different varieties or forms.
The species is easy to recognize, among the other annulate species, by its more prominent spore-ornamentation; the warts are up to 1 mm high, whereas the other species in the section have an ornamentation which never exceeds 0.5 mm.
2. Lactarius zenkeri (Henn.) Singer
Singer, Ann. Mycol. 40: 111 (1942). − Lactariopsis zenkeri Henn., Bot. Jahrb. Syst. 30: 51 “1901” (1902). − Type: Cameroon, Korup National Park, Jan. 1988, I. Alexander 16 (E, neotype).
Excl.: Lactarius zenkeri ss. Singer (1946), = L. annulatoangustifolius.
Selected descriptions and icons. Verbeken, Edinburgh J. Bot. 53: 50–54 (1996a); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 145–146 & Pl. 77–81 (1996d).
Pileus 30–60 mm diam., first convex, then applanate and infundibuliform; margin thin, incurved, then straight; pellis submembranaceous, pruinose, whitish, yellowish brown to pale ochraceous, darker in the centre, paler towards the margin, rupturing towards the margin and leaving small whitish flocks, underlaying layer smooth and striate. Secondary velum closed in the young basidiomes (development velangiocarpic), then forming an annulus near the apex of the stipe, membranaceous, thin, whitish. Lamellae adnate to decurrent, crowded, narrow to medium broad (1–3 mm), waxy, ochraceous, paler than pileus; edge entire, concolorous. Stipe 20–40 × 5–20 mm, cylindrical to subclavate, ochraceous, dry, smooth, paler at the apex, smooth, solid, sometimes becoming fistulous. Context firm, white, not changing; smell not remarkable. Latex quite abundant, white; taste unknown. Spore-deposit white.
Basidiospores ellipsoid, sometimes subglobose, 6.4–7.1–7.8–8.6 × 4.9–5.5–6.3–6.9 mm (Q = 1.09–1.19–1.29–1.45; n = 100); ornamentation amyloid, low, composed of irregular sized and shaped warts; warts less then 0.2 mm high, isolated, connected by lines or aligned, sometimes forming a faint, partial reticulum; plage mostly not amyloid, sometimes with a centrally, slightly amyloid spot. Basidia 30–40(–50) × 9–10 mm, cylindrical to slightly clavate, 4-spored, rarely 2-spored. Pleurocystidia absent. Pleuropseudocystidia rather scarce, 15–25 mm diam. in the upper part, clavate to conical, tapering upwards, mucronate, very emergent, with needle-like to granular content. Lamellar edge sterile; marginal cells 15–40 × 5–10 mm, narrowly utriform to conical, in some specimens tortuous and often dichotomously branched. Hymenophoral trama composed of sphaerocytes and numerous laticiferous hyphae; narrow, hyaline hyphae becoming more abundant near the edge. Pileipellis a lamprotrichopalisade to lampropalisade; elements of suprapellis very thick-walled (1–2 mm), 50–110 × 4–8 mm, long, slender, hair-shaped and tapering upwards, septate and sometimes branched; subpellis composed of irregular to isodiametric cells, 5–15(–20) mm, with some elements of the upper layer penetrating. Stipitipellis a lamprotrichopalisade; elements of suprapellis 50–120 × 7–10 mm, very often branched, thick-walled (2–3 mm); with numerous lactifers. Annulus composed of narrow hyphae which are thin-walled, hyaline, septate and sometimes branched, (2–)3–5(–7) mm diam. Clamp-connections absent. – Fig. 26
Ecology. Lowland rainforest, semi‑evergreen forest with Afzelia, Uapaca bojeri woodland.
Distribution. Known from Cameroon, Gabon, Madagascar and Senegal.
Revised specimens
Cameroon. South Western Prov., near Mundema, Korup National Park, transect P16-27, Jan. 1988, I. Alexander 16 (E, neotype); Ibid., transect P23, Jan. 1989, R. Watling 21471 (E); Ibid., P23, R. Watling 21480 (E); Ibid., P18, May 1989, R. Watling 22185 (E); Ibid., trail to transect P, R. Watling 22186 (E); Ibid., P18, March 1991, R. Watling 23135 (E); Ibid., P15, R. Watling 23136 (E); Ibid., transect before P, R. Watling 23137 (E); Ibid., P13, R. Watling 23139 (E); Ibid., P21, R. Watling 25312 (E).
Gabon. Research Station of Ipassa-Makokou, 15 March 2005, A. Murhula Cizungu 20 (BR 164546–34).
Madagascar. Arivonimamo, 30 Jan. 1997, B. Buyck & al. 97-092 (PC); Ibid., 5 Feb. 1997, B. Buyck & al. 97-286 (PC).
Senegal. Basse Casamance Prov., 10 m alt., Sept. 1987, legit A. Bâ, D. Thoen 7885 (BR 66456–11).
Notes. The original type material (Zenker 2230, with water colour) is lost at Berlin (Horak 1968) and no isotype-material could be traced at BR, PC, K(M), E, FH or S. Singer (Singer & al., 1983) refers to "the type" of Lactarius zenkeri designating a collection from Liberia which he described as L. zenkeri, although this exsiccatum clearly represents L. annulatoangustifolius (q.v.), a species which differs particularly in having more bright orange colours and larger spores with a more pronounced ornamentation. To avoid further misunderstanding we proposed the collection Alexander 16 (E) as the neotype for Lactarius zenkeri (Verbeken 1996a). This collection was made in Korup National Park (Cameroon) where several other fungi occur which were described by Hennings from Cameroon (Watling, pers. comm.).
The macroscopic description is based on the description of Hennings (1902) and the fieldnotes from Watling. The microscopic description is based on Alexander 16, Watling 23135 and Thoen 7885.
Lactarius zenkeri is recognizable as a small and tiny annulate species with a very fragile context and small, lowly ornamented spores.
3. Lactarius pelliculatus (Beeli) Buyck
Buyck, Bull. Jard. Bot. Nat. Belg. 59: 242 (1989). – Armillaria pelliculata Beeli, Bull. Soc. Roy. Bot. Belg. 59: 111 (1927). – Type: Democratic Republic of Congo, Lisala, Dec. 1925, M. Goossens-Fontana 546 (BR 7715–52, holotype).
Lactarius pandani R. Heim f. intermedius R. Heim, Bull. Jard. Bot. Etat Bxl 25: 20 (1955, as "var. intermedius" in Fl. Icon. Champ. Congo 4: 87 (1955), invalid.
Lactarius pandani f. pallidus R. Heim, Bull. Jard. Bot. Etat Bxl 25: 17 (1955), invalid.
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 17–23 & Pl. 1, figs. 2–4 (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 13, fig. 2a–b & 3 (1955); Verbeken, Edinburgh J. Bot. 53: 56–60 (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 146–147 & Pl. 82–88 (1996d).
Pileus 40–80 mm diam., fleshy, firm, convex, then applanate, infundibuliform when older; margin thin, striate, first incurved, then straight; pellis viscous and with radial, gelatinized veins when wet, greyish and pellicular when dry, slightly gelatinized, ferruginous red, yellowish ochre to pale yellowish, leaving fine, whitish flocks towards the margin, dehiscent near the margin. Secondary velum closed in the young basidiomes (velangiocarp development), then forming an annulus near the apex of the stipe, membranaceous, thin, whitish. Lamellae adnate to decurrent, rather broad, rather thick, brittle, with some anastomosing veins near the margin, sometimes dichotomously branched, pale ochraceous. Stipe 30–50 × 9–14 mm, cylindrical, short, tapering downwards, smooth, with some longitudinal veins near the apex, ochraceous, paler than the pileus, fleshy, solid, then fistulous. Context white to pale yellowish; taste acrid; smell pepper-like. Latex scarce, transparent, watery, not present in older basidiomes; taste acrid to very acrid. Spore-deposit white. (According to Heim 1955a). – Pl. 2
Basidiospores ellipsoid, 7.1–8.0–8.6–9.1(–9.9) × 5.8–6.4–7.0–7.5(–8.3) mm (Q = 1.13–1.23–1.24–1.35, n = 60); ornamentation amyloid, composed of irregularly sized and shaped warts; warts less then 0.5 mm high, isolated, connected by lines or aligned, sometimes forming a faint, partial reticulum, warts always distinct; plage inamyloid. Basidia 40–50 × 9–10 mm, cylindrical to slightly clavate, 4-spored, rarely 2-spored. Pleurocystidia absent. Pleuropseudocystidia rather scarce, 15–25 mm diam. in the upper part, clavate to conical, tapering upwards, mucronate, very emergent, with needle-like to granular content. Lamellar edge sterile; marginal cells 15–40 × 5–10 mm, narrowly utriform to conical. Hymenophoral trama composed of sphaerocytes and numerous laticiferous hyphae; narrow, hyaline hyphae becoming more abundant near the edge. Pileipellis a lamprotrichopalisade; elements of suprapellis very thick-walled (1–2 mm), 50–130 × 4–8 mm, long, slender, hair-shaped and tapering upwards, septate and sometimes branched; subpellis composed of irregular to isodiametric cells, 5–15(–20) mm, with some elements of the upper layer penetrating. Stipitipellis a lamprotrichopalisade to lampropalisade, with larger isodiametric cells than the pileipellis; elements of suprapellis 50–150 × 7–10 mm, sometimes branched, with 2–3 mm thick walls; with numerous lactifers. Annulus composed of narrow hyphae which are thin-walled, hyaline, septate and sometimes branched, (2–)3–5(–7) mm diam. Clamp-connections absent. – Fig. 27
Ecology. Guineo‑Congolian rainforest, Uapaca woodland.
Distribution. Known from Democratic Republic of Congo, Gabon, Guinea and Madagascar.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Lisala, 350-400 m alt., on the ground, Dec. 1925, M. Goossens-Fontana 546 (BR 7715–52, holotype); Binga, 380 m alt., Aug. 1928, M. Goossens-Fontana 807 (BR 12321–02); Ibid., Dec. 1941, M. Goossens-Fontana 2069 (BR 12320–01); Ibid., Oct. 1942, M. Goossens-Fontana 3036 (BR 12319–00).
Gabon. Research Station of Ipassa-Makokou, 16 March 2005, S. Dibaluka Mpulusu 28 (BR 164539–27); Ibid., 16 March 2005, A. Murhula Cizungu 21 (BR 164546–34).
Guinea. Forêt humide de Ziama, arboretum des stagiaires, July 1999, A. Bâ 62 (GENT).
Madagascar. Toamasina, RN7, 32 km after Antsirabe, Fiadanana village, 10 Feb. 2000, B. Buyck & al. 00-1335 (PC); Ambotantihely, N of Ankazobe, sentier botanique, 15 Feb. 2000, B. Buyck & al. 00-1389a (PC).
Notes. Lactarius pelliculatus is very closely related to L. zenkeri, but differs by the somewhat larger basidiomes (pileus up to 8 cm diam.) and the slightly larger spores. The marginal cells are not tortuous or branched, as is often the case in Lactarius zenkeri. The ornamentation of the spores is more pronounced than in Lactarius zenkeri, although still very low and faint when compared to L. annulatoangustifolius.
Buyck (1989a) synonymized Lactarius pandani R. Heim with L. pelliculatus (Beeli) Buyck, without any comment. It has been demonstrated that only part of the specimens described as Lactarius pandani (“f. pallidus” and “f. intermedius”) is conspecific with L. pelliculatus, while the type of L. pandani and a specimen described as L. pandani f. aurantiacus represent L. annulatoangustifolius (Verbeken 1996a).
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 201 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 30 March 1994, A. Verbeken 94–465 (BR 57671–53, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 147–149 & Pl. 89–92 (1996d); Karhula & al., Karstenia 38: fig. 13 (1998); Härkonen & al., Norrlinia 10: 87 (2003).
Pileus 30–90 mm diam., firm and fleshy, convex to planoconvex, depressed in the centre; margin strongly involute when young, then deflexed downwards; pellis very thin, arachnoid, pruinose, greyish orange to brownish orange (5AB3–8, 6BC3–5), rather pale yellow (4A4–5) near the margin, orange white to light yellow or dark orange when young (4A4–5, 6A2), rupturing at the margin, showing a more shiny underlaying layer. Secondary velum forming an annulus near the apex of the stipe, arachnoid, thin, whitish. Lamellae adnate to slightly decurrent, unequal with lamellulae, moderately distant (L+l = 6+6 to 8+2/cm), medium broad (2–4 mm), thin, fragile, yellowish white, 3A2–3; edge concolorous, entire. Stipe 25–35 × 8–15 mm, cylindrical, slightly tapering downwards, felty, dry, orange white to cream (5A2), firm and solid. Context firm, (1)2–3 mm in the pileus, white to cream, unchanging; taste spicy to very acrid; smell agreeable, sweet. Latex not abundant, white or watery, taste mild or bitter to more or less acrid. Spore-deposit white. – Pl. 3
Chemical reactions. Context greenish (Buyck 3200) or unchanging (Verbeken 94-465) with FeSO4; blue-green with gaiac (Buyck 3200).
Basidiospores ellipsoid to elongate, 7.6–8.7–10.7–11.6 × 5.9–6.7–7.4–8.3 mm (Q = 1.16–1.30–1.49–1.64; n = 240); ornamentation amyloid, very dense, composed of low warts and ridges, < 0.2 mm high, aligned and connected by fine lines, forming an incomplete to almost complete reticulum with very small meshes; plage inamyloid. Basidia 45–55 × 10–12 mm, subclavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia scarce, 10–18 mm diam., emergent up to 50 mm, cylindrical, obtuse and with rounded apex; content densely granular; some pseudocystidia connected with oleiferic hyphae, 5–7 mm diam., content oleiferic. Lamellar edge sterile; marginal cells 15–25 × 7–11 mm, shortly cylindrical to clavate, thin-walled, sometimes septate. Hymenophoral trama composed of sphaerocytes and hyaline hyphae; lactifers abundant and up to 15 mm diam. Pileipellis a lamprotrichopalisade; elements of suprapellis (10–)30–200 × 5–10 mm, short and subclavate or long, slender, cylindrical and hair-shaped, often septate and branched, very thick-walled, 2–3(–4) mm; subpellis composed of irregular to isodiametric elements, very poorly developed with a lot of penetrating elements from suprapellis who are connected with hyaline, thin-walled hyphae. Stipitipellis a lamprotrichopalisade, as pileipellis. Annulus composed of interwoven, hyaline, thin-walled hyphae, 4–6(–9) mm diam., septate and sometimes branched. Clamp-connections absent. – Fig. 28
Ecology. Zambezian miombo woodland. Collected under various trees such as Brachystegia, Baphia, Parinari, Isoberlinia, Uapaca, Albizia, Julbernardia, Marquesia.
Distribution. Known from Burundi, Malawi, Tanzania, Zambia and Zimbabwe.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, Dec. 1993, B. Buyck 5358; Ibid., March 1994, A. Verbeken 94-003 (BR 57555–34), 94-115 (BR 57598–77), 94-158 (BR 57612–91), 94-209 (BR 57619–01), 94-210 (BR 57620–02), 94-440 (BR 57666–48) & 94-465 (BR 57671–53, holotype; GENT, isotype); Ibid., April 1994, A. Verbeken 94-560 (BR 57703–85); Nkayamba, just N of Rumonge, Dec. 1991, B. Buyck 4050 & 4051 (all PC); Ibid., Jan. 1992, B. Buyck 4187 (PC); Ibid., March 1992, B. Buyck 4239 & 4285 (all PC); Ibid., Dec. 1992, B. Buyck 4652 (PC); Ibid., B. Buyck 4686 & 4724 (all PC); Ibid., Jan. 1993, B. Buyck 4849 (PC); Ibid., Feb. 1993, B. Buyck 4972 (PC); Mugara, Nov. 1978, J. Rammeloo 5868 (BR 8779–49) & 5908 (BR 8773–43); Ibid., Dec. 1978, J. Rammeloo 5997 (BR 18928–13), 6006 (BR 18927–12) & 6073 (BR 8777–47).
Malawi. Central Prov., Dedza distr., Chongoni, 11 Feb. 1972, Williamson 594 (K(M) 38476).
Tanzania. Western Prov., Kahama distr., Mpunze, Forest Reserve, 10 Dec. 1991, T. Saarimäki & al. 1034 & 1042 (all H); Ibid., Tabora distr., Lulanguru, 14 Dec. 1991, T. Saarimäki & al. 1111 (H); Southern Highlands Prov., Iringa distr., Itagutwa, 40 km NE of Iringa, 8 Feb. 1993, T. Saarimäki & al. 1600 (H).
Zambia. Copperbelt Prov., Chati‑forest, near Kitwe, Dec. 1990, B. Buyck 3122, 3127, 3139, 3159, 3162, 3189, 3191 & 3200 (all PC); Ibid., B. Buyck 3203 & 3209b (all PC); Luapala Prov., Mansa, Kwanfumwe, 2 Jan. 1991, B. Buyck 3316 (PC); Kabunda mission, 3 Jan. 1991, B. Buyck 3354 & 3355 (all PC); Western Prov., Mwinilunga, 29 Jan. 1991, B. Buyck 3515 (PC).
Zimbabwe. Midlands Prov., Mvuma, Central Estates, south side of Beacon Hill, 27 Jan. 1999, A. Verbeken 99-029 (GENT); Ibid., Baru section, 27 Jan. 1999, A. Verbeken 99-036 (GENT); Midlands Prov., Mvuma, Mtao forest, 28 Jan. 1999, A. Verbeken 99-044 (GENT); Ibid., 11 Feb 1999, A. De Kesel 2451 (BR 112621–04)
Notes. The macroscopic description is based on fieldnotes of Rammeloo, Verbeken & Buyck. The microscopic description is based on Verbeken 94-465, 94-210, 94-003 and Buyck 3189.
Among the other annulate species, Lactarius heimii is easily recognized by its elongate spores which are ornamented with a dense, almost complete reticulum with hardly any warts distinguishable.
This species is consumed by some local populations (Karhula & al. 1998, Verbeken & al. 2000).
5. Lactarius velutissimus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 212 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 23 March 1994, A. Verbeken 94-291 (BR 57639–21, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 149–150 & Pl. 93–96 (1996d); Karhula & al., Karstenia 38: fig. 14 (1998); Ryvarden & al., An introduction to the larger fungi of South Central Africa: 100 (1994, as Lactarius pandani); Verbeken, Micol. Veget. Medit. 16: fig. 6 (2001).
Pileus 45–140 mm diam., first convex, then applanate and planoconcave to infundibuliform; margin involute when young, straight and even deflexed upwards when older, then sometimes striate; pellis thick, felty, pale orange to light yellow in the centre (4A3–5A3), yellowish white (4A2) towards the margin, rupturing star-shaped and showing a shiny, striate, somewhat darker layer. Secondary velum forming an annulus near the apex of the stipe, membranaceous, arachnoid when older, not mobile, white to cream (4A2). Lamellae decurrent, unequal with a few lamellulae of different lengths, sometimes dichotomously branched, waxy to fragile, thin, pale yellow to light yellow (3A2–3 to 4A2–4); edge concolorous, entire. Stipe 25–35 × 6–17 mm, shortly cylindrical, tapering downwards, felty, pale orange, light orange to cream (4A3–5A3). Context firm, 4–6 mm in the pileus, solid in stipe, white, changing light orange in the base of the stipe; taste acrid; smell agreeable. Latex rather abundant, white, taste acrid. Spore-deposit white. – Pl. 4
Chemical reactions. Context pale orange with FeSO4.
Basidiospores ellipsoid, 7.6–8.4–9.2–10.4 × 6.3–6.8–7.2–8.1 mm (Q = 1.15–1.21–1.32–1.40; n = 100); ornamentation amyloid, composed of irregularly sized and shaped warts and ridges, sometimes aligned, sometimes connected by fine lines, never forming a reticulum, < 0.2 mm high, wall between the ornamentation rugose; plage inamyloid. Basidia 48–60 × 9–10 mm, subclavate, 4-spored; content guttate or needle-like. Pleurocystidia absent. Pleuropseudocystidia not abundant, 12–15 mm diam., emergent up to 30 mm, fusiform, tapering upwards, mucronate; content granular and needle-like. Lamellar edge sterile; marginal cells 15–25 × 8–12 mm, shortly cylindrical or clavate, sometimes septate. Hymenophoral trama composed of sphaerocytes and hyaline hyphae, lactifers abundant. Pileipellis a lamprotrichopalisade; elements of suprapellis (20–)50–200 × 5–6(–10) mm, short and clavate or long, slender and hair-shaped, often septate and branched, very thick-walled, 2–3(–4) mm; subpellis composed or irregular to isodiametric elements, sometimes slightly thick-walled. Stipitipellis a lamprotrichopalisade, as pileipellis; elements of suprapellis 60–200 × 4–6 mm. Annulus composed of narrow hyphae, thin-walled, hyaline, septate and sometimes branched. Clamp-connections absent. – Fig. 29
Ecology. Zambezian miombo woodland.
Distribution. Known from Burundi, Democratic Republic of Congo, Kenya, Malawi, Tanzania, Zambia and Zimbabwe.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, Dec. 1993, B. Buyck 5358b; Ibid., March 1994, A. Verbeken 94-007 (BR 57559–38), 94-075 (BR 57585–64), 94-291 (BR 57639–21, holotype; GENT, isotype) & 94-439 (BR 57665–47); Cankuzo Prov., Murango, March 1994, B. Nzigidahera 58 & 61 (all PC).
Democratic Republic of Congo. Shaba region, Kipopo, 1250 m alt., 5 Dec. 1971, D. Thoen 5084 (BR 66272–21); Luiswishi, terrestrial, Feb. 1986, J. Schreurs 1121 (BR 7954–00) & 1165 (BR 7982–28); Fungurume, 1350 m alt., 26 Feb. 1986, J. Schreurs 1211 (BR 8015–61).
Kenya. Coast Prov., Kwale distr., Arabuko-Sokoke Forest Reserve, plot 5, 15 May 1987, M.H. Ivory 674 (K(M) 38466); Ibid., plot 1, 29 Nov. 1994, M.H. Ivory 5036 (BR 49371–95).
Malawi. Northern Prov., Mzimba distr., Mtanga tanga Forest, 15 March 2005, F. Noe 05/538, (GENT); Ibid., Ntanga Forest Reserve, 30 Jan. 2005, A. Verbeken 05-147 (GENT); Ibid., Perekezi Forest Reserve, 28 Jan. 2005, A. Verbeken 05-093 (GENT); Ibid., 29 Jan. 2005, F. Noe 05/11 (GENT); Ibid., 13 Feb. 2005, F. Noe 05/225 (GENT); Ibid., 18 Feb. 2005, F. Noe 05/351 (GENT); Ibid., 9 March 2005, F. Noe 05/453 (GENT); Ibid., 11 March 2005, F. Noe 05/496 (GENT); Ibid., 16 March 2005, F. Noe 05/615 (GENT); Southern Prov., Machinga distr., Liwonde Forest Reserve, 22 Jan. 2005, A. Verbeken 05-019 (GENT).
Tanzania. Southern Prov., Songea distr., 29 km from Songea towards Mbinga, 29 Jan. 1993, T. Saarimäki & al. 1454 (H); Southern Highlands, Njombe distr., Mbalali village, 11 km S of Kidugala, 1 Feb. 1993, T. Saarimäki & al. 1515 (H).
Zambia. Copperbelt Prov., Misaka Forest Reserve, near Kitwe, April 1991, R. Watling 25920 (E); Feb. 1982, G. Piearce FP712/1 (K(M) 38498); Luapala Prov., near Kawamba, road near Ntumbachushi falls, 3 Jan. 1991, B. Buyck 3431 (PC); Mansa, Kabunda mission, 6 Jan. 1993, B. Buyck 3353 (PC); Western Prov., Mwinilunga, 29 Jan. 1991, B. Buyck 3507 (PC); Northern Prov., Mpika distr., North Luangwa National Park, Mwaleshi camp, 18 Jan. 1995, D. Shah‑Smith 126 (K(M) 35510); farm Gibsons, 20 Jan. 1996, B. Buyck & G. Eyssartier 96-147 (PC); Chibuli, 31 Jan. 1996, B. Buyck & G. Eyssartier 96-254 (PC).
Zimbabwe. Manicaland Prov., Chipinge, Busi farm, around Scott’s house, 12 Feb. 1999, A. Verbeken 99-185 (GENT).
Notes. The macroscopic description is based on fieldnotes of Verbeken and Schreurs; the microscopic description is mainly based on the specimens collected by Verbeken in Burundi and Zimbabwe.
The species is characterized by the very thick, whitish and stellately rupturing pileipellis, which even in exsiccata remains recognizable.
This species is locally eaten in the miombo area (e.g. Karhula & al. 1998).
Lactarius sect. Chamaeleontini Verbeken, Mycotaxon 66: 393 (1998)
− Type species: Lactarius chamaeleontinus R. Heim
Note. The distinction between this section and L. sect. Lactariopsis is based on absence versus presence of annular veil. This grouping is artificial but maintained for practical reasons.
Key to tropical African species
1. Hymenial leptocystidia rather abundant, 65–80 × 8–10 µm, cylindrical to subclavate, moniliform to rostrate. Veil present in young basidiomes as an arachnoid indusium between the stipe and the margin of the pileus, sometimes remaining as a very small annulus in older basidiomes; thick-walled hair-shaped elements in the pileipellis (very) scarce; context changing pinkish or reddish, then greyish green .............................................................. 6. L. indusiatus
Pleurocystidia absent; thick-walled hair-shaped elements in pileipellis absent to abundant; velar remnants never forming an annulus or absent .................................................. 2
2. Pileipellis without thick-walled elements ................................................................. 10. L. laevigatus
Pileipellis with more or less abundant hair-shaped, thick-walled elements, sometimes difficult to find ................... 3
3. Fruit body firm; cap dirty white to yellowish cream, often covered with soil particles ........ 13. L. emergens
Not with this combination of characters ............................................................. 4
4. Young cap and stipe entirely whitish pruinose, thick-walled terminal elements in pileipellis numerous ........ 11. L. pruinatus
Cap and stipe not entirely pruinose, thick-walled terminal elements of the pileipellis scarce or numerous ......................... 5
5. Small species, cap < 2 cm diam.; all terminal elements of pileipellis thick-walled, subfusiform to subcylindrical, 15–35 × 6–9 µm, spores on average 10 µm long .................... 12. L. uapacae
Cap most often > 2 cm diam.; thick-walled elements in pileipellis scattered, longer and slender, spores on average shorter ... 6
6. Basidiocarp more or less entirely bright yellowish orange, often sticky; pseudocystidia tapering or finely capitate. Species associated with Brachystegia spp. ................... 14. L. brachystegiae
Basidiocarp differently coloured; pseudocystidia not finely capitate .............................................. 7
7. Spore ornamentation low, not more than 0.2 µm high .......................................... 9. L. sesemotani
Spore ornamentation up to 0.5 µm high ................................................ 7
8. Pileus up to 5 cm diam., thin and fragile, pale orange, ochraceous or reddish brown; margin regularly striate to grooved ....................................................................... 7. L. chamaeleontinus
Pileus up to 8 cm diam., firmer, pale to straw yellow; margin irregular ....... 8. L. madagascariensis
Species descriptions
6. Lactarius indusiatus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 201 (1996). – Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 11 March 1994, A. Verbeken 94–122 (BR 57599–78, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 151-152 & Pl. 97-99 (1996d); Verbeken, Mycotaxon 55: 394-399 (1998).
Pileus 30–65 mm diam., rather thick, firm, convex to applanate, then infundibuliform; margin involute when young, then straight, sometimes slightly crenulate; pellis not or hardly dehiscent, smooth, soon desiccating, greyish orange to pale orange or leather-brown in the centre (6E4–5) and greyish yellow (4B3–4) near the margin. Lamellae slightly decurrent, unequal with lamellulae of different lengths (1–3 lamellulae between two lamellae), rather distant (L+l = 4+2/cm), moderately broad (2–4 mm), thick, firm, tough, without venation, very seldom dichotomously branched, orange-yellow, pale orange to light orange (5A3–4); edge entire, slightly darker. Stipe 24–43 × 7–19 mm, cylindrical, rather short and broad, slightly tapering downwards, smooth, greyish orange to pale orange, brownish orange (5C3) near the base, yellowish grey (4B2) in the upper part or paler towards the base and the apex, firm, solid. Context firm, white, changing salmon pink or dull red (8C3), greyish green when cut or bruised; smell agreeable, sweet; taste acrid, burning. Latex scarce, white; taste acrid. Velum present in young specimens as an arachnoid indusium between the stipe and the margin of the pileus; sometimes remaining as a small annulus in the older specimens. Spore-deposit not observed. – Pl. 4
Chemical reactions. Context unchanging with FeSO4.
Basidiospores subglobose to ellipsoid, 6.9–7.8–8.2–9.1 × 5.5–6.3–7.0–8.0 mm (Q = 1.09–1.16–1.22–1.38, n = 40); almost smooth; ornamentation very faint, composed of some very small warts and ridges; plage inamyloid. Basidia (50–)60–80 × 8–9(–10) mm, cylindrical, long, slender, 4-spored, sometimes 2-spored. Pleuroleptocystidia rather abundant, 65–80 × 8–10 mm, cylindrical to subclavate, moniliform to rostrate, thin-walled to slightly thick-walled; content guttate. Pleuropseudocystidia abundant, 6–9 mm diam., cylindrical to subfusiform, tapering upwards, often emergent; content oleiferic, slightly needle-like. Lamellar edge sterile; marginal cells long, slender, cylindrical, with rounded apex, 20–35 × 3–5(–6) mm, often septate, thin-walled, hyaline; pseudocystidia present. Hymenophoral trama composed of sphaerocytes and rather abundant lactifers. Pileipellis a mixed trichopalisade; terminal elements 10–20(–25) × 4–10(–12) mm, subclavate to clavate, thin-walled, hyaline; hair-shaped, thick-walled elements scarce, but present. Stipitipellis a trichoderm; terminal elements 15–50(–60) × 4–6 mm, long, cylindrical or tapering upwards, thin-walled or slightly thick-walled, hyaline, recumbent or ascending. Velum composed of long, narrow, parallel, thin-walled hyphae, 2–4(–5) mm diam., septate. Clamp-connections absent. – Fig. 30
Ecology. Miombo woodland.
Distribution. Only known from Burundi.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, 11 March 1994, A. Verbeken 94-122 (BR 57599–78, holotype; GENT, isotype); Nkayamba, just N of Rumonge, 12 March 1992, B. Buyck 4211 (PC); Ibid., 26 March 1992, B. Buyck 4290 (PC).
Notes. The macroscopic description is based on fieldnotes of Verbeken. The microscopic description is based on Verbeken 94-122, Buyck 4211 & 4290.
Besides the presence of an indusium, Lactarius indusiatus differs especially from L. laevigatus by the changing context, the longer basidia, the extremely lowly ornamented spores, the immediately burning acrid context and the presence of thick-walled hairs in the pileipellis.
7. Lactarius chamaeleontinus R. Heim
Heim, Bull. Jard. Bot. Etat Bxl 25: 87 (1955). – Type: Democratic Republic of Congo, Binga, 11 April 1934, M. Goossens-Fontana 942 (BR 7721–58, holotype).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 87–89 & Pl. 6, fig. 4a–c (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 15, fig. 8 (1955); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 156–157 & Pl. 107–109 (1996d).
Pileus 20–40(–50) mm diam., thin, convex, then depressed; margin inflexed, deeply, regularly striate to grooved; pellis dehiscent, dry, very finely tomentose, pale orange, ochraceous or reddish brown. Lamellae adnate to decurrent and even prolonging on the stipe, unequal with lamellulae of different lengths (one lamellula between two lamellae), sometimes dichotomously branched, distant (L+l = 3+1 to 3+2/cm), rather broad (3–7 mm), thin, brittle, pale yellow, pale orange, ochraceous yellow; edge entire, concolorous. Stipe 40–65 × 3–8 mm, cylindrical, slender, sometimes sinuose, tapering downwards, very finely tomentose, pale orange, ochraceous or reddish brown, whitish at the apex and at the base, soon fistulous. Context granular, spongy, brittle, white, unchanging. Latex white, becoming ochraceous. Spore-deposit white. – Pl. 5
Chemical reactions. Context between orange and green with FeSO4, intense and soon blue-green with gaiac (Goossens-Fontana 4030), faintly and slowly blue-green with gaiac (Buyck 3218), unchanging with NH4OH, slowly violet-blue with phenol (Goossens-Fontana 4030).
Spores subglobose to ellipsoid, 8.8–9.4–9.5–10.3(–10.5) × (6.8–)7.0–7.7–8.3–8.8 mm (Q = 1.06–1.15–1.23–1.35, n = 60); ornamentation amyloid, composed of irregularly sized and shaped warts, up to 0.5 mm high, sometimes aligned, mostly connected by fine connective lines, forming an incomplete reticulum; wall rugose, slightly amyloid; plage sometimes with a central amyloid spot. Basidia 45–50 × 10–13 mm, subclavate to clavate, 4-spored, rarely 2-spored; content guttate, sometimes needle-like. Pleurocystidia absent. Pleuropseudocystidia scarce, emergent, 12–16 mm, subclavate or fusiform, rounded or tapering at apex, sometimes mucronate; content oleiferic, guttate. Lamellar edge sterile; marginal cells 20–35 × 5–10(–12) mm, subcylindrical, clavate, fusiform or irregularly tortuous, often septate, thin-walled, hyaline. Hymenophoral trama composed of sphaerocytes and lactifers. Pileipellis a mixed trichopalisade; suprapellis composed of short clavate to subcylindrical, thin-walled, hyaline elements, 10–15(–30) × 5–10(–15) mm, forming chains; and long, slender, hair-shaped, thick-walled elements, 20–90(–110) × 4–7 mm; subpellis ixocutis-like, composed of slender, narrow, thin-walled, more or less parallel hyphae, 2–4 mm diam. Stipitipellis a cutis, with some arising, hair-shaped, thick-walled terminal elements, 20–90(–120) × 4–7 mm; lactifers abundant. Clamp-connections absent. – Fig. 31
Ecology. Found in different forest types, such as Guineo-Congolian rainforest with Gilbertiodendron dewevrei, wetter Zambezian miombo woodland (found under Brachystegia, Baphia, Julbernardia etc.), dry Sudanian woodland with Afzelia africana.
Revised specimens
Benin. Atacora Prov., Bassila, 4 Oct. 2000, A. De Kesel 2998 (BR 129210–06).
Democratic Republic of Congo. Equateur Prov., Binga, 11 Apr. 1931, M. Goossens-Fontana 942 (BR 7721–58, holotype); Ibid., Dec. 1945, M. Goossens-Fontana 4030 (BR 5002–55).
Zambia. Copperbelt Prov., Chati‑forest near Kitwe, 21 Dec. 1990, B. Buyck 3218 (PC); Ibid., 22 Dec. 1990, B. Buyck 3235, 3245 & 3246 (all PC); Ibid., 14 Jan. 1991, B. Buyck 3472 (PC); Western Prov., Mwinilunga, road to Solwezi, km 50, 30 Jan. 1991, B. Buyck 3534 (PC).
Notes. The type-specimen consists of small pieces of basidiomes and is in very bad condition. The macroscopic description is based on the fieldnotes of Goossens-Fontana 942 & 4030 and Buyck 3218. The microscopic description is based on Goossens-Fontana 942 & 4030 and Buyck 3218 & 3534.
The taste and smell of the context is described as acrid (?) in Goossens-Fontana 942; mild, then slightly acrid in Goossens-Fontana 4030. In Buyck 3218 the taste is said to be mild, the smell particular and chemical.
The Zambezian specimens have slightly larger basidiomes and the stipe is not as slender.
This species looks like a small version of Lactarius sesemotani, from which it also differs by the clearly bigger spores.
8. Lactarius madagascariensis Verbeken & Buyck
Buyck & al., Mycol. Res. 111: 791 (2007). – Type: Madagascar, Ranomafana National Park, 3. Feb. 1999, B. Buyck & G. Eyssartier 99-417 (PC, holotype; GENT, isotype).
Pileus 40–80 mm diam., planoconcave, soon depressed to irregular infundibuliform; surface smooth, shiny, pale to straw yellow, darker in the center, paler near the margin; margin distinctly and strongly striate to sulcate, up to half of the pileus radius. Stipe 24–44 × 12–16 mm, cylindric, concolorous with pileus or slightly paler. Lamellae broadly adnate to decurrent and decurrent with tooth, concolorous with pileus, very distant, with lamellulae, locally irregularly forked and with anastomosing veins, very broad (up to 1 cm). Context whitish to pale yellowish, unchanging, thin-fleshy in pileus, brittle in stipe; taste mild; smell not remarkable. Latex absent. – Pl. 5
Spores broadly ellipsoid, 8.0–8.6–8.9–10.1 × 6.6–7.3–7.5–8.5 µm (Q = 1.08–1.19–1.20–1.32, n = 40); ornamentation amyloid, composed of low, irregular warts, up to 0.5 µm high, sometimes aligned, but rarely connected with thin ridges; plage not amyloid. Basidia 50–65 × 9–11 µm, subclavate, 4-spored; sterigmata 6–9 × 2–3 µm. Pleurocystidia absent. Pleuropseudocystidia rare, slightly emergent, irregularly cylindrical, 7–11 µm diam., with rounded, slightly tapering apex, with dense, needle-like content. Lamellar edge sterile; marginal cells 12–35 × 5–8 µm, cylindrical, somewhat irregular and slightly tortuous, thin-walled, hyaline. Hymenophoral trama cellular. Pileipellis an irregular and loosely interwoven cutis to trichoderm, up to 100 µm thick, with abundant, thick-walled, septate hyphae orientated in all directions. Stipitipellis an irregular and loosely interwoven, thin cutis, resting on a layer of sphaerocytes. Clamp-connections absent. – Fig. 32
Ecology. Associated with Uapaca louvellii in wet mountain forest.
Distribution. Only know from the type locality.
Revised specimens
Madagascar. Ranomafana National Park, 3 Feb. 1999, Buyck & Eyssartier 99-409 (PC), 99-417 (PC, holotype; GENT, isotype), 99-424, 99-431 & 99-432 (all PC).
Notes. Lactarius madagascariensis differs from L. chamaeleontinus by the somewhat larger, firmer and paler basidiocarps and the more irregular cap margin, which is regularly striate to grooved in L. chamaeleontinus. The spores of L. madagascariensis are somewhat shorter (compared to 8.8–9.4–9.5–10.5 µm long in L. chamaeleontinus) and the pileipellis lacks the rounded to short clavate cells present in L. chamaeleontinus. Lactarius madagascariensis shares most of its field-characters with L. sesemotani, from which it is easily distinguished by the heavier ornamented spores: spore ornamentation of L. sesemotani never exceeds 0.2 µm height, while the spore ornamentation in L. madagascariensis is composed of warts up to 0.5 µm. In the field, Lactarius madagascariensis could be mistaken also for a representative of Russula sect. Archeinae. This is due to a general similarity in habitus and to the absence of latex exudation in this species. Microscopically, the abundance of lactifers in the gill trama places this species unambiguously in the genus Lactarius.
9. Lactarius sesemotani (Beeli) Buyck
Buyck, Bull. Jard. Bot. Nat. Belg. 59: 241 (1989). – Russula sesemotani Beeli, Bull. Soc. Roy. Bot. Belg. 60: 169 (1928). − Type: Democratic Republic of Congo, Diobo Akuba, Dec. 1925, M. Goossens-Fontana 505a (BR 5019–72, lectotype).
Selected descriptions and icons. Verbeken, Edinburgh J. Bot. 53: 66–69 (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 153–155 & Pl. 100–103 (1996d).
Pileus 50–100(–130) mm diam., thick, fleshy, hemispherical to convex when young, then applanate and infundibuliform; margin incurved when young, straight when older, strongly and deeply sulcate; pellis dehiscent, smooth, shiny, viscose, finely fibrous in the centre, yellowish ochre, then ochraceous brown (4A4–7 to 5AB4), paler towards the margin (3A2), greyish when older. Lamellae adnate, decurrent when older, unequal with a few lamellulae (0–1 between 2 lamellae), fairly distant (L+l = 3+1 to 5+1/cm), rather broad (7–9 mm), rather thick, sometimes forked, cream to pale yellowish (4A3–4); edge entire, concolorous. Stipe (25–)40–100 × 15–25 mm, cylindrical, subbulbous, faintly striate, faintly velvety, whitish, pale ochre, firm, solid, later sometimes fistulous. Context firm, white to cream, unchanging, but turning pale orange under the pileipellis; taste mild, but then acrid by the latex; smell not remarkable, agreeable. Latex scarce, white, fluid; taste acrid. Spore-deposit pale, but not purely white. – Pl. 6
Chemical reactions. Context unchanging or faintly greyish brown with FeSO4, unchanging with HCl, NaOH and NH4OH.
Basidiospores mostly ellipsoid, sometimes subglobose, 7.2–8.2–8.6–9.7 × 5.6–6.7–7.2–7.9 mm (Q = 1.09–1.19–1.24–1.43; n = 80); ornamentation amyloid, composed of irregularly shaped and sized warts, aligned or connected by fine lines, never forming a reticulum, not exceeding 0.2 mm height; plage sometimes with a faintly amyloid spot in the centre. Basidia 35–40 × 8–10 mm, cylindrical to subclavate, 4-spored, sometimes 2-spored. Pleurocystidia absent. Pleuropseudocystidia rather scarce, 8–12 mm diam., cylindrical to fusiform, with rounded or tapering apex, emergent up to 30 mm; content oleiferic or granular and needle-like. Lamellar edge sterile; marginal cells 15–40 × 5–8 mm, fusiform, tapering upwards, often septate, sometimes branched. Hymenophoral trama composed of sphaerocytes and lactifers. Pileipellis a mixed trichopalisade; suprapellis composed of cylindrical or isodiametric to fusiform, hyaline elements, 10–20 × 5–8 mm, and hair-shaped, slender, long, slightly tortuous elements with thickened wall (1 mm), 50–120 × 4–8 mm, sometimes branched; subpellis composed of loosely interwoven, hyaline hyphae, (1–)2–3 mm diam., forming an ixocutis. Stipitipellis a cutis with some ascending, hair-shaped, thick-walled terminal elements, 20–100 × 4–7 mm; lactifers abundant. Clamp-connections absent. – Fig. 33
Ecology. Found in several forest types, such as swamp forest, drier Guineo-Congolian rainforest, wetter Zambezian miombo woodland.
Distribution. Widespread in tropical Africa. Known from Burundi, Democratic Republic of Congo, Ivory Coast, Malawi, Zambia and Zimbabwe.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, near Rumonge, March 1994, A. Verbeken 94-082 (BR 57592–71), 94-154 (BR 57608–87), 94-155 (BR 57609–88), 94-471 (BR 57677–59) & 94-476 (BR 57682–64).
Democratic Republic of Congo. Equateur Prov., Eala, May 1923, M. Goossens-Fontana 143a (BR 5020–73); Djongo Akuba, Dec. 1925, M. Goossens-Fontana 505a (BR 5019–72, lectotype); Shaba Prov., Mengé, 4 April 1986, J. Schreurs 1592 (BR 8761–31).
Ivory Coast. Abidjan Prov., Forêt du Bianco, 4 Jan. 1976, Aké Assi 435 (K(M) 38465).
Malawi. Northern Prov., Mzuzu distr., Katoto E, 31 March 1973, Pawek in J. Williamson 638 (K(M) 38479).
Zambia. Copperbelt Prov., Chati forest, 14 Jan. 1991, B. Buyck 3471 (PC); Western Prov., Mwinilunga, road to Solwezi km 50, 30 Jan. 1991, B. Buyck 3531 (PC); Northern Prov., Mpika distr., North Luangwa National Park, Mwaleshi camp, 9 Feb. 1995, D. Shah‑Smith 199 (K(M) 35504); Chibuli, 31 Jan. 1996, B. Buyck & G. Eyssartier 96-249 (PC).
Zimbabwe. Mashonaland East Prov., Bromley, Liemba farm, 3 Feb. 1999, A. Verbeken 99-107 (GENT); Manicaland Prov., Eastern Highlands, Mguzu, 5 Feb. 1999, A. De Kesel 2409 (BR 112579–59).
Notes. The original description has been based on a heterogeneous exsiccatum. Both Goossens-Fontana 143 & 505 are a mixture of Russula and Lactarius material. One water-colour of Goossens-Fontana 505 represents clearly the Lactarius, the water-colour of Goossens-Fontana 143 represents a smaller Russula. We separated the collections renumbering the material as Goossens-Fontana 143a & 143b and Goossens-Fontana 505a & 505b. Goossens-Fontana 505a is designated as lectotype (Verbeken 1996a). The specimens are in rather good condition and agree perfectly with the more recent collected material from Democratic Republic of Congo and Zambia.
The macroscopic description is based on the field notes of Goossens-Fontana, Buyck and Verbeken. The microscopic description is based on Goossens-Fontana 143a & 505a, Buyck 3471 and A. Verbeken 94-082.
In the field this species can very easily be confused with Lactarius laevigatus. It differs mainly by the presence of thick-walled hair-like elements in the pileipellis. The species differs from Lactarius chamaeleontinus by its firmer and bigger habit and its smaller spores.
10. Lactarius laevigatus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 203 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 31 March 1994, A. Verbeken 94-535 (BR 57694–76, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 155–156 & Pl. 104–106 (1996d); Karhula & al., Karstenia 38: fig. 19 (1998).
Pileus 10–70 mm, thick, but brittle, infundibuliform; margin deflexed upwards, strongly striate to sulcate; pellis not dehiscent, smooth, wet, not viscose, finely fibrous in the centre, light orange to greyish orange (4A5 to 5AB5), locally more yellow, darker when older. Lamellae decurrent, unequal with mostly 1 lamellula between 2 lamellae, rather distant (L+l = 2+2 to 3+4/cm), rather thick, broad (7 mm), very brittle, with numerous small veins, sometimes anastomosing, pale yellow (3A3); edge entire, concolorous. Stipe 40–50 × 10–20 mm, cylindrical, tapering towards the base, dry, wrinkling towards the apex and the base, pale yellow (4A3), solid but brittle. Context white to cream; taste mild, then acrid; smell not particular. Latex scarce, white. Spore-deposit white. – Pl. 7
Chemical reactions. Context unchanging or greyish to greenish with FeSO4, faintly greenish blue with gaiac, unchanging with NH4OH. Pileipellis olive-yellow with KOH.
Basidiospores ellipsoid, 7.9–8.9–9.3–9.9(–10.2) × 6.4–7.2–7.3–7.9 mm (n = 80, Q = 1.15–1.23–1.28–1.37); ornamentation amyloid, composed of irregularly shaped and sized warts, aligned or connected by fine connective lines, never forming a reticulum, very low, not exceeding 0.2 mm height; plage sometimes with a faintly amyloid spot in the centre. Basidia (40–)50–60 × 9–12 mm, subclavate to clavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia rather abundant, 10–17 mm diam., cylindrical to fusiform, sometimes mucronate, emergent up to 50 mm; content densely oleiferic and needle-like. Lamellar edge sterile; marginal cells 20–40 × 4–7 mm, cylindrical, with rounded apex, slender, sometimes branched. Hymenophoral trama composed of sphaerocytes and scarce lactifers, with narrow, hyaline hyphae towards the edge. Pileipellis ixocutis-like to trichoderm-like, with recumbent and locally ascending chains of short cylindrical elements, 20–30(–40) × 5–10 mm, thin-walled or very slightly thick-walled; terminal elements rounded, sometimes mucronate, remarkably dark and band-wise coloured in cresyl-blue. Stipitipellis cutis to ixocutis, composed of narrow, thin-walled hyphae, 2–10 mm diam., often septate. Clamp-connections absent. – Fig. 34
Ecology. Zambezian miombo woodland. Collected under Caesalpiniaceae and Uapaca spp.
Distribution. Known from Burundi, Malawi, Tanzania, Zambia and Zimbabwe.
Revised specimens
Burundi. Bururi Prov., Rumonge Forest Reserve, Nyamirambo, May 1993, B. Buyck 5138 (PC); Ibid., March 1994, A. Verbeken 94-082b (BR 57593–72), 94-163 (BR 57616–95) & 94-535 (BR 57694–76, holotype; GENT, isotype); Ibid., 1 April 1994, A. Verbeken 94-579 (BR 57706–88); Nkayamba, just N of Rumonge, 11 Dec. 1992, B. Buyck 4737 (PC); Mugara, 29 Nov.1978, J. Rammeloo 5932 (BR 8775–45).
Malawi. Southern Prov., Blantyre, ± 1000 m, Dec. 1971, Williamson 536 (K(M) 38473).
Zambia. Copperbelt Prov., Chati‑forest, near Kitwe, Dec. 1990, B. Buyck 3117-3121, 3123-3125, 3185, 3207 & 3211 (all PC).
Zimbabwe. Mashonaland East Prov., Liemba farm, 2 Feb. 1999, A. Verbeken 99-079 & 99-080 (all GENT); Midlands Prov., Mvuma, Central Estates, paddock 77, 5 Feb. 1999, A. Verbeken 99-127 & 99-129 (all GENT); Masvingo Prov., south of Lake Mutirikwe, around Norma Jean’s lodge, 16 Feb. 1999, A. Verbeken 99-223 (GENT).
Notes. On a young basidiome of Buyck 3211, an indusium (referred to as "cortina" in the fieldnotes) is observed. It becomes a fugitive and very thin annulus near the apex of the stipe, which disappears later.
Both species, Lactarius sesemotani and L. laevigatus are recognized by the striate or sulcate and shiny pileus, even in the exsiccata. Lactarius laevigatus differs from L. sesemotani by the lack of thick-walled and hair-shaped elements in the pileipellis, but resembles the species in every other character. This difference is not correlated with the state of development (thick-walled hairs disappearing with age), since both species are studied in both young and old condition.
This species is locally consumed, e.g. in Tanzania (Karhula & al. 1998).
11. Lactarius pruinatus Verbeken & Buyck
Verbeken, Mycotaxon 66: 399 (1998). − Type: Zambia, Copperbelt, Chati-forest near Kitwe, 24 Dec. 1990, B. Buyck 3248 (GENT, holotype; PC, isotype).
Selected descriptions and icons. Verbeken, Mycotaxon 66: 399–403 (1998).
Pileus 17–25 mm diam., almost applanate, slightly depressed in the centre, with central papilla; margin straight, not really striate but radially wrinkled when drying; pellis mat, dry, pruinose, with very short hairs, yellowish brown to pale brown (4AB6 to 5C4–6). Stipe 18–25 × 5–10 mm, strongly flattened, with irregular folds and a bit tortuous, fragile and completely hollow; pellis velvety, with short hairs, white to cream, pure white at the base. Lamellae clearly decurrent, moderately distant, with lamellulae, cream (3A2–3); edge entire, concolorous. Context white to cream, unchanging; taste very acrid. Latex white, unchanging or a bit watery; taste very acrid. Spore-deposit white. – Pl. 8
Chemical reactions. Context unchanging to pale greyish with FeSO4.
Basidiospores subglobose to ellipsoid, 8.4–9.8–11 × 7.4–8.2–9.0 mm (n = 20, Q = 1.07–1.19–1.30); ornamentation amyloid, very regularly reticulate, composed of ridges, up to 0.5(–1) mm high, forming an almost complete to complete reticulum; wall clearly and typically rugose between the meshes; plage not amyloid. Basidia 4-spored, seldom 2-spored, 55–65 × 12–14 mm, subclavate; sterigmata 8–10 × 2–3 mm. Pleurocystidia absent. Pleuropseudocystidia not very abundant, mostly emergent up to 35 mm, 7–18 mm broad, fusiform, sometimes with moniliform apex, thin-walled or with slightly refringent walls; content slightly needle-like. Lamellar edge sterile: marginal cells cylindric or fusiform, 17–40 × 2–7 mm, thin-walled, hyaline. Hymenophoral trama composed of sphaerocytes mainly; some hyaline hyphae and lactifers present. Subhymenium cellular. Pileipellis a lamprotrichoderm; elements of the suprapellis cylindric to subcylindric, with obtuse or more or less acute apex, 20–50(–70) × 4–7 mm, thick-walled and sometimes septate; subpellis composed of hyaline hyphae. Stipitipellis a lamprotrichoderm; elements of the suprapellis cylindric to subcylindric, with obtuse or more or less acute apex, sometimes branched, 20–80 × 3–10 mm, thick-walled and sometimes septate; subpellis composed of hyaline hyphae. Clamp-connections absent. – Fig. 35
Ecology. Found in wetter Zambezian miombo woodland.
Distribution. Only known from type locality.
Revised specimens
Zambia. Copperbelt Prov., near Kitwe, Chati-forest, 24 Dec. 1990, B. Buyck 3248 (GENT, holotype; PC, isotype).
12. Lactarius uapacae Verbeken & Stubbe
Verbeken & al., Cryptog. Mycol. 29: 140 (2008). − Type: Cameroon, forêt de Dja, 30 Jan. 1993, A. Verbeken 07- 48 (GENT, holotype).
Selected descriptions and icons. Verbeken & al., Cryptog. Mycol. 29: 140-142 (2008).
Pileus 10–14 mm diam., applanate with slight depression or slightly curved upwards in the centre; surface transparently striate-grooved, minutely felty, opaque, slightly greasy, lacking veins, very pale yellow to yellowish white (4A3) at the margin and when older, in centre and in young specimens slightly darker buff-yellowish to pinkish buff. Lamellae broadly adnate, distant, with 3 lamellulae between 2 lamellae, with regular short-long-short pattern, yellowish white; edge entire, concolorous. Stipe 7–10 × 1–3 mm, subcyindrical or strongly curved, tapering downwards, sometimes irregular, pale buff to whitish. Context very thin-fleshed, hollow in stipe, dirty pale yellow, unchanging; smell not remarkable; taste unknown. Latex not observed. – Pl. 8
Spores subglobose to broadly ellipsoid, 8.5–9.9–11.4 × 7.8–8.9–9.9 µm (Q = 1.05–1.12–1.22, n = 20); ornamentation amyloid, subreticulate, composed of wings up to 1(–1.5) µm high; plage not amyloid. Basidia 35–45 × 12–15 µm, (2) 4-spored, clavate. Pleurocystidia absent. Pseudopleurocystidia abundant, very emergent, locally up to 20 µm diam., fusiform or irregular, sometimes branching. Lamellar edge sterile; marginal cells 20–37 × 4–6 µm, subfusiform to subcylindrical, thin-walled, hyaline. Hymenophoral trama cellular, composed of rather large sphaerocytes and lactifers. Pileipellis a lampropalisade, 60–80 µm thick; terminal elements subfusiform to subcylindrical, 15–35 × 6–9 µm, thick-walled; subpellis 40–50 µm thick. Stipitipellis a lamprotrichopalisade to lamprotrichoderm; terminal elements longer and more variable than in the pileus. – Fig. 36
Ecology. Guineo-Congolian rainforest. Found on branchlets under Uapaca.
Distribution. Only known from the type locality.
Revised specimens
Cameroon. Eastern Prov., Réserve de faune du Dja, campement dans la brousse, 10 April 2007, A. Verbeken 07-48 (GENT, holotype).
Notes. Striking characteristics for this species are the pale yellow and small basidiocarps. Microscopically the species is characterized by the large spores with subreticulate ornamentation. The broad and emergent pseudocystidia are typical for representatives of Lactarius sect. Chamaeleontini Verbeken, as are the yellowish colours and the sulcate pileus. The species differs from all other members of this section by its small basidiocarps and the pure lampropalisade.
13. Lactarius emergens Verbeken
Verbeken & al., Syst. Geogr. Pl. 70: 192 (2000). − Type: Zimbabwe, Mvuma, Central Estates, Beacon Hill section, 26 Jan. 1999, A. Verbeken 99-006 (GENT, holotype).
Selected descriptions and icons. Verbeken & al., Syst. Geogr. Pl. 70: 192–194 (2000).
Pileus 40–80 mm diam., convex to planoconvex and slightly depressed, a bit irregular; pellis smooth, glabrous, radially subfibrillose, rather sticky, yellowish white to light yellow (4A2–5), darkest in the center, paler outside; margin first smooth, strongly striate, grooved in older specimens (up to 10 mm from margin). Lamellae slightly decurrent, very distant (L+l = 7–9/cm at midradius in young basidiomes, L+l = 4/cm in older ones), rather thick, brittle, anastomosing at some places, yellowish white (3A2 to 4A4); edge slightly darker. Stipe 30–50 × 12–17 mm, more or less cylindric, a bit curved and irregular; pellis smooth, glabrous, white, becoming pale greyish. Context rather thick but thin near margin, with 2 chambers in stipe, white to cream, unchanging; smell agreeable, reminding of Russula fellea; taste acrid, but disappearing soon. Latex very scarce, white, unchanging, unchanging with KOH. Spore-deposit not observed. – Pl. 9
Chemical reactions. Context very faint blueish grey with gaiac, unchanging with FeSO4.
Basidiospores subglobose to ellipsoid, 7.0–8.0–9.0–9.9 × 6.0–6.6–6.7–7.3 µm (Q = 1.08–1.21–1.33–1.43, n = 40); ornamentation amyloid, extremely weakly developed, composed of low warts (less than 0.2 µm high) and some ridges; plage not amyloid. Basidia 50–65 × 9–11 µm, 4-spored, subclavate, with guttate content; sterigmata very short (up to 5(–7) µm) and thick (up to 3 µm). Pleurocystidia absent. Pleuropseudocystidia rather abundant, sometimes emergent, up to 10(–12) µm diam., tortuous, subcylindric, with tapering apex, with needle-like content. Hymenophoral trama cellular and with abundant lactifers. Lamellar edge sterile; marginal cells subcylindric, subfusiform, 15–25 × 6–9 µm, thin-walled, hyaline. Pileipellis trichopalisade-like, with abundant thick-walled elements, which are long, cylindric, with rounded but tapering apex, 40–90 × 4–6 µm, often septate; some swollen to subglobose elements present; diverticulate knobs present on some terminal elements and some hyphae. Stipitipellis a trichoderm, composed of thin-walled hyphae where some knobs are present. Clamp-connections absent. – Fig. 37
Ecology. Found in drier Zambezian Brachystegia woodland and under Uapaca in Isoberlinia woodland.
Distribution. Known from Benin and Zimbabwe.
Revised specimens
Benin. Borgou Prov., Wari Maro, 19 June 2000, A. De Kesel 2834 (BR 126387–93).
Zimbabwe. Midlands Prov., Mvuma, Lovedale Ranch, Beacon Hill section, at the ostrich incubator, on the ridge, 26 Jan. 1999, A. Verbeken 99-005 (GENT), 99-006 (GENT, holotype) & 99-012 (GENT).
Notes. The species is closely related to Lactarius sesemotani (Beeli) Buyck (Lactarius section Chamaeleontini Verbeken) from which it differs macroscopically by the paler and more whitish colours and the chambered stipe. Microscopically the thick-walled hairs in the pileipellis of L. sesemotani are more pronounced (walls 1 µm thick) and longer. Diverticulate knobs have never been observed in the pileipellis-elements of L. sesemotani. Pleuropseudocystidia are more abundant in Lactarius emergens and less emerging as in L. sesemotani. The basidia are clearly larger in L. emergens (35–40 × 8–10 µm in L. sesemotani) and the spores have never been observed to have an amyloid spot on the plage as in L. sesemotani.
Lactarius emergens emerges typically under the soil and pushes the soil upwards when growing mature. Much soil remnants remain on the rather sticky cap.
14. Lactarius brachystegiae Verbeken & C. Sharp
Verbeken & al., Syst. Geogr. Pl. 70: 187 (2000). – Type: Zimbabwe, Mvuma, Central Estates, Beacon Hill, 26 Jan. 1999, A. Verbeken 99-002 (GENT, holotype).
Selected descriptions and icons. Verbeken & al., Syst. Geogr. Pl. 70: 187–190 (2000); Verbeken, Micol. Veget. Medit. 16: fig. 3 (2001).
Pileus 32–110 mm diam., first convex, undeeply depressed and with strongly recurved margin, then becoming more planoconvex and with a deeper depression, finally infundibuliform and with an irregular, wavy margin; margin straight in older specimens, grooved, striate in older specimens (up to 20 mm long); pellis especially in young specimens soft, pruinose, almost velvety and in center sometimes very finely cracked, but towards margin and also when older smooth, glabrous, a bit sticky (leave remnants sticking on the pileus) and radially fibrillose, never really wet, golden yellow-orange to deep orange (5AB6–7 to 5A8), light yellow to reddish yellow (4A5–6) near margin, light brown to reddish brown (6D7 to 8D8) in center, when rottening dark reddish brown from margin on. Lamellae broadly adnate to shortly decurrent, rather spaced (L+l = 6–7/cm halfway), with a lot of short lamellulae, anastomosing and furcated lamellae present, 3–6 mm broad, rather thick, brittle, pale orange to orange white (5A2–3); edge staining a bit darker. Stipe 10–35 × 10–22 mm, relatively short, cylindrical, in some specimens distinctly broader at the base, in others distinctly narrower (like a triangle); pellis smooth, soft, glabrous, with very fine, darker, vertical lines (like varicose veins), pale yellow to pale orange (4A2–3 to 5A2–3), some with darker spots (but no scrobicules), some with a whitish zone at the extreme base and some with a slightly darker narrow zone at the extreme top. Context thick and firm, up to 4 mm in pileus, solid, hard and robust in stipe (sometimes soft because of damage by insects), white to very pale orange, especially under the pileipellis, pale, whitish to cream in the stipe, a bit more flesh-coloured after a while, smell apple-like, strongly sweetish, like Russula fellea, Lactarius evosmus; taste first mild, soon acrid and slightly astringent, burning, but disappearing soon. Latex very scarce to rather abundant, watery white, unchanging (Verbeken 99-004) or golden yellow (Verbeken 99-132) with KOH; taste as context. Spore-deposit not observed. – Pl. 10
Chemical reactions. Context bright blue after some minutes with gaiac, very slightly salmon or unchanging with FeSO4.
Basidiospores broadly ellipsoid to ellispoid, 9.5–10.5–10.6–11.5 × 7.6–8.2–8.4–9.0 µm (Q = 1.15–1.26–1.28–1.38, n = 40); ornamentation amyloid, composed of irregular, more or less rounded warts, sometimes aligned, sometimes connected by fine lines, about 0.5 µm high; plage not amyloid. Basidia 65–75 × 10–12 µm, subcylindric to subclavate, 4-spored; sterigmata 5–8 × 2–4 µm. Pleurocystidia absent. Pleuropseudocystidia abundant, emergent up to 40 µm above the hymenium, up to 18 µm broad, fusiform to clavate but with tapering or capitate apex; content needle-like. Hymenophoral trama cellular, with abundant lactifers. Lamellar edge sterile, composed of marginal cells; marginal cells 13–30 × 6–10 µm; irregularly cylindric to subclavate, thin-walled, hyaline. Pileipellis (ixo)cutis-like but with abundant swollen compartiments; terminal elements mostly pericline and usually cylindric, sometimes clavate or irregular, 20–50 × 5–8 µm; some slightly thick-walled elements present; pseudocystidia present, 8–10 µm diam., with needle-like content, capitate; diverticulate knobs or bulges present on both terminal elements and hyphae of the subjacent layer. Stipitipellis a trichoderm; terminal elements mostly anticline, thin-walled, cylindric to subclavate, 12–40 × 5–7 µm, with diverticulate knobs. Clamp-connections absent. – Fig. 38
Ecology. Associated with Brachystegia species (e.g. B. glaucescens, B. spiciformis) in Zambezian miombo woodland. In central Zimbabwe Lactarius brachystegiae seems a common species. It was mostly observed in well-developed Brachystegia leaf litter, especially under Brachystegia glaucescens on hills.
Distribution. Known from Malawi, Tanzania and Zimbabwe.
Revised specimens
Malawi. Southern Prov., Makwawa, Zomba, 7 Jan. 1991, B. Morris 91-23 (PC); Ibid., 9 April 1994, B. Buyck 3978-3981 (all PC).
Tanzania. Southern Highlands Prov., Mbeya distr., Ipembe Hill, 28 March 1991, T. Saarimäki & al. 747 (H, judging from photograph in unpublished thesis of Karhula 1997: 62).
Zimbabwe. Midlands Prov., Mvuma, Central Estates, Beacon Hill Homestead, garden Sharp, 26 Jan. 1999, A. Verbeken 99-002 (GENT, holotype); Ibid., 4 Jan. 1997, C. Sharp 518/97 (GENT); Central Estates, Beacon hill section, at the ostrich incubator, 26 Jan. 1999, A. Verbeken 99-004 (GENT); Ibid., 9 Jan. 1997, C. Sharp 539/97 (GENT); Midlands Prov., Mvuma, Central Estates, along road north of the ostrich incubator, leg. R. Walleyn 6 Feb. 1999, A. Verbeken 99-132 (GENT).
Notes. The description is based on the collections from Zimbabwe.
This species is included here in Lactarius sect. Chamaeleontini but the characters of the pileipellis are more reminescent of species of L. sect. Russulopsideae.
Lactarius sect. Russulopsidei Verbeken, Mycotaxon 77: 441 (2001).
− Type species: Lactarius ruvubuensis Verbeken.
Key to tropical African species
1. Hymenophoral trama mainly composed of sphaerocytes .................................................................. 2
Hymenophoral trama irregular to regular, mainly composed of hyaline, more or less parallel hyphae, occasionally with a few sphaerocytes ....................... 6
2. Spore-ornamentation composed of conical, pointed to slightly rounded spines, up to 1.5 mm high, connected by fine connective lines, forming an almost complete reticulum ... 22. L. corbula
Spore-ornamentation composed of irregular, rounded and obtuse warts, isolated or connected by fine connective lines ..... 3
3. Lamellar edge reddish brown ........................................................................... 19. L. rufomarginatus
Lamellar edge concolorous ..................................................... 4
4. Taste mild; elements of the pileipellis without remarkable bulges ..... 20. L. pseudotorminosus
Taste very acrid; elements of the pileipellis with remarkable diverticulate bulges ........................ 5
5. Stipe shortly cylindrical; lamellae rather distant (3+7 to 4+8/cm); pileus becoming greyish dark brown from the margin to the centre when old and rottened; latex unchanging; pleurocystidia mostly present and abundant ....... 15. L. ruvubuensis
Stipe long and slender; lamellae very crowded (22–32/cm); pileus becoming paler when older; latex sometimes greyish to blackish when drying; pleurocystidia absent ...... 16. L. longipes
6. Latex blue-greenish when drying; spores very lowly ornamented; pileus unicolorous .......... 17. L. cyanovirescens
Latex unchanging when drying; spore-ornamentation 0.2–0.5 mm high; pileus clearly darker in the centre and paler towards the margin ................. 7
7. Pileipellis with clearly moniliform dermatocystidia; lamellae crowded and narrow (2–2.5 mm); taste of latex slightly acrid ............................................................................................... 21. L. claricolor
Some of the elements of the pileipellis with lactiferous content and reminding dermatocystidia; lamellae rather distant and broader (3–13 mm); taste of latex burning acrid ...... 18. L. urens
Species descriptions
15. Lactarius ruvubuensis Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 208 (1996). − Type: Burundi, Ruvubu National Park, 6 Apr. 1994, A. Verbeken 94-599 (BR 57708–90, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 207–208 & Pl. 208–215 (1996d).
Pileus (40–)55–90 mm diam., quite thick, slightly to deeply infundibuliform; margin incurved to deflexed downwards when young, then straight and deflexed upwards, striate when older; pellis dehiscent at the margin, dry, finely tomentose in the centre, finely fibrose towards the margin, light brown in the centre (5D5–6), brownish orange to reddish towards the margin (6C4–6), becoming greyish dark brown from the margin to the centre when old and rottened. Lamellae strongly decurrent, unequal with lamellulae between 2 lamellae (1–3, irregular pattern), dichotomous near the margin, quite distant (L+l = 3+7 to 4+8/cm), thick, medium broad (3–8 mm), brittle, sometimes with finely anastomosing veins, white; edge entire, concolorous. Stipe (20–)35–45 × 6–22 mm, short, tapering downwards, dry, smooth, reddish brown to brownish orange, broadly fistulous. Context firm in pileus, brittle in stipe, white, changing greyish brown when rottening; taste faint mild to bitter, then acrid; smell apple-like, sweet, unpleasantly sweet when older. Latex abundant, fluid, white, unchanging; taste very acrid. Spore-deposit white. – Pl. 11
Chemical reactions. Context salmon, light orange with FeSO4. Pileipellis red with KOH.
Basidiospores subglobose to ellipsoid, 7.1–7.9–8.1–8.7(–8.9) × 6.3–6.8–7.0–7.6 mm (Q = 1.07–1.13–1.17–1.27; n = 80); ornamentation amyloid, composed of irregular warts and connective lines, never forming a reticulum, some warts isolated, up to 0.3 mm high; plage inamyloid. Basidia 45–60 × 9–10 mm, slightly clavate, 4-spored. Pleurocystidia scarce to abundant, 35–50 × 3–5 mm, tortuous to irregularly branched and diverticulate. Pleuropseudocystidia scarce to abundant, 3–5 mm diam., tortuous to irregularly branched and diverticulate. Lamellar edge sterile; marginal cells 25–35(–45) × 3–6 mm, irregularly clavate to tortuous, often diverticulate and branched at the apex. Hymenophoral trama composed of isodiametric cells; lactifers scarce to rather abundant, in some specimens even absent. Pileipellis a cutis, composed of hyaline hyphae, often septate, 3–7 mm diam., with diverticulate bulges; lactifers present; dermatocystidia present, capitate, 35–45 × 4–6 mm, with needle-like content (dark-brown pigmentation in upper layer). Stipitipellis a trichoderm; terminal elements fusiform; at the base of the stipe consisting of slender, often dichotomously branched, slightly thick-walled hyphae. Clamp-connections absent. – Fig. 39
Ecology. Found in damp forest, under Uapaca guineensis.
Distribution. Only known from Burundi and Guinea.
Revised specimens
Burundi. Ruyigi Prov., Ruvubu National Park, April 1994, A. Verbeken 94-598 (BR 57707–89), 94-599 (BR 57708–90, holotype; GENT, isotype), 94-600 (BR 57709–91), 94-601(BR 57710–92) & 94-617 (BR 57711–93).
Guinea. Forêt humide de Ziama, 31 July 2000, A. Bâ 185 (GENT); Ibid., 1 Aug. 2000, A. Bâ 305 (GENT).
Notes. Both macroscopic and microscopic descriptions are based on the collections from Burundi.
In some specimens (e.g. Verbeken 94-599 & 94-617) lactifers in the trama are rather abundant and pleuropseudocystidia are also rather abundant, while true pleurocystidia are scarce. In other specimens (e.g. Verbeken 94-600) lactifers and pseudocystida could not be observed in the trama of the lamella, which makes the trama look exactly like that of a Russula species. There was latex observed in fresh condition, and there are lactifers in the trama of the pileus. The true pleurocystidia become more abundant as the pseudocystidia are absent. Those true cystidia have exactly the same size and shape as the pseudocystidia in the other specimens. They are not connected with lactiferous hyphae, although they arise rather deep in the hymenium.
16. Lactarius longipes Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 203 (1996). − Type: Democratic Republic of Congo, Kivu, Irangi, 6 April 1972, J. Rammeloo Z242 (GENT, holotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 208–209 & Pl. 215–217 (1996d).
Pileus 30–60 mm diam., firm, moderately thick, regular, slightly depressed to infundibuliform; margin sharp, involute when young, slightly striate and irregularly undulate when old; pellis adnate, not dehiscent, mat, almost smooth, slightly concentrically zonate, radially wrinkled, not unicolorous, with darker and brighter spots of reddish golden and brownish orange (6C8), light brown (6D6–8), dark brown (6F6–8) or darker in the centre and paler towards the margin, greyish orange to reddish blond (5BC6) when older. Lamellae adnate to subdecurrent, unequal with numerous lamellulae of different lengths, frequently forked, very dense (22–32/cm), very narrow (1–1.5 mm), thin, greasy, pale yellow (1A3) to yellowish white (3A2–3) when young, orange white (5A3) when older; edge entire, concolorous. Stipe 35–80 × 6–15 mm, cylindrical, long and slender, sometimes tapering downwards, slightly longitudinally grooved, light brown to brown in upper part (6DE8), brownish orange to reddish golden in lower part (6C5–6), whitish near the base, firm, solid. Context firm, white, yellowish to ochraceous under the pellis, unchanging; smell sickly or sweetish, particular; taste very acrid. Latex abundant, white, becoming transparent, sometimes greyish to blackish when drying; taste very acrid. Spore-deposit cream to pale yellowish (not observed in fresh condition). – Pl. 12
Chemical reactions on context. Anilated H2O: pale ochraceous, pale brown to brown after a while. Phenol, TL4 NH4OH: nihil.
Basidiospores subglobose to ellipsoid, 6.7–7.5–7.9–8.8 × 5.0–6.5–6.6–7.3 mm (Q = 1.06–1.14–1.15–1.24, n = 60); ornamentation amyloid, composed of irregular warts, isolated or connected by fine connective lines, up to 0.5 mm high; plage inamyloid or with slightly amyloid spot. Basidia 35–45 × 7–10 mm, cylindrical to narrowly clavate, 4-spored, seldom 2-spored. Pleurocystidia absent. Pleuropseudocystidia abundant, emergent, 4–5(–7) mm diam., cylindrical and tortuous, often branched and with bulges; content oleiferic to needle-like. Lamellar edge sterile; marginal cells 15–30 × 4–6(–7) mm, cylindrical, tortuous, sometimes septate, hyaline, thin-walled. Hymenophoral trama composed of sphaerocytes; narrow, hyaline hyphae near the edge; lactifers abundant. Pileipellis a cutis, composed of hyaline hyphae, often septate, 3–7 mm diam., sometimes tortuous or with some bulges; lactifers present, penetrating in the upper layer; dermatocystidia very abundant, capitate or subcapitate, (25–)30–45 × 4–6 mm, with needle-like content. Stipitipellis idem as pileipellis; abundant caulocystidia. Clamp-connections absent. – Fig. 40
Ecology. Found in Guineo‑Congolian rainforest with Gilbertiodendron dewevrei and Scaphopetalum thonneri, and riverine forest with Uapaca guineensis.
Distribution. Known from the Democratic Republic of Congo, Togo and Gabon.
Revised specimens
Democratic Republic of Congo. Tshopo Prov., 5 km NNE of Batiabongena, April 1984, B. Buyck 1336 (BR 8766–36); Ibid., B. Buyck 1345 (BR 12745–38); Ibid., B. Buyck 1346 (BR 12746–39); Kivu Prov., Irangi, March 1972, J. Rammeloo Z228 (GENT); Ibid., April 1972, J. Rammeloo Z242 (GENT, holotype); Ibid., June 1972, J. Rammeloo Z271 (GENT).
Gabon. Research Station of Ipassa-Makokou, border of Ivindo river, 21 March 2005, J. Degreef 303 (BR 164551–39); Research Station of Ipassa-Makokou, 25 March 2005, M. Nguele 29 (BR 164545–33).
Togo. Central Prov., Forêt Classée de Alédjo, Gallery forest with Berlinia grandiflora, Uapaca guineensis, Pentadesma and Breonadia, 12 July 2007, A. Dekesel 4315 (BR 163825–89).
Notes. The type consists of 5 basidiomes in good condition.
The species reminds Lactarius claricolor by the habit and the dermatocystidia, differs however because of the darker colour of the pileus and the stipe, the very acrid taste, the heteromerous trama (subregular in L. claricolor) and the spore ornamentation (which looks more pronounced in L. kivuensis, but was more difficult to observe in L. claricolor, due to the alcohol-conservation).
The species is closely related to Lactarius ruvubuensis. They are microscopically almost identical, especially with regard to the abundant pileocystidia and the very remarkable shape of the pleurocystidia. However in Lactarius longipes, the pleurocystidia are never true cystidia, only pseudocystidia, and the bulges and diverticulate elements in the pileipellis are scarce, whereas they are very abundant in L. ruvubuensis. These differences, combined with the more striking difference in number and spacing of the lamellae, make the two taxa specifically different.
According to Rammeloo (herbarium notes), this species is locally eaten in the Democratic Republic of Congo.
17. Lactarius cyanovirescens Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 199 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 11 March 1994, A. Verbeken 94-080 (BR 57590–69, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 214–215 & Pl. 233–237 (1996d).
Pileus 65–95 mm diam., firm, fleshy, convex to planoconvex and depressed in the centre to slightly infundibuliform; margin incurved, then straight and irregularly crenulate, faintly striate or sulcate; pellis not or slightly dehiscent, smooth, dry to very slightly sticky, mat, brownish orange to light brown (6D7), greyish red (7C3 to 7D4 or 8CD4), dull red (8B3–4), locally or in the centre paler (3AB3). Lamellae adnate to subdecurrent, unequal with lamellulae of different lengths (1–3 lamellulae between two lamellae), exceptionally forked, distant (L+l = 2+1, 3+8 to 4+4/cm), moderately broad to broad (4–9 mm), thick, brittle, sometimes with venation towards the margin of the pileus, white to cream, pale yellow (3A3) or pale brownish yellow (4A4), staining blue-green to greyish green by the latex; edge entire, concolorous. Stipe 25–40 × 8–20 mm, shortly cylindrical, tapering downwards, smooth, pale yellow (4A4–5), pale orange, orange to reddish brown (6CD5), whitish near the apex, with a whitish felty layer at the base, soon chambered. Context firm, 4–7 mm in pileus, white, staining blue-green by latex-contact; taste mild; smell sometimes agreeable, sometimes slightly H2S-like. Latex scarce to rather abundant, white, blue-green (25C4–7) when drying; taste spicy to very acrid. Spore-deposit white to pale cream. – Pl. 12
Chemical reactions. Context slowly salmon red with FeSO4. Pileipellis pale yellowish green with KOH.
Basidiospores ellipsoid, 7.3–8.2–8.6–9.4 × 6.3–6.9–7.2–8.2 mm (Q = 1.04–1.16–1.23–1.33, n = 100); ornamentation not very distinct, very low and irregular, composed of some irregularly shaped and sized warts, often aligned or connected by connective lines; plage inamyloid. Basidia 40–55 × 9–10(–11) mm, cylindrical to subclavate, 4-spored; content guttate. Pleurocystidia absent. Pleuropseudocystidia rather abundant, emergent, cylindrical, with rounded apex; content needle-like. Lamellar edge sterile; marginal cells 25–55 × 4–7(–10) mm, tortuous, with bulges, sometimes branched, often septate, thin-walled, hyaline. Hymenophoral trama heteromerous, composed of sphaerocytes, a few hyaline, thin-walled hyphae and abundant lactifers. Pileipellis a cutis, composed of a layer of interwoven, more or less parallel hyphae, thin-walled, hyaline, septate, 3–5 mm diam, locally swollen up to 15 mm; pileocystidia scarce, 20–30 × 3–5 mm, with very dense oleiferic yellowish brown content. Stipitipellis trichoderm-like to subcellular with hymeniform aspect, composed of a few isodiametric to more elongated cells, mostly forming chains; terminal elements 15–35 × 2–5(–7) mm, cylindrical, slender, thin-walled; base of the stipe covered with long and slender hyphae, 3–5(–8) mm diam. Clamp-connections absent. – Fig. 41
Ecology. Miombo woodland, dry evergreen forest (muhulu), often under Brachystegia spp.
Distribution. Known from Burundi, Democratic Republic of Congo, Malawi, Tanzania and Zambia.
Revised specimens
Burundi. Bururi Prov., Rumonge Forest Reserve, Nyamirambo, Nov. 1993, B. Buyck 5191 & 5225 (all PC); Ibid., March 1994, A. Verbeken 94-080 (BR 57590–69, holotype; GENT, isotype), 94-300 (BR 57642–24), 94-514 (BR 57684–66), 94-515 (BR 57685–67) & 94-516 (BR 57686–68); Ibid., Nkayamba, near Rumonge, Dec. 1991, B. Buyck 4045 & 4046 (all PC); Ibid., Jan. 1992, B. Buyck 4174 (PC); Ibid., April 1992, B. Buyck 4379 & 4428 (all PC); Ibid., March 1993, B. Buyck 4984, 4989 & 4999 (all PC); Ibid., April 1994, A. Verbeken 94-151 (BR 57605–84), 94-152 (BR 57606–85) & 94-164 (BR 57617–96); Cankuzo Prov., Murango, Dec. 1994, B. Buyck 5665 (PC).
Democratic Republic of Congo. Shaba Prov., Kipopo, 5 Dec. 1971, D. Thoen 5089 (BR 66274–23); Shaba Prov., Lubumbashi, Feb. 1986, J. Schreurs 1055 (BR 7896–39); Shaba Prov., Luiswishi, 14 Dec. 1971, D. Thoen 5127 (BR 66281–30).
Malawi. Northern Prov., Mzimba distr., Mtanga tanga forest, 15 March 2005, F. Noe 05/562-565 & 05/567 (all GENT); Ibid., 16 March 2005, F. Noe 05/629-633 (all GENT); Perekezi Forest Reserve, 9 March 2005, F. Noe 05/454 & 05/466 (all GENT); Ibid., 16 March 2005, F. Noe 05/634 (GENT).
Zambia. Northern Prov., Mpika distr., North Luangwa National Park, Mwaleshi Camp, 10 Feb. 1995, D. Shah‑Smith 214 (K(M) 35512); Mbereshi National Forest, NF 265, 22 Feb. 1998, herb. M. Groenendaal L2; farm Gibsons, 19 Jan. 1996, B. Buyck & G. Eyssartier 96-124 (PC); Lake Kashyba, 24 Jan. 1996, B. Buyck & G. Eyssartier 96-167 (PC).
Notes. The macroscopic description is based on fieldnotes from Verbeken, Buyck and Schreurs; the microscopic description is especially based on Verbeken 94-080 & 94-151, Schreurs 1055 and Buyck 4045.
The taste of the context was noted to be acrid in Schreurs 1055.
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 211 (1996). – Type: Burundi, Mugara, 2 Dec. 1978, J. Rammeloo 6002 (BR 18874–56, holotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 211–213 & Pl. 223–229 (1996d); Verbeken & al., Syst. Geogr. Pl. 70: 208–211 (2000).
Pileus 40–120(–150) mm diam., firm, medium thick, convex to planoconvex, with a depressed centre to infundibuliform when older; margin involute to incurved when young, then waving and irregularly crenulate, broadly and longly striate when old; pellis dehiscent up to halfway the radius, finely felty, with very small squamules, smooth to even shiny when old, with remarkable difference in colour between the centre and the margin, brownish orange to light brown (6CD5–7) in the centre, greyish orange (5CD7), golden yellow (5B4–6), buff to cream (3A2) in the outer zone, with remaining reddish brown squamules. Lamellae adnate when young, decurrent when older, unequal with lamellulae of different lengths (1–3 lamellulae between two lamellae, if three then regular short-long-short pattern), moderately distant to distant (L+l = 2+2 to 5+5/cm), moderately broad to broad [(3–)5–10(–13) mm)], rather thick, firm to brittle, sometimes with venation, seldom dichotomously branched towards the apex of the stipe, cream to pale yellow; edge entire, concolorous. Stipe (20–)30–55 × 8–20(–25) mm, shortly cylindrical, tapering downwards, finely felty to smooth, slightly wrinkling, brownish orange when young (6C3–4 to 6D4–5), greyish orange (5A3 to 5B4) when old, paler at the apex, solid, firm. Context firm, white, changing to cream (4A2) when cut; smell agreeable and fruit-like to unagreeable and slightly chemical; taste mild and nut-like in the stipe, often acrid by the latex. Latex rather abundant, white, fluid, burning acrid, turning bright yellow-orange with KOH. Spore-deposit white (0–Ia). – Pl. 13
Chemical reactions. Context immediately dark blue with gaiac, weakly salmon with FeSO4, unchanging with formol, NH4OH and phenol.
Basidiospores ellipsoid, 7.7–8.6–9.4–10.2 × 6.4–7.1–7.8–8.5 mm (Q = 1.11–1.20–1.21–1.30, n = 80); ornamentation amyloid, composed of rather regular and rounded warts, connected by fine connective lines, forming an almost complete reticulum; warts up to 0.5 mm high; plage inamyloid. Basidia 55–70 × 10–12 mm, subclavate, 4-spored; content often guttate, sometimes needle-like. Pleurocystidia very abundant, emergent, 65–130 × 8–10 mm, cylindrical to fusiform, rounded, tapering or mucronate at the apex; content needle-like or granular; arising often very deep in the hymenium and thus easily to confuse with pseudocystidia. Pleuropseudocystidia very abundant, 7–10 mm diam., cylindrical to fusiform, mostly with rounded apex; content needle-like or granular. Lamellar edge sterile; cheilopseudocystidia present; marginal cells 16–40 × 5–12 mm, cylindrical to clavate, thin-walled, with guttulate content. Hymenophoral trama irregular, composed of interwoven hyaline hyphae and lactifers. Pileipellis one-layered, composed of interwoven horizontal to ascending hyphae; hyphae thin-walled to very slightly thick-walled, septate, often branched or with short bulges; terminal elements 20–80 × 5–8 mm, cylindrical to tortuous, with rounded apex, with yellowish or reddish brown content (only in the centre of the pileus); laticiferous hyphae abundant in the upper layer; pileocystidia rather abundant, never capitate, cylindrical, with a laticiferous content, 40–80 × 5–7 mm. Stipitipellis idem as pileipellis. Clamp-connections absent. – Fig. 42
Ecology. Zambezian miombo woodland. Found under Caesalpiniaceae and Uapaca. In Central Zimbabwe, Lactarius urens is confined to richer miombo woodland with Brachystegia glaucescens on hill ridges. Rather common.
Distribution. Known from Burundi, Democratic Republic of Congo, Kenya, Malawi and Zambia, Zimbabwe.
Revised specimens
Burundi. Bururi Prov., Rumonge Forest Reserve, Nyamirambo, 9 May 1993, B. Buyck 5089 & 5090 (PC); Ibid., 18 May 1993, B. Buyck 5136 (PC); Ibid., 7 Dec. 1993, B. Buyck 5265 (PC); Ibid., 7 March 1994, A. Verbeken 94-008 (BR 57560–39), 94-014 (BR 57565–44) & 94-016 (BR 57566–45); Ibid., 11 March 1994, A. Verbeken 94-070 (BR 57581–60) & 94-081 (BR 57591–70); Ibid., 15 March 1994, A. Verbeken 94-156 (BR 57610–89); Ibid., 18 March 1994, A. Verbeken 94-212 (BR 57621–03); Ibid., 23 March 1994, A. Verbeken 94-295 (BR 57641–23); Ibid., 25 March 1994, A. Verbeken 94-383 (BR 57652–34) & 94-384 (BR 57653–35); Ibid., 29 March 1994, A. Verbeken 94-443 (BR 57667–49); Ibid., 30 March 1994, A. Verbeken 94-473 (BR 57680–62); Ibid., 31 March 1994, A. Verbeken 94-518 (BR 57688–70); Ibid., 1 April 1994, A. Verbeken 94-558 (BR 57701–83); Mugara, in garden, 16 Nov. 1978, J. Rammeloo 5712 (BR 8767–37); Mugara, 2 Dec. 1978, J. Rammeloo 6002 (BR 18874–56, holotype); Nkayamba, Rumonge, 18 Dec. 1976, F. Dossin 76 (BR 6114–03); Ibid., 14 Dec. 1991, B. Buyck 4026 (PC); Ibid., 12 March 1992, B. Buyck 4245 (PC); Ibid., 20 April 1992, B. Buyck 4377 (PC); Ibid., 29 April 1992, B. Buyck 4424 & 4434 (all PC); Ibid., 15 March 1994, A. Verbeken 94-162 (BR 57615–94); Cankuzo Prov., Murango, 11 March 1994, B. Nzigidahera 59 (PC); Ibid., Dec. 1994, B. Buyck 5625, 5648, 5651 & 5696 (all PC).
Democratic Republic of Congo. Shaba Prov., along road between Lubumbashi and Likasi, 2 Feb. 1986, J. Schreurs 1000 (BR 7849–89); Tshilongo, 3 March 1986, J. Schreurs 1248 (BR 8046–92); Kipopo, 25 km WNW of Lubumbashi, 20 Feb. 1971, D. Thoen 4548 (BR 66224–70); Ibid., 6 March 1971, D. Thoen 4622 (BR 8763–33).
Kenya. Coast Prov., Kwale distr., Arabuko-Sokoke forest reserve, plot 1, 3 Aug 1994, M.H. Ivory 936 (BR 49372–96).
Malawi. Southern Prov., Zomba distr., near R.C. Cathedrale, 5 Jan. 1974, J. Williamson 763 (K(M) 38483); Mulanje distr., nabij Likaubula village, 9 Feb. 2005, F. Noe 05/137 (GENT); Northern Prov., Mzimba distr., Perekezi forest reserve, 28 Jan. 2005, A. Verbeken 05-096 (GENT); Ibid., 18 Feb. 2005, F. Noe 05/353 (GENT); Ibid., 9 March 2005, F. Noe 05/492, 05/460, 05/465 & 05/474-475 (all GENT); Ibid., Ntanga tanga forest reserve, 31 Jan. 2005, F. Noe 05/20 (GENT); Ibid., 15 March 2005, F. Noe 05/551, 05/553 & 05/576-578 (all GENT);
Tanzania. Lake Prov., Mwanza distr., W of Geita, 13 Jan. 1974, D.L. Ebbels F98 (K(M) 38490).
Zambia. Luapala Prov., Mansa, Kwanfumwe, 5 Jan. 1991, B. Buyck 3400 (PC); near Kawambwa, 1431 m alt., 6 Jan. 1991, B. Buyck 3442 (PC); Copperbelt Prov., Mwekera‑forest, between Kitwe & Ndola, 1274 m alt., 21 Jan. 1991, B. Buyck 3481 (PC); Western Prov., Mwinilunga, 1484 m alt., 29 Jan. 1991, B. Buyck 3506 (PC); Northern Prov., Mpika distr., North Luangwa National Park, Mwaleshi Camp, 8 Feb. 1995, D. Shah‑Smith 182 (K(M) 35502); Mbereshi National Forest, 22 Feb. 1998, herb. M. Groenendaal L6.
Zimbabwe. Manicaland Prov., Chipinge, Busi farm, around Scott’s house, 12 Feb. 1999, A. Verbeken 99-201 (GENT); Midlands Prov., Mvuma, Central Estates, Beacon Hill section, at the ostrich incubator, 26 Jan. 1999, A. Verbeken 99-014 (GENT); Ibid., Beacon Hill ridge, 26 Jan. 1999, A. Verbeken 99-014 (GENT); Ibid., 28 Jan. 1999, A. Verbeken 99-038 (GENT).
Notes. The description is based on the type and collections of Verbeken from Burundi and Zimbabwe.
Lactarius urens is easily recognized by its two-coloured cap: reddish brown in the centre with a paler yellowish zone towards the margin, and by its extremely acrid taste. The latex causes a rather heavy burning sensation on unprotected skin, which may last more than one hour.
19. Lactarius rufomarginatus Verbeken & Van Rooij
Van Rooij, Nova Hedwigia 77: 235 (2003). − Type: Benin, Borgou Prov., Wari Maro, 21 June 2006, A. De Kesel 3011 (BR 129223–19, holotype; GENT, isotype).
Selected descriptions and icons. Van Rooij & al., Nova Hedwigia 77: 235–236, 244 (2003).
Pileus 61–83 mm diam., slightly infundibuliform to convex and broadly depressed in the centre; margin sometimes striate, undulate and slightly crenulate in older specimens, slightly pruinose in young basidiomes; pellis somewhat wax-like, mat; orange pink to orange brown when young, red to reddish brown when older (8D8–7D8). Lamellae broadly adnate to strongly decurrent, distant (5–8 L+l/cm), broad (6 mm), with a strong venation between the lamellae towards the stipe, forked and veined, pale yellow to yellowish orange (4A3–5A3); lamellulae with an irregular pattern (but usually one lamellula between 2 lamellae); edge reddish brown. Stipe 40–66 × 9–15 mm, long, slender and regularly cylindric; pellis tomentose and mat, orange (6B7–8) to orange brown (6C5–8), becoming orange white towards the base of the stipe, sometimes with a white mycelium. Context (very) thin-fleshed in pileus, solid in the stipe, yellowish white to slightly darker at margin; taste faintly acrid to mild. Latex not observed. Spore-deposit not observed. – Pl. 13
Basidiospores subglobose to broadly ellipsoid, sometimes globose or ellipsoid, 7.1–7.9–8.2–8.9 × 6.0–6.7–7.3–8.2 µm (Q = 1.02–1.12–1.17–1.33, n = 60); ornamentation amyloid, composed of warts up to 1 µm high, subspherical to subconical, aligned or connected with fine connective lines, forming a distinct reticulum, warts never isolated; plage inamyloid. Basidia 63–75 × 6–10 µm, cylindric to subclavate, sometimes slightly thick-walled, 4-spored (seldom 2-spored), content with oil-like droplets. Pleurocystidia absent. Pleuropseudocystidia abundant, emergent, (3–)5–9.5 µm diam., irregularly cylindrical, apex more or less moniliform, sometimes rounded; content needle-like, sometimes granular and with oil-like droplets. Lamellar edge sterile; marginal cells 20–30 × 4–14 µm, subcylindric, thin-walled, hyaline, with a reddish brown pigmentation. Hymenophoral trama cellular with abundant lactifers. Pileipellis ixotrichoderm-like; terminal elements 10–40 × 2–5(–7) µm, narrowly cylindric, fusiform to slightly conical, sometimes thick-walled; hyphae sometimes with little knobs, embedded in a thin slime-layer. Stipitipellis idem, terminal elements more frequently slightly thick-walled and not embedded in a slime-layer. Clamp-connections absent. – Fig. 43
Ecology. Sudanian riverine forest with Berlinia grandiflora, Lonchocarpus sericeus, Uapaca somon and Pterocarpus erinaceus.
Distribution. Only known from the type locality.
Revised specimens
Benin. Atacora Prov., Bassila, 21 June 2000, A. De Kesel 2843 (BR 126396–05); Atacora Prov., Bassila, 5 Oct 2000, A. De Kesel 3011 (BR 129223–19, holotype; GENT, isotype) ; Ibid. 11 June 2002, A. De Kesel 3358 (BR 152171–75) ; Ibid. 26 June 2002, A. De Kesel 3470 (BR 152263–70); Ibid., 27 June 2002, A. De Kesel 3481 (BR 152163–67); Ibid. 28 June 2002, A. De Kesel 3487 (BR 152172–76); Ibid., 7 Oct. 2002, A. De Kesel 3524 (BR 152004–05); Ibid. 15 June 2004, A. De Kesel 3613 (BR 157040–94).
Notes. The ixotrichoderm structure and the hyphae with little knobs are reminiscent of Lactarius urens Verbeken. The spore ornamentation with rounded warts, connected with fine lines and the irregular cylindrical pleuropseudocystidia are typical for Lactarius section Russulopsidei. Because of its reddish brown cap, the species can be placed next to L. cyanovirescens Verbeken, L. ruvubuensis Verbeken, L. longipes Verbeken and L. urens Verbeken, which all share this character.
20. Lactarius pseudotorminosus R. Heim
Heim, Candollea 7: 379 (1938). – Type: Madagascar, S of Soanierana, 8 Dec. 1934, R. Heim G84 (PC, holotype).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 54-56 (1938); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 209–210 & Pl. 218–220 (1996d).
Pileus 50–60 mm diam., rather thick (4–6 mm halfway radius), subtranslucid towards the margin, convex, then uneven and irregular, largely depressed when mature; margin incurved, not involute, straight or deflexed upwards when older; pellis dehiscent, completely and finely tomentose, dry, pale pinkish orange or bright orange. Lamellae decurrent, unequal with abundant lamellulae, rather crowded, narrow (2 mm), thick, cream; edge concolorous, irregularly sinuose. Stipe 30–50 × 10–15 mm, robust, firm, smooth, creamish orange or yellowish white, becoming ochraceous in alcohol and brownish at the base, solid. Context firm, whitish or cream; smell not particular; taste mild. Latex abundant, white; taste mild. Spore-deposit not observed. (According to Heim 1938b).
Chemical reactions. Context immediately lilac with FeSO4, intense and positive with gaiac and gaiacol, immediately olive-grey with pyramidon. (According to Heim 1938b).
Basidiospores ellipsoid, (6.2–)6.5–7.7–8.8 × (4.7–)5.3–6.2–7.1 mm (Q = 1.15–1.23–1.36, n = 30); ornamentation amyloid, composed of rounded, obtuse warts which are mostly isolated, sometimes aligned or connected by fine connective lines, up to 0.3 mm high; plage inamyloid. Basidia 34–45(–50) × 8–11 mm, cylindrical to subclavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia abundant, sometimes emergent, (2–)3–5 mm, subcylindrical to tortuous, obtuse or mucronate and tapering upwards, often branched; content oleiferic to granular. Lamellar edge sterile; some very short, deformed basidia present; marginal cells 15–20 × 5–8 mm, fusiform, cylindrical or subclavate, thin-walled; content yellowish, oleiferic. Hymenophoral trama heteromerous, composed of sphaerocytes and some parallel, narrow, hyaline hyphae; lactifers present. Pileipellis a cutis, composed of narrow, hyaline, interwoven hyphae; dermatocystidia abundant, 20–40 × 4–6 mm, cylindrical and mucronate; content granular. Stipitipellis cutis-like, composed of slightly thick-walled hyphae, septate and very frequently branched. Clamp-connections absent. – Fig. 44
Ecology. Malagasy lowland rainforest.
Distribution. Only known from the type locality.
Revised specimens
Madagascar. Betsimisaraka Prov., south of Soanierana, terrestrial, isolated, 8 Dec. 1934, R. Heim G84 (PC, holotype).
Notes. The type-specimen consists of one basidiome in quite good condition, but preserved in alcohol.
21. Lactarius claricolor R. Heim
Heim, Candollea 7: 381 (1938). – Type: Madagascar, North of Antanambé, 21 Dec. 1934, R. Heim J18bis (PC, holotype).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 62–64 & Pl. 1f (1938); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 210–211 & Pl. 221–222 (1996d).
Pileus 45–60 mm diam., rather thick in the centre, first subumbonate, then strongly depressed and irregular; margin sharp, inflexed, finely and rather long grooved or wrinkled; pellis not dehiscent, tough, almost smooth, finely spotted, dry, pinkish lilac, locally paler, paler at the margin. Lamellae subdecurrent, unequal with numerous lamellulae of different lengths (often short), crowded, narrow (2–2.5 mm), thin, frequently wrinkled in the upper part, ivory white (brownish lilac when preserved in alcohol). Stipe 50–70 × 8–14 mm, long, cylindrical to subfusiform, irregular, wrinkled to grooved, creamish orange to orange-ochraceous, tomentose and whitish at the base with large, membranaceous subiculum, solid. Context white (brownish ochraceous in alcohol), unchanging, tough; smell not particular; taste mild, then acrid. Latex not abundant, white; taste slightly acrid. Spore-deposit white. (According to Heim 1938b). – Pl. 14
Chemical reactions. Context immediately pink with FeSO4, positive with gaiac and gaiacol, slightly pale lilac with pyramidon. (According to Heim 1938b).
Basidiospores ellipsoid, (6.4–)6.8–7.3–7.8(–8.6) × (5.0–)5.3–5.8–6.3–7.3 mm (Q = 1.17–1.26–1.35, n = 30); ornamentation amyloid, composed of irregularly sized and shaped warts, 0.2–0.5 mm high, which isolated, aligned or connected by fine connective lines, never forming a complete reticulum; plage inamyloid. Basidia 35–45 × 7–9 mm, narrowly clavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia abundant, (2.5)3–4.5 mm diam., cylindrical to tortuous, with rounded apex, often branched; content densely oleiferic. Lamellar edge sterile; marginal cells 15–35 × 4–6 mm, cylindrical, subcylindrical or tortuous, 2- or 3-septate, with rounded apex, hyaline, thin-walled. Hymenophoral trama subregular, composed of hyaline, narrow hyphae, more or less parallel, lactifers present but rather scarce. Pileipellis composed of interwoven, recumbent to ascending hyphae; terminal cells 13–25 × 2–4 mm, cylindrical, with rounded apex; hyphae sometimes branched and with swollen and isodiametric elements; dermatocystidia rather abundant, 26–36 × 2–4 mm, long, slender, cylindrical, mucronate, capitulate, with some dark brown guttules. Stipitipellis idem. Clamp-connections absent. – Fig. 45
Ecology. Found in Malagasy lowland rainforest.
Distribution. Only known from the type locality.
Revised specimens
Madagascar. Betsimisaraka Prov., North of Antanambé, terrestrial, 21 Dec.1934, R. Heim J18bis (PC, holotype).
Notes. The type-specimen consists of one basidiome in rather good condition, preserved in alcohol, which makes it difficult to get a clear picture of the spore-ornamentation.
22. Lactarius corbula R. Heim & Gooss.‑Font.
Heim, Bull. Jard. Bot. Etat Bxl 25: 64 (1955). – Democratic Republic of Congo, Binga, Dec. 1941, M. Goossens-Fontana 2070 (BR 7728–65, holotype).
Misappl.: Lactarius gymnocarpus ss. Heim (1955) pro parte.
Selected descriptions and icons. Bull. Jard. Bot. Etat Bxl 25: 64–66, fig. 2a–b (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 13, fig. 1b & Pl. 15, fig. 5 (1955); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 213–214 & Pl. 230–233 (1996d).
Pileus 35–90 mm diam., fleshy, convex, then deeply depressed, finely infundibuliform, uneven in the centre; margin deeply grooved up to halfway the radius; pellis partly dehiscent, thin, dry, granulose in the centre, ochraceous brown. Lamellae decurrent, unequal with lamellulae of different lengths (one lamellula between two lamellae), distant (L+l = 2+2/cm), rather broad (up to 9 mm), thick, brittle, anastomosing with fine veins, pale yellow with lemmon shade. Stipe 20–35(–70) × 7–15 mm, shortly cylindrical, tapering downwards, whitish to pale yellowish, solid. Context white, then pale yellowish; taste very acrid, graphite-like; smell not very particular. Latex not abundant in the context, abundant in the lamellae, white, opaque, thick; taste acrid. Spore-deposit white (cream?). (According to Heim 1955a). – Pl. 14
Chemical reactions. Context unchanging with alfa-naftol, NH3 and aniline; slowly pinkish with FeSO4; immediately blue-green with gaiac; wine-colour with phenol and immediately chocolate-brown with pyrogallol. (According to Heim 1955a).
Basidiospores ellipsoid, 7.0–8.5–9.0–10.2(–10.4) × 5.3–6.8–7.3–8.5 mm (Q = 1.13–1.24–1.26–1.35, n = 60); ornamentation amyloid; composed of conical, pointed to slightly rounded spines, up to 1.5 mm, connected by fine connective lines, forming an almost complete reticulum; plage inamyloid. Basidia 43–60 × 7–12 mm, cylindrical to subclavate, 4-spored; content slightly granular. Pleurocystidia very abundant, often emergent, often arising deep in the subhymenium, 47–70 × 6–10 mm, subcylindrical, with rounded, tapering or slightly mucronate apex; content oleiferic, granular to needle-like. Pleuropseudocystidia rather abundant, often emergent, 6–10 mm diam., subcylindrical, rounded, with tapering or slightly mucronate apex; content oleiferic, needle-like. Lamellar edge sterile; marginal cells 13–37 × 4–6 mm, cylindrical, subclavate or fusiform, sometimes septate; content sometimes granular. Hymenophoral trama composed of narrow, hyaline hyphae and very abundant lactifers. Pileipellis cutis-like; terminal elements 30–50 × 5–7 mm, cylindrical, with rounded apex, sometimes with slightly thickened wall; pileocystida rather abundant, 20–60 × 5–9 mm, with granular or lactiferous content, never capitate. Stipitipellis idem. Clamp-connections absent. – Fig. 46
Ecology. Found in drier Guineo‑Congolian rainforest with Gilbertiodendron dewevrei.
Distribution. Only known from the type locality.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Binga, Apr. 1928, M. Goossens-Fontana 714 (BR 12325–06); Ibid., Sept. 1940, M. Goossens-Fontana 2042 (BR 4921–71); Ibid., Dec. 1941, M. Goossens-Fontana 2070 (BR 7728–65, holotype).
Notes. The type consists of numerous basidiomes in mouldy condition.
Goossens-Fontana 2042 shows the same microscopic characters, however the macroscopic description seems to be based on a Russula-specimen. Especially because of the lack of lamellulae, this can hardly be considered as a Lactarius-description, although, all the macrochemical reactions are the same as those of Goossens-Fontana 2070. Probably, a confusion of specimens has occurred. The microscopic description is based on all revised specimens.
The species reminds somewhat the species of Lactarius section Chamaeleontini by its deeply grooved pileus and its pileipellis- and stipitipellis-structure, mainly composed of thin-walled hyphae but with some thick-walled terminal elements. However, we do not include Lactarius corbula in L. section Chamaeleontini because of the different aspect of the pileus in the exsiccata (not typical shiny, but rather granulose), the presence of true pleurocystidia, the hymenophoral trama composed of hyphae and not of sphaerocytes and the more cutis-like pilei-and stipitipellis-structure.
The conspicuous spore-ornamentation, composed of conical, pointed spines, connected by fine connective lines, is only found in Lactarius corbula and L. russuliformis but the pileipellis structure of the latter is completely different.
Lactarius sect. Edules Verbeken, Belg. J. Bot. 132: 176 “1999” (2000).
Lactarius sect. Russuliformes Verbeken, Mycotaxon 77: 441 (2001).
− Type species: Lactarius edulis Verbeken & Buyck.
Notes. As molecular data indicate that L. roseolus fits into L. sect. Edules, the section Russuliformes (type species: L. roseolus) becomes a synonym.
Key to tropical African species
1. Pileipellis entirely composed of interwoven, long and slender hyphae ......................................... 2
Pileipellis composed of chains of subcylindrical to subspherical elements .................................. 3
2. Taste burning acrid; lamellae distant, pale yellow to golden yellow; pleurocystidia absent ......... 26. L. aureifolius
Taste mild; lamellae very crowded, cream-colour; pleurocystidia rather abundant ........... 27. L. densifolius
3. Pileus < 3 cm diam., pink to reddish brown, basidiocarp often brittle and delicate ........... 30. L. roseolus
Pileus larger, basidiocarp mostly firm ...................................... 4
4. Lamellae moderately to very distant; stipe short and thick ........................... 5
Lamellae (very) crowded; stipe long and slender ...................................................... 28. L. inversus
5. Pileus somewhat with an olivaceous shade; lamellae dichotomously branched towards the margin; margin of pileus translucently striate .......................................... 29. L. phlebophyllus
Pileus never with an olivaceous shade; lamellae not dichotomously branched; margin sometimes striate in old specimens, not translucid ............................................... 6
6. Pileus, especially when young, with reddish brown and cinnamon-brown colours; lamellae orange, yellowish orange to pale brownish orange; elements of pileipellis often with small, intracellular, dark granules ........... 25. L. latifolius
Pileus mainly yellowish to cream-colour, with reddish shade only present in exsiccatum; lamellae cream-colour; elements of pileipellis without intracellular granules ........................................ 7
7. Spore-ornamentation up to 0.3 µm high, composed of verrucose elements more or less connected; macrocystidia absent ..................... 23. L. edulis
Spore-ornamentation less pronounced, never higher than 0.2 µm; macrocystidia very abundant and as the pseudocystidia with knotty bulges ................................. 24. L. nodosicystidiosus
23. Lactarius edulis Verbeken & Buyck
Buyck, Champignons comestibles de l'Ouest du Burundi: 103 (1994). – Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 7 March 1994, A. Verbeken 94-004 (BR 57556–35, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Russulales News 3: 17–22 (1995); Härkonen & al., Karstenia 35, Suppl.: fig. 73 (1995); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 159–161 & Pl. 110–112 (1996d); Karhula & al., Karstenia 38: fig. 20 (1998); De Kesel & al., Guide des champignons comestibles du Bénin: 163 (2002); Härkonen & al., Norrlinia 10: 85 (2003).
Pileus 50–150 mm diam., firm and fleshy, planoconvex to applanate and depressed, irregular; margin incurved when young, then straight to irregularly crenulate, yellow to orange when young, paler when old; pellis not dehiscent, dry, slightly felty, cracking towards the margin when older, brownish or greyish orange to yellow, ochraceous, lemmon–colour or cream (5A5, 5B4–5, 6D4–5), with remarkable reddish tinge in exsiccatum. Lamellae slightly decurrent, unequal with lamellulae of different lengths (regular pattern, one or three lamellulae between two lamellae), distant (L+l = 4+3 to 3+7/cm), thick, broad (5–7 mm), cream; edge concolorous, entire. Stipe 30–50 × 20–30 mm, short cylindrical, firm, solid, dry, slightly felty, concolorous to pale orange or yellow (4A3–4, 5A3–4), paler towards the apex. Context very firm, solid, even hard in young basidiomes, white, unchanging, becomes first grey then black when rottening; smell agreeable, sweetish; taste mild. Latex fairly abundant, white; taste mild. Spore-deposit white. – Pl. 14
Chemical reactions. Context salmon orange with FeSO4.
Basidiospores ellipsoid, 6.8–7.6–7.8–8.3 × 5.6–6.2–6.4–7.0 mm (Q = 1.09–1.19–1.24–1.35, n = 60); ornamentation composed of verrucose, convex to cylindrical elements, < 0.3 mm high, more or less connected, sometimes forming short ridges; plage inamyloid. Basidia 60–80 × 9–11 mm, cylindrical to subclavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia abundant, 3–5 mm diam., cylindrical, rounded or mucronate. Lamellar edge sterile; marginal cells 15–25 × 5–7 mm, cylindrical, subclavate or slightly fusiform. Hymenophoral trama composed of sphaerocytes; some hyaline hyphae and lactifers present. Pileipellis a trichopalisade, 40–120 mm thick, composed of chains of subcylindrical to subspherical elements, 10–40 × 10–40 mm; terminal elements 10–30 × 5–10, cylindrical to subclavate, sometimes with slightly thickened walls. Stipitipellis a trichopalisade, 40–80 mm thick, composed of chains of subcylindrical to cylindrical elements, 20–30(–40) × 5–10 mm; terminal elements 10–30 × 8–12 mm, cylindrical to subclavate or fusiform. Clamp-connections absent. – Fig. 47
Ecology. Miombo woodland, found a.o. under Brachystegia and Uapaca spp.
Distribution. Widespread and common in the Zambezian region. Known from Benin, Burundi, Democratic Republic of Congo, Malawi, Tanzania, Zambia and Zimbabwe.
Revised specimens
Benin. Atacora Prov., Koussou Kouangou, 5 Sept. 1997, A. De Kesel 2063 (BR 74896–12).
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, April 1993, B. Buyck 5067 (PC); March 1994, A. Verbeken 94-004 (BR 57556–35, holotype; GENT, isotype), 94-009 (BR 57561–40), 94-017 (BR 57567–46), 94-079 (BR 57589–68), 94-083 (BR 57594–73), 94-143 (BR 57600–79), 94-144 (BR 57601–80), 94-214 (BR 57622–04), 94-217 (BR 57625–07), 94-275 (BR 57632–14), 94-276 (BR 57633–15) & 94-447 (BR 57668–50); Ibid., April 1994, A. Verbeken 94-573 (BR 57705–87); Nkayamba, just N of Rumonge, 8 April 1978, J. Lambinon 78/99 (LG); Ibid., April 1992, B. Buyck 4404 (PC); Ibid., Jan. 1993, B. Buyck 4923 (PC); Ibid., Feb. 1993, B. Buyck 4957, 4960 & 4970 (all PC); Ibid., March 1994, A. Verbeken 94-161 (BR 57614–93); Mugara, Feb. 1979, J. Rammeloo 6632 (BR 8778–48); Ruyigi Prov., Ruvubu National Park, April 1994, A. Verbeken 94-646 (BR 57714–96); Cankuzo Prov., forêt claire de Murangu, March 1994, B. Nzigidahera 56 (PC).
Democratic Republic of Congo. Shaba Prov., Biano, Lubudi, 18 March 1990, J. Degreef 90/192 (BR 5115–71); Shilatembo, 12 Dec. 1972, D. Thoen 5579 (BR 7769–09); road between Likasi and Lubumbashi, bought along road side, Jan. 1972, D. Thoen 5287 (Herb. Thoen); Ibid., 3 Dec. 1973, D. Thoen 5742 (BR 66367–19) & 5743 (BR 66368–20).
Malawi. Northern Prov., Mzuzu distr., Katoto, Feb. 1972, J. Williamson 608 (K(M) 38478); Ibid., March 1973, Pawek in J. Williamson 632 (K(M) 38471); Northern Prov., Mzimba distr., Perekezi Forest Reserve, 15 Feb. 2005, F. Noe 05/328 (GENT); Ibid., 11 March 2005, F. Noe 05/502 (GENT); Ibid., 16 March 2005, F. Noe 05/628 (GENT); Southern Prov., Zomba distr., near R.C. Cathedrale, 1 Jan. 1974, J. Williamson 736 (K(M) 38481).
Tanzania. Western Prov., Tabora distr., Tabora, purchased at market, 12 Dec. 1991, T. Saarimäki & al. 1057a (H); Southern Highlands Prov., Iringa distr., Mafinga, Sao Hill Misitu, 6 Feb. 1993, T. Saarimäki & al. 1588 (H).
Zambia. Western Prov., Mwunilinga, 29 Jan. 1991, B. Buyck 3514 (PC); Ibid., 30 Jan 1991, B. Buyck 3536 (PC); Mbereshi National Forest, 22 Feb. 1998, herb. M. Groenendaal L4; road Kawe-Kiposhi, 14 Jan 1996, B. Buyck & G. Eyssartier 96-005 (PC); road Mpongwe-Lusaka (Mikati), 18 Jan. 1996, B. Buyck & G. Eyssartier 96-118 (PC); Miputu, 22 Jan. 1996, B. Buyck & G. Eyssartier 96-005 (PC).
Zimbabwe. Midlands Prov., Mvuma, Mtao Forest Reserve, 28 Jan. 1999, A. Verbeken 99-041 (GENT); Manicaland Prov., Stapleford J. Meikle, 2 Feb. 1999, A. De Kesel 2385 (BR 112555–35).
Notes. The species is easily recognized by the firm context, the agreeable sweetish smell and taste and the reddish colour in dried condition. It is a popular edible species throughout the Zambezian miombo region (e.g. Buyck 1994, Härkonen & al. 2003, Parent & Thoen 1977).
24. Lactarius nodosicystidiosus Verbeken & Buyck
Buyck & al., Mycol. Res. 111: 794 (2007) − Type : Madagascar, Arivonimamo, 30 Jan 1997, B. Buyck, G. Eyssartier & P.A. Moreau 97-092 (PC, holotype; GENT, isotype).
Pileus 50–100 mm diam., convex to planoconvex, first with incurved margin, later with rather irregular, wavy margin; surface smooth, dry, slightly velvety and pruinose, cream to pale yellowish orange. Stipe 20–40 × 10–15 mm, cylindric, robust, solid; surface similar to pileipellis. Lamellae broadly adnate, distant, with lamellulae of different lenghts, cream; edge entire, concolorous. Context firm, pale.
Spores subglobose to broadly ellipsoid, rarely ellipsoid, 7.5–8.5–8.6–9.8 × 6.2–7.3–7.6–8.7 µm (Q = 1.03–1.13–1.18–1.39, n = 40; ornamentation amyloid, but very low and weakly developed, composed of indistinct, very low warts, sometimes aligned or connected by narrow lines, not exceeding 0.2 µm; plage inamyloid. Basidia 50–70 × 8–12 µm, 4-spored, rarely 2-spored, long, narrow, subcylindrical; sterigmata often long and sometimes swollen in the middle. Pleuromacrocystidia and pleuropseudocystidia very abundant and very similar. Pleuromacrocystidia slightly emergent, often originating deep in the hymenium, 75–110 × 5–9 µm, thin-walled, subcylindric, somewhat tapering near apex, often with very irregular apex, with bulges or moniliform; content opaque oil-like or needle-like. Pleuropseudocystidia slightly emergent, similar in shape to the pleuromacrocystidia, with irregular apex, often moniliform or with bulges and branching, up to 7(–12) µm broad; content oil-like or needle like. Lamellar edge sterile; marginal cells subcylindric to somewhat irregular or slightly subclavate, 20–45 × 6–9 µm, thin-walled, hyaline. Pileipellis a trichopalisade, composed of chains of small (usually less than 15 µm diam.), rounded cells, with long, narrow cylindric elements on top; terminal elements 30–60 × 3–5 µm, regularly cylindric, usually oblique to pericline, forming a rather densily interwoven layer of 30–60 µm thick on top of the sphaerocytes. Stipitipellis similar. Clamp-connections absent. – Fig. 48
Ecology. Found in Uapaca bojeri woodland.
Distribution. Only known from Madagascar.
Revised specimens
Madagascar. Arivonimamo, 30 Jan. 1997, B. Buyck & al. 97-072 (PC, holotype; GENT, isotype), B. Buyck & al. 97-045b (PC); Ibid., 5 Feb. 1997, B. Buyck & al. 97-273 (PC); Fiadanana village along RN7, 32 km from Antsirabe direction Ambositra, 10 Feb. 2000, B. Buyck 00-1336 (PC).
Notes. The microscopical description is based on Buyck 97-273 & 97-072.
Lactarius nodosicystidiosus is very similar to L. edulis. Both species share the pale overall colour, the robust, firm habit and the distant, broad lamellae. Under the microscope, however, the pileipellis structure of L. edulis shows a thicker, more loosely arranged pileipellis having the cylindrical terminal elements more dispersed and not forming a distinct superficial layer. Also the spore-ornamentation is more pronounced in L. edulis and spores are slightly smaller (on average 7.7 × 6.3 µm). Pseudocystidia as well as macrocystidia are strikingly abundant in L. nodosicystidiosus, while only pseudocystidia are present in L. edulis.
25. Lactarius latifolius Gooss.‑Font. & R. Heim
Heim, Bull. Jard. Bot. Etat Bxl 25: 82 (1955). – Type: Democratic Republic of Congo, Binga, Feb. 1935, M. Goossens-Fontana 1009 (BR 7726–63, holotype).
Lactarius cinnamomeus Gooss.‑Font. & R. Heim in Heim, Bull. Jard. Bot. Etat Bxl 25: 32 (1955), illegit., non Lactarius cinnamomeus W.F. Chiu (1945).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 32–35, 82–85 & Pl. 2, fig. 2a–d & Pl. 4, fig 3a–c (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 14, fig. 1 & Pl. 15, fig. 1 (1955); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 113–116 & Pl. 161–162 (1996d); Verbeken & Walleyn, Belg. J. Bot. 132: 177, 179–181 (2000).
Pileus 40–150 mm diam., firm, fleshy, convex to applanate, soon depressed, finally subinfundibuliform; margin first involute and curved, then very irregular and undulate, sometimes with cerebriform, sinuose and concentrical veins, striate to grooved when older; pellis not dehiscent, felty, finely cracking, orange-brown to reddish brown when young, cinnamon-brown, then rather polychrome, yellowish ochraceous, lemon-yellow in the centre, with violet-cinnamon shade in exsiccatum. Lamellae adnate to decurrent with teeth, unequal with a few lamellulae of different lengths, very distant (40/total), very thick, moderately broad, brittle, anastomosing with thick veins, orange, yellowish orange to pale brownish orange, with lilac shade in exsiccatum. Stipe 40 × 20–25 mm, short, thick, cylindrical, slightly tapering downwards, finely felty, brownish pale pink, cinnamon-colour or orange-brown, paler at the base, cream and faded lilac in exsiccatum, firm, solid. Context brittle, white, sometimes becoming yellowish; taste variable; smell not particular. Latex abundant, white; taste variable. Spore-deposit whitish, cream. (According to Heim 1955a). – Pl. 14
Chemical reactions. Context immediately blue (Goossens-Fontana 3062) or nihil (Goossens-Fontana 2074 & 3025) with aniline; immediately pale pink with FeSO4 (Goossens-Fontana 2074, 3025 & 3062); slowly blue-green (Goossens-Fontana 2074, 3025 & 3062) or immediately blue-green (Goossens-Fontana 1009) with gaiac; unchanging with naphthol (Goossens-Fontana 2074) and NH4OH (Goossens-Fontana 2074, 3025 & 3062); wine-colour to blackish with phenol (Goossens-Fontana 2074 & 3062); immediately brown (Goossens-Fontana 2074 & 3062) or slowly yellowish brown (Goossens-Fontana 1009 & 3025) with pyrogallol. Lamellae immediately brown with FeSO4 (Goossens-Fontana 1009). (According to Heim 1955a).
Basidiospores subglobose to ellipsoid, 6.2–6.5–7.9–8.8(–9.2) × 5.0–5.7–6.9–7.6(–8.0) µm (Q = 1.03–1.14–1.19–1.28, n = 80); ornamentation amyloid, low, composed of rounded to irregularly shaped warts, connected by fine connective lines, forming an almost complete reticulum, < 0.3 µm high; plage inamyloid. Basidia (35–)50–67 × (5–)8–12 µm, cylindrical to subcylindrical, long and slender, 4-spored; content granular. Pleurocystidia absent. Pleuropseudocystidia abundant, slightly emergent, 7–12 µm diam., cylindrical or fusiform, with tapering or rounded apex; content oleiferic. Lamellar edge sterile; marginal cells 16–45 × 4–5 µm, cylindrical to tortuous, with rounded or tapering, sometimes mucronate apex, thin-walled, hyaline. Hymenophoral trama composed of sphaerocytes; some hyaline hyphae and lactifers present. Pileipellis a trichopalisade, composed of chains of isodiametric to irregular cells, often with incrusting pigments; isodiametric cells 20–30 µm diam.; terminal elements (20–)30–50 × 5–6 µm, cylindrical, slender, with rounded apex, sometimes fusiform and up to 12 µm diam., often with small, intracellular, dark granules (not indicated in figure). Stipitipellis cutis-like, composed of hyaline hyphae; hyphae branched, septate, more or less parallel; lactifers present; hyphae at the base of the stipe with slightly thickened walls. Clamp-connections absent. – Fig. 49
Ecology. Drier Guineo‑Congolian rainforest.
Distribution. Known from the Democratic Republic of Congo and Gabon.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Binga, ca. 1928, M. Goossens-Fontana 866 (BR 4968–21); Ibid., Feb. 1935, M. Goossens-Fontana 1009 (BR 7726–63, holotype, consisting of 5 basidiomes in very poor condition); Ibid., Nov. 1945, M. Goossens-Fontana 2074 (BR 5007–60); Ibid., Jan. 1943, M. Goossens-Fontana 3025 (BR 7730–67); Ibid., Nov. 1946, M. Goossens-Fontana 3062 (BR 7731–68).
Gabon. Research Station of Ipassa-Makokou, 24 March 2005, S. Dibaluka Mpulusu 37 (BR 164536–24).
Notes. Heim (1955b) already mentioned that Lactarius cinnamomeus Gooss.-Font. & R. Heim has identical microscopic characters as L. latifolius, but indicated the absence of gelified veins in L. latifolius and differences in taste of the context (acrid in L. cinnamomeus) as discriminating characters. As for the taste, we consider this character as very doubtful, as in fact the fieldnotes accompanying Goossens-Fontana 3062 mention a mild taste. As to the gelified veins on the pileus, they are probably only present in young basidiomes and the exsiccata of the syntypes of Lactarius cinnamomeus show smooth and very regular pileal surfaces, just like those of L. latifolius.
26. Lactarius aureifolius Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 198 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 11 March 1994, A. Verbeken 94-005 (BR 57557–36, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 164–165 & Pl. 125–128 (1996d); Verbeken & Walleyn, Belg. J. Bot. 132: 177–179 (2000); Verbeken, Micol. Veget. Medit. 16: fig. 2 (2001).
Pileus 30–120 mm diam., thick, fleshy, firm, convex when young, then planoconvex to applanate, finally depressed to infundibuliform; margin involute when young, then straight or slightly deflexed upwards, regular or crenulate; pellis dehiscent up to halfway the pileus, dry, mat, finely felty, cracking when older, pale yellow (4A3–4) when young, pale orange to greyish orange (5A3 to 5B4), brownish in the centre when older, sometimes with pinkish shade. Lamellae adnate to slightly decurrent, unequal with lamellulae of two different lengths (regular pattern, 3(–5) lamellulae between two lamellae), distant (L+l = 2+5 to 3+8/cm, exceptionally 5+15/cm), broad (3–10 mm), rather thick, firm then brittle, sometimes anastomosing and dichotomously branched, pale yellow to deep yellow, orange yellow (4A5–8); edge entire, brighter than lamella when young, more orange when old. Stipe (15–)25–55 × 15–35 mm, short cylindrical, sometimes tapering downwards, dry, mat, finely cracking, pale yellow to pale orange, somewhat paler than pileus, whitish downy at the base, solid. Context firm, 8–10 mm in pileus, white to cream, changing locally pale orange when cut; smell agreeable, sweetish, rhubarb-like, apple-like; taste very acrid, burning. Latex abundant to very abundant, fluid, white, unchanging; taste very acrid, burning. Spore-deposit white. – Pl. 15
Chemical reactions. Context salmon pink to orange to greyish with FeSO4; unchanging with HCl; unchanging or yellow with KOH. Pileipellis olive brown with KOH, unchanging or somewhat yellow with NH4OH.
Basidiospores ellipsoid, 6.7–7.6–7.8–8.8 × 5.7–6.2–6.5–7.2 µm (Q = 1.12–1.21–1.22–1.33, n = 60); ornamentation amyloid, composed of low, irregularly sized and shaped warts, often aligned or connected by fine lines, never forming a complete reticulum, < 0.2 µm; plage with central amyloid spot. Basidia 50–80 × 6–8 µm, cylindrical to subclavate, long and slender, 4-spored; content sometimes guttate. Pleurocystidia absent. Pleuropseudocystidia rather abundant, 4–9 µm diam., cylindrical to clavate, with rounded apex. Lamellar edge sterile; marginal cells 15–25 × 3–6 µm, cylindrical to subfusiform, hyaline, thin-walled. Hymenophoral trama composed of small sphaerocytes and abundant lactifers, a few hyaline hyphae present. Pileipellis a trichoderm, composed of interwoven hyphae; hyphae 4–6 µm diam., long, slender, often septate, sometimes branched, very slightly thick-walled; terminal elements not aberrant, 15–30 × 4–6 µm, cylindrical, rounded at the apex. Stipitipellis idem; some intracellular granules observed. Clamp-connections absent. – Fig. 50
Ecology. Zambezian miombo woodland with dominance of Brachystegia spp.
Distribution. Known from Burundi, Democratic Republic of Congo, Malawi, Zambia and Zimbabwe. Not common.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, March 1994, A. Verbeken 94-005 (BR 57557–36, holotype; GENT, isotype), 94-032 (BR 57572–51), 94-076 (BR 57586–65), 94-273 (BR 57630–12), 94-350 (BR 57649–31) & 94-536 (BR 57695–77); Ibid., April 1994, A. Verbeken 94-556 (BR 57698–80).
Democratic Republic of Congo. Shaba Prov., Kipopo, 9 Jan. 1961, M.C. Schmitz-Levecq 293 (BR 12294–72).
Malawi. Southern Prov., Machinga distr., Liwonde Forest Reserve, 4 Feb. 2005, A. Verbeken 05-207 & 05-209 (all GENT).
Zambia. Farm Gibsons, 23 Jan. 1996, B. Buyck & G. Eyssartier 96-157 (PC).
Zimbabwe. Manicaland Prov., Penhalonga, hill behind Harris’, 9 Feb. 1999, A. Verbeken 99-153 (GENT); Manicaland Prov., Chipinge, Busi farm, around Scott’s house, 12 Feb. 1999, A. Verbeken 99-199 (GENT).
Notes. This species is easily recognized in the field by its deep yellow to golden yellow lamellae and the very acrid to burning taste of the latex. These outstanding differences not taken in account, it could be confused with Lactarius edulis, from which it is microscopically also very different by the lack of isodiametric or inflated elements in the pileipellis.
27. Lactarius densifolius Verbeken & Karhula
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 200 (1996). – Type: Zambia, Mwunilunga, 30 Jan. 1991, B. Buyck 3601 (BR 38487–75, holotype; PC, isotype).
Misappl.: Lactarius inversus ss. Buyck (1994) & Buyck & Nzigidahera (1996); L. piperatus ss. Pegler & Piearce (1980).
Excl.: L. densifolius ss. Van Rooij & al. (2003), De Kesel & al. (2002), = Lactarius sp.
Selected descriptions and icons. Buyck, Champignons comestibles de l'Ouest du Burundi: 105, fig. 79 (1994: as "inversus"); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 165–167 & Pl. 129–133 (1996d) ; Verbeken & Walleyn, Belg. J. Bot. 132: 177, 179–180 (2000); Karhula & al., Karstenia 38: fig. 21 (1998); Härkonen & al., Norrlinia 10: 84 (2003).
Pileus 30–70 mm diam., firm, fleshy, planoconvex to applanate, then deeply depressed to infundibuliform; margin involute, sometimes becoming reflexed when older; pellis not dehiscent, dry, finely tomentose, typically rupturing into small, felty fields, pale brownish yellow or apricot orange to cream. Lamellae decurrent, unequal with lamellulae of different lengths, very dense, very narrow, cream; edge entire, concolorous. Stipe 40–60 × 12–25 mm, cylindrical, concolorous with the pileus, firm, solid. Context firm, white, unchanging; taste mild. Latex white to transparent, unchanging; taste mild. Spore-deposit white.
Basidiospores subglobose to ellipsoid, (7.5–)8.0–8.7–9.4(–9.7) × (6.5–)6.8–7.6–8.5(–8.7) µm (Q = 1.04–1.14–1.25, n = 40); ornamentation amyloid, composed of small and irregularly shaped warts, aligned or connected by fine lines, not exceeding 0.2 µm height; plage inamyloid. Basidia 55–70 × 9–11 µm, subcylindrical to subclavate, long and slender, 4-spored, sometimes 2-spored; content guttate. Pleurocystidia rather abundant, arising deep in the subhymenium, sometimes slightly emergent, (80–)90–110(–120) × 4–6 µm, long and slender, cylindrical, almost moniliform at the apex, sometimes branched; content hyaline or oleiferic. Pleuropseudocystidia rather abundant, slightly emergent, 4–6 µm, slender, cylindrical or slightly tortuous, almost moniliform at the apex, sometimes branched; content oleiferic. Lamellar edge sterile; marginal cells 15–50 × 4–7 µm, cylindrical to subclavate, with rounded apex, thin-walled, hyaline. Hymenophoral trama composed of sphaerocytes; sphaerocytes especially abundant in the central part, also mixed with thin-walled, hyaline hyphae which are more abundant near the subhymenium; lactifers present. Pileipellis a trichoderm; elements of the suprapellis 20–80 × 3–5 µm, long and slender, cylindrical, frequently septate, with rounded apex, sometimes with incrustrated wall, sometimes with small bulges, thin-walled, hyaline, but sometimes guttate; subpellis composed of hyphae, with some isodiametric or irregularly shaped cells. Stipitipellis trichoderm-like; elements of the suprapellis 20–50 × 3–5 µm, cylindrical, slender, with rounded apex, frequently septate, sometimes branched, with slightly thickened wall, becoming much more longer and more thick-walled near the base of the stipe; subpellis composed of parallel hyphae and lactifers, no isodiametric cells. Clamp-connections absent. – Fig. 51
Ecology. Zambezian miombo woodland.
Distribution. Widespread in the Zambezian region. Known from Burundi, Democratic Republic of Congo, Malawi, Tanzania and Zambia.
Revised specimens
Burundi. Cankuzo Prov., forêt claire de Murangu, Dec. 1994, B. Buyck s.n. (PC).
Democratic Republic of Congo. Shaba Prov., Kipopo, 14 March 1959, M.C. Schmitz‑Levecq 105 (BR 6123–12).
Malawi. Southern Prov., Makwawa, Zomba, 29 Jan. 1980, B. Morris 82 (K(M) 38468); Southern Prov., Machinga distr., near Machinga, Malosa Forest Reserve, 25 Jan. 2005, A. Verbeken 05-062 (GENT); Northern Prov., Mzimba distr., Perekezi Forest Reserve, 9 March 2005, F. Noe 05/455 (GENT); Ibid., 11 March 2005, F. Noe 05/504, 05/510 & 05/514 (all GENT).
Tanzania. Southern Prov., Songea distr., Matomondo, 29 km from Songea towards Mbinga, 29 Jan. 1993, T. Saarimäki & al. 1449 (H); Ibid., Lirondo, 30 Jan. 1993, T. Saarimäki & al. 1468 (H); Southern Highlands Prov., Kidugala, 30 Jan. 1993, T. Saarimäki & al. 1474 (H); Ibid., Njombe distr., Ngaramiro, 2 km N of Kidugala, 31 Jan. 1993, T. Saarimäki & al. 1479 & 1489 (all H); Ibid., between Sengele and Masaulwa, N of Kidugala, 2 Feb. 1993, T. Saarimäki & al. 1524 & 1543 (all H).
Zambia. Western Prov., Mwunilunga, 30 Jan. 1991, B. Buyck 3601 (BR 38487–75, holotype; PC, isotype); Luapala Prov., Mansa, Kwanfumwe, 2 Jan. 1991, B. Buyck 3311 (PC); Copperbelt Prov., Kitwe, 21 Dec. 1977, G. Piearce FP600/1a (K(M) 34213); Miputu, 22 Jan. 1996, B. Buyck & G. Eyssartier 96-150 (PC).
Notes. This species is edible, and well appreciated in the Zambezian miombo area (e.g. Buyck 1994, Härkonen & al. 2003, Pegler & Piearce 1980).
The species was recorded for Benin (Van Rooij & al. 2003, De Kesel & al. 2002) but the concerning specimen (Borgou Prov., Wari Maro, 24 Sept 1998, De Kesel 2324) represents another species. The spore ornamentation consists of larger and more regular warts with less connections.
28. Lactarius inversus Gooss.‑Font. & R. Heim
Heim, Bull. Jard. Bot. Etat Bxl 25: 78 (1955). – Type: Democratic Republic of Congo, Binga, Nov. 1945, M. Goossens-Fontana 3063 (BR 7732–69, holotype).
Excl.: Lactarius inversus ss. Buyck (1994, = Lactarius densifolius).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 78–82 & Pl. 3, fig. 2 (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 15, fig. 2 (1955); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 163–164 & Pl. 120–124 (1996d).
Pileus 30–100 mm diam., convex, becoming more and more depressed, finally infundibuliform; margin incurved, then straight and irregular; pellis not dehiscent, slightly velvety, tomentose, cracking into small, felty, darker flocks, sienna-colour, brownish to yellowish orange, becoming more orange when older, more reddish in exsiccatum; veins towards the margin slender, concolorous, concentrical and sinuose, later varicose and radial. Lamellae decurrent with long teeth, unequal with lamellulae of two different lengths, veined to anastomosing, crowded, narrow (< 4 mm), pale yellow to yellowish cream; edge entire, concolorous. Stipe (30–)40–65 × 10–15 mm, firm, long cylindrical, rounded at the base, felty, with longitudinal veins in the upper third, brownish orange as pileus, more yellowish orange when young, cinnamon with some red areas when old. Context firm, brittle, white, becoming slightly yellowish; taste mild; smell acrid. Latex abundant, white, transparent, unchanging; taste mild, sweet. Spore-deposit not observed. – Pl. 16
Chemical reactions. Context unchanging with aniline; slowly blue-green with gaiac; slowly pinkish with Fe SO4; brown with pyrogallol; immediately wine-colour with phenol; unchanging with NH3. (according to Heim 1955a)
Basidiospores subglobose to ellipsoid, 6.3–7.3–8.0–8.9 × 5.3–6.1–6.9–7.6 mm (Q = 1.06–1.17–1.20–1.35, n = 50); ornamentation amyloid, composed of globose to irregular warts, often isolated, sometimes connected by fine connective lines, < 0.2 mm; plage sometimes with central amyloid spot. Basidia 40–55 × 7–10 mm, cylindrical to subclavate, 4-spored, often 2-spored and then sterigmata up to 17 mm. Pleurocystidia scarce, 50–65 × 6–8 mm, cylindrical to subclavate with remarkable mucronate to rostrate apex, hyaline, thin-walled, giving the impression to be deformed, monosterigmatic basidia. Pleuropseudocystidia moderately abundant, 4–8 mm diam., cylindrical to tortuose, hardly emergent; content oleiferic. Lamellar edge mixed; basidia present, 30–40 × 7–9 mm; marginal cells 10–40 × 5–10 mm, cylindrical to clavate, hyaline, thin-walled. Hymenophoral trama composed of sphaerocytes and lactifers. Pileipellis a trichopalisade, composed of isodiametric to pyriform or irregular cells which are 30–40 mm diam., with slightly thickened and often incrustated walls, mostly forming chains; terminal cells 15–35 × 5–10 mm, cylindrical to pyriform, with slightly thickened and often incrustated walls. Stipitipellis idem as pileipellis. Clamp-connections absent. – Fig. 52
Ecology. Drier Guineo‑Congolian rainforest, lowland humid rainforest.
Distribution. Known from Cameroon, Democratic Republic of Congo and Guinea.
Revised specimens
Cameroon. South Western Prov., near Mundema, Korup National Park, transect P10, 27 March 1991, R. Watling 24174 (E).
Democratic Republic of Congo. Equateur Prov., Binga, terrestrial, Nov. 1945, M. Goossens-Fontana 3063 (BR 7732–69, holotype).
Guinea. Forêt humide de Ziama, arboretum des stagiaires, 1 Aug. 1999, A. Bâ 168 (GENT).
Notes. The type consists of 4 basidiomes in rather good condition.
The macroscopic description is based on the description of Heim (1955a), completed with fieldnotes of Watling. The microscopic description is based on both specimens.
The species is recognized in the section by the dense lamellae, a character shared with Lactarius densifolius, but differs from the latter by the reddish colours of the pileus and the pileipellis which is here a trichopalisade whereas a trichoderm in L. densifolius.
29. Lactarius phlebophyllus R. Heim
Heim, Candollea 7: 378 (1938). – Type: Madagascar, Anosibé, S of Soanierana, 8 Dec. 1934, R. Heim G87 (PC, holotype).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 52–53 (1938); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 162–163 & Pl. 117–119 (1996d); Buyck & al., Mycol. Res. 111: 792–794 (2007).
Pileus 30–100 mm diam., thin, convex to planoconcave and slightly depressed in the centre, then depressed, irregular; margin translucid, striate by the lamellae; pellis tomentose, spotted, cracking into small areolate fields in mature specimens, smooth in the centre, dry, ochraceous orange, brownish orange, subolivaceous, very pruinose and pale especially in young specimens, in older specimens more brownish orange in the centre; margin somewhat brighter and golden yellow in young specimens, becoming ochraceous and irregularly undulate in older specimens. Lamellae broadly adnate to decurrent with small tooth, rather crowded to distant, brittle, narrow (up to 3 mm), thin, anastomosing with abundant lateral veins, dichotomously branched near the margin or often forked at different distances, cream, creamish orange; edge entire, concolorous or slightly paler. Stipe 35–50 × 10–20 mm, robust, tapering downwards, sometimes irregular and compressed, finely tomentose, orange, slightly ochraceous, pale ochraceous and somewhat more yellowish than the pileus, whitish at the apex, solid. Context firm, cream, unchanging; smell not particular; taste mild. Latex white, rather abundant in young specimens, unchanging; taste mild. Spore-deposit white. – Pl. 16 & 17
Basidiospores subglobose to ellipsoid, seldom globose, 6.4–7.7–7.9–9.0 × 5.5–6.4–7.0–7.6 mm (Q = 1.01–1.13–1.22–1.40, n = 50); ornamentation amyloid, very low, up to 0.3(–0.5) µm high, composed of irregular warts which are isolated or sometimes aligned or connected by fine lines, forming an incomplete reticulum; plage inamyloid. Basidia 40–60 × 7–12 mm, cylindrical to subclavate, 4-spored, with rather broad sterigmata which are 5–8 µm long, slightly thick-walled at the base. Pleurocystidia absent. Pleuropseudocystidia not very abundant, not or hardly emergent, subcylindrical to irregularly tortuose and with often very tortuous, branching apex, sometimes with some bulges, 5–10(–12) µm diam.; content densely oleiferic, slightly guttate. Lamellar edge sterile; marginal cells 10–75 × 4–8 mm, narrowly cylindrical, subclavate, thin-walled, hyaline. Hymenophoral trama composed of rather small sphaerocytes; lactifers scarce. Pileipellis a trichopalisade, up to 150 µm thick, composed of isodiametric to elongate cells, 10–45 × 7–20 mm, mostly forming chains; terminal elements 15–60 × 5–15 mm, cylindrical or fusiform, tapering upwards. Stipitipellis similar to pileipellis structure. Clamp-connections absent. – Fig. 53
Distribution. Only known from Madagascar.
Ecology. Described from Malagasy lowland rainforest, and found recently in dry dense forest with dominance of Uapaca densifolia, Sarcolaena multiflora and Leptolaena pauciflora.
Revised specimens
Madagascar. Betsimisaraka Prov., Anosibé, S of Soanierana, terrestrial, Dec. 1934, R. Heim G87 (PC, holotype); Ambohitantely Natural Reserve, N of Ankazobe, 15 Feb. 2000, B. Buyck 00-1388 & 00-1400 (all PC).
Notes. The type of L. phlebophyllus, consists of half a basidiome in bad condition. This description is based on the type material and on the recently collected specimens from Madagascar (Buyck & al. 2007).
Lactarius phlebophyllus is a striking species because of the overall uniform, yellowish colour, the areolate pileus and the very narrow, almost regularly forked gills. Heim (1938b) described the gills as ‘distant’, but our own observations on the type contradicted this and showed both collections to be exactly comparable in this respect.
Spore characters and pileipellis structure are strictly comparable, but there are some minor microscopical differences between our observations made on the type specimen and those on the recently collected material. Pleuropseudocystidia are present in both, but more scarce in the type material. The narrow and very long marginal cells of the gill edge in the recent collection have not been observed on the type.
30. Lactarius roseolus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 207 (1996). – Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 23 March 1994, A. Verbeken 94-274 (BR 57631–13, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 222–223 & Pl. 252–254 (1996d); Verbeken & al., Syst. Geogr. Pl. 70: 203–205 (2000); Verbeken, Micol. Veget. Medit. 16: fig. 4 (2001).
Pileus (5–)10–31 mm diam., very thin, very delicate and brittle, planoconvex to applanate and slightly infundibuliform, weakly depressed, sometimes with a very small central umbo inside that depression; margin regular, finely but distinctly crenulate, faintly striate, grooved in older specimens; pellis not dehiscent, soft, dry, chamois-leather-like to felty, sometimes slightly rugulose in older specimens, pale orange to pastel red (6A3 to 7A4 to 8A5), dark reddish brown (7E–F8), brownish orange (6D6), paler orange when older, paler and almost white towards the margin. Lamellae adnate to slightly or strongly decurrent, unequal with lamellulae of different lengths (0–3 lamellulae between 2 lamellae), distant (total number L = 32–40), narrow (1–2 mm), fairly thick, white to cream (3A2 to 4A2), with some furcations; edge entire, concolorous. Stipe 10–30 × 1–6 mm, a bit irregularly cylindric, slender, often tapering downwards; pellis smooth or more often finely cracked or a bit sqamulose which makes it looks spotted on a whitish background, concolorous with pileus or a bit paler, pastel red to greyish red (7A4–5), whitish at the base, very contrasting with the white lamellae and sometimes with a distinct separation in colour just underneath the sometimes decurrent teeth. Context thin in pileus, fistulose or multi-chambered in stipe, fragile, white, unchanging; taste fresh, mild, slightly menthol-like; smell not particular. Latex quite abundant, white or watery, unchanging; taste mild. Spore-deposit white. – Pl. 18
Chemical reactions. Context salmon with FeSO4, unchanging with gaiac.
Basidiospores subglobose to ellipsoid, 7.0–7.8–7.9–8.6 × 6.0–6.9–7.0–7.6(–7.9) mm (Q = 1.05–1.11–1.13–1.25; n = 60); ornamentation not very distinct, very low and irregular, composed of irregularly shaped and sized warts and connective lines; plage inamyloid. Basidia 50–70 × 7–10 mm, cylindrical to subcylindrical, long and slender, 4-spored. Pleurocystidia absent. Pleuropseudocystidia scarce, 3–5 mm diam., cylindrical to tortuous, sometimes branched; content oleiferic. Lamellar edge fertile, no marginal cells observed. Hymenophoral trama heteromerous, composed of sphaerocytes; lactifers fairly abundant. Pileipellis a palisade, hymeniderm-like; elements of suprapellis 20–60 × 5–10 mm, cylindrical to fusiform, sometimes septate, hyaline, thin-walled; subpellis pseudoparenchymatous, composed of isodiametric cells. Stipitipellis a trichoderm; terminal elements 10–50 × 4–10 mm, cylindrical to fusiform, sometimes septate, hyaline, thin-walled, connected with slender hyphae which are forming an underlaying layer. Clamp-connections absent. – Fig. 54
Ecology. Miombo woodland with Brachystegia spp.
Distribution. Only known from Burundi and Zimbabwe.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, 11 March 1994, A. Verbeken 94-064 (BR 57575–54); Ibid., 23 March 1994, A. Verbeken 94-274 (BR 57631–13, holotype; GENT, isotype); Ibid., 25 March 1994, A. Verbeken 94-343 (BR 57644–26); Road Kigwena to Vianda, 6 May 1982, C. Cocquyt 336 (GENT).
Zimbabwe. Manicaland Prov., Chipinge, Busi farm, around Scott’s house, 12 Feb. 1999, A. Verbeken 99-207 & 99-208 (all GENT); Manicaland Prov., Burma valley road, 20 km from turnoff “cloudland”, 10 Feb. 1999, A. Verbeken 99-159 & 99-160 (all GENT); Manicaland Prov., above Bridal Veil Road, Chimanimani, 26 Jan. 1999, D. Arora 99-50 (SFSU); Manicaland Prov., Eastern Highlands, Mguzu, 5 Feb. 1999, Gansemans in A. De Kesel 2740 (BR 115760–39); Ibid., 6 Feb. 1999, A. De Kesel 2421 (BR 112591–71).
Notes. The collection Verbeken 99-159 from Zimbabwe consisted of stouter, darker coloured and more reddish brown (the colour of the red banded ironstone) specimens than the small pale pinkish specimens from Burundi, that were mostly collected as single specimens. Because the Burundi and Zimbabwe specimens are microscopically fully identical, we do not believe that these differences have taxonomic value.
Specimens of one collection, Arora 99-50, had a very strong smell of grated coconut (macaroons), which was absent in the other collections. Conspicuous differences in smell were also observed in other African Lactarius species such as L. saponaceus.
Lactarius sect. Aurantiifolii Verbeken, Mycotaxon 77: 441 (2001).
− Type species: Lactarius aurantiifolius Verbeken.
31. Lactarius aurantiifolius Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 197 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 11 March 1994, A. Verbeken 94-063 (BR 57574–53, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 167–168 & Pl. 134–138 (1996d); Verbeken & al., Syst. Geogr. Pl. 70: 185–187 (2000); Verbeken, Micol. Veget. Medit. 16: fig. 1 (2001).
Pileus 17–50 mm diam., thin, convex to planoconvex and applanate, finely infundibuliform, slightly depressed; margin incurved when young, then straight and regular; pellis not dehiscent, dry, very pruinose when young, felty (with distinct erect hairs in young specimens, lens!) to smooth when older, concentrically zoned (more distinct in center, often vague towards margin), orange white to pale orange (5A2–7), orange in the centre (6AB6), whitish pruinose (5A2–3) when young. Lamellae adnate to slightly decurrent, unequal with 1–3 lamellulae between 2 lamellae (regular pattern), crowded to moderately distant (L+l = 3+7 to 5+13/cm), with abundant lamellulae from different lenghts, thin, narrow (2–3 mm), paper-like, veined when older, sometimes anastomosing, deep orange (5A6–7); edge paler, whitish, appendiculate to fimbriate, with watery tears. Stipe 10–70 × 4–16 mm, cylindrical, strongly tapering downwards, knotty, dry, felty, orange white to pale orange (4A4 to 5A2–4), whitish pruinose (remaining so in the exsiccatum), fragile, chambered (with up to 8 chambers). Context brittle, white to cream, pale orange near the pellis, unchanging; taste mild, pleasant; smell pleasant to fruity. Latex not abundant, white, unchanging; taste mild. Spore-deposit white. – Pl. 18
Chemical reactions. Context pale salmon pink with FeSO4, changing yellow then olive brown with KOH, yellow with NH4OH, unchanging with HCl and gaiac.
Basidiospores ellipsoid, (7.4–)7.5–8.0–8.4–9.6 × 5.9–6.0–6.5–7.3 mm (Q = 1.17–1.23–1.27–1.33; n = 40); ornamentation faintly amyloid, composed of small warts and low ridges forming an obscure broken reticulum; prominences extremely low and hardly distinct; plage inamyloid. Basidia 45–65(–70) × 8–10(–11) mm, cylindrical to slightly clavate, 4-spored, sometimes 2-spored. Pleurocystidia 60–70 × 12–15 mm, clavate, slightly thick-walled. Pleuropseudocystidia not abundant, 3–5 mm, cylindrical, obtuse. Lamellar edge sterile; marginal cells 20–80(–150) × 5–8 mm, cylindrical, obtuse, septate, sometimes branched. Hymenophoral trama composed of small sphaerocytes and hyaline narrow hyphae; laticiferous hyphae not abundant. Pileipellis a trichoderm, composed of interwoven and ascending hyphae, up to 150 mm, hyaline, 4–7 mm diam., with rounded apex, septate, thin-walled. Stipitipellis as pileipellis but less thick (up to 70 mm thick). Clamp-connections absent, although in the stipitipellis some clamp-connections-like structures were observed; they are not more frequent towards the base but best observed halfway the stipe. – Fig. 55
Ecology. Found in Zambezian miombo woodland dominated by Brachystegia utilis, Sudanian rivulet associated woodland with dominance of Uapaca somon, Uapaca bojeri woodland.
Distribution. Widespread in tropical African woodlands. Known from Benin, Burundi, Madagascar and Zimbabwe.
Revised specimens
Benin. Atacora Prov., Kota, 23 Sept. 2000, A. De Kesel 2928 (BR 129139–32).
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, 18 May 1993, B. Buyck 5175 (PC); Ibid., 11 Feb. 1994, B. Buyck 5470 (PC); Ibid., 11 March 1994, A. Verbeken 94-063 (BR 57574–53, holotype; GENT, isotype); Ibid., 31 March 1994, A. Verbeken 94-513 (BR 57683–65); Nkayamba, just N of Rumonge, 29 Jan. 1992, B. Buyck 4180 (PC); Ibid., 29 April 1992, B. Buyck 4398 (PC).
Madagascar. Arivonimamo, 30 Jan. 1997, B. Buyck & al. 97-060 (PC).
Zimbabwe. Manicaland Prov., Chipinge, Busi farm, forest around Scott’s house, 12 Feb. 1999, A. Verbeken 99-186 (GENT); Manicaland Prov., Penhalonga, hill behind Harris’ garden, 9 Feb. 1999, A. Verbeken 99-152 (GENT).
Notes. This species is characterized by a lot of outstanding and unique characters, not observed in any other African Lactarius species: the bright orange lamellae, the white and fimbriate lamellar edge, the zoned pileus, the very pruinose pileus and the chambered, tapering stipe.
Sect. Rubroviolascentini (Singer) Verbeken
Lactarius sect. Rubroviolascentini Verbeken, Mycotaxon 66: 380 (1998). Lactarius subsect. Rubroviolascentini Singer, Ann. Mycol. 40: 114 (1942).
− Type species: Lactarius rubroviolascens R. Heim.
Notes. The distinction between this section and L. sect. Pseudogymnocarpi is only based on blackening context. This grouping is artificial but maintained for practical reasons.
Key to tropical African species
1. Pileus brownish ochre to orange reddish; context becoming blackish, then reddish; spores extremely lowly ornamented to almost smooth; plage not amyloid .......... 32. L. rubroviolascens
Pileus pale buff; context becoming red, then lilac and finally strongly blackening by the latex; spores with a distinct incomplete reticulum, plage centrally amyloid ............. 33. L. denigricans
32. Lactarius rubroviolascens R. Heim
Heim, Candollea 7: 377 (1938). – Type: Madagascar, South of Soanierana, near Manompana, 9 Dec. 1934, R. Heim G85 (PC, holotype).
Selected descriptions and icons. R. Heim, Prodr. Fl. Mycol. Madagascar 1: 46–49 & Pl. 1c (1938); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 138–139 & Pl. 64–68 (1996d).
Pileus 60–80 mm diam., firm, fleshy, first convex and with slight central depression, then planoconvex to irregular; margin inflexed, irregularly crenulate; pellis dehiscent, thick, tomentose (handlens), smooth or with local small excavations, brownish ochre with orange tinge, pale pinkish, greyish pink, slightly hygrophaneous, greyish tinges more obvious when dry, more orange-reddish when wet, sometimes with some blackish spots, often with local whitish pruinose aspect. Lamellae broadly adnate to subdecurrent, unequal with lamellulae of two different lengths, very distant (L+l = 6 to 11/cm), large (7–8 mm), very thick, brittle or papery, cream to slightly greyish yellow (3A2), in some specimens at first blackening, finally violet-blue; edge concolorous or distinctly pale orange (especially in young specimens). Stipe 30–45 × 18–28 mm, cylindrical, slightly tapering downwards, dry, velvety, orange to yellow with pinkish tinge (5A5–6), orange-like near the apex, pale in the middle, brownish orange in the lower part, white and tomentose at the base, sometimes greyish or pinkish at base, solid, firm. Context firm, white, pale orange under the stipitipellis, becoming blackish then reddish in the pileus, or first pale grey, then pink-reddish and finally greyish black, in some specimens unchanging, unchanging in the stipe; taste mild; smell not particular. Latex abundant, water-like, almost translucid, or first white, then heterogenous watery with white clouds, with greyish pink tinge, unchanging or reddening, drying greyish black; taste mild. Spore-deposit white. – Pl. 19
Chemical reactions. Context unchanging with gaiac and gaiacol; pink with FeSO4; pale lilac with pyramidon. Pileipellis green with FeSO4. (According to Heim 1938b).
Basidiospores ellipsoid, (7.7–)8.2–8.8–9.4–10.3 × 6.3–6.9–7.4–7.9 mm (Q = 1.20–1.27–1.28–1.40; n = 50); ornamentation not very distinct, very low and irregular; composed of some irregularly shaped and sized warts and connective lines; plage inamyloid. Basidia 40–60 × 8–11 mm, cylindrical to slightly flexuous or slightly subclavate, 4-spored; sterigmata 5–7 × 1–2(–3) mm. Pleurolamprocystidia very abundant, 110–140 × 10–15 mm, very emergent (up to 70 mm), cylindrical to setiform, obtuse or tapering upwards; wall thickened (1–3(–4) mm); content sometimes granular. Pleuropseudocystidia scarce, (2–)4–6 mm diam.; cylindrical, obtuse, sometimes mucronate, sometimes branched; content oleiferic. Lamellar edge without distinct cells, with some lamprocystidia and a lot of dark brown, amorphous flocks. Hymenophoral trama irregular and heteromerous, composed of delicate, hyaline hyphae (3–6 mm diam.) and numerous sphaerocytes; lactifers abundant. Pileipellis a palisade to trichopalisade; suprapellis 50–90 mm thick; subpellis 20–30 mm thick; elements of suprapellis 20–50(–90) × 4–6 mm, cylindrical and obtuse to tortuose, septate and branched, with slightly thickened wall and slightly granular content; subpellis pseudoparenchymatous, with isodiametric or irregular cells. Stipitipellis a lamprotrichoderm to lamprotrichopalisade, 100–150 mm thick; elements of the suprapellis resembling the lampropleurocystidia, (50–)100–135 × 4–8 mm, cylindrical to setiform, slightly flexuous, obtuse; wall thickened (1–3 mm); content granular; elements at the base of stipe with remarkable globose and swollen apex; subpellis pseudoparenchymatous, rudimentary; isodiametric cells scarce. Clamp-connections absent. – Fig. 56
Ecology. Found in various forest types, such as lowland rainforest, miombo woodland, Uapaca bojeri woodland, dry evergreen forest (muhulu), lowland rainforest. Not common.
Distribution. Known from Cameroon, Democratic Republic of Congo, Madagascar, Malawi and Zambia.
Revised specimens
Cameroon. Western Prov., near Mundeba, Korup National Park, transect P, 1984, I. Alexander 13 (E).
Democratic Republic of Congo. Shaba Prov., Kipopo, March 1971, D. Thoen 4635 (BR 66235–81).
Madagascar. Betsimisaraka Prov., South of Soanierana, Dec. 1934, R. Heim G85 (PC, holotype); Arivonimamo, 5 Feb. 1997, B. Buyck & al. 97-266 (PC).
Malawi. Northern Prov., Mzimba distr., Mtanga tanga forest, 15 March 2005, F. Noe 05/550 (GENT); Ibid., Perekezi forest reserve, 28 Jan. 2005, A. Verbeken 05-086 (GENT); Ibid., 15 Feb. 2005, F. Noe 05/324 (GENT).
Zambia. Miputu, 22 Jan. 1996, B. Buyck & G. Eyssartier 96-153 (PC); n.l., Feb. 1996, B. Buyck & G. Eyssartier 96-292a (PC).
Notes. The type-specimen is conserved in alcohol and consists of one basidiome in rather good condition. The type-material was also studied by Lalli & Pacioni (1992) who excluded the species from Lactarius section Lactifluus.
The macroscopical description is adapted from Heim (1938b), supplemented with our field notes from Malawi. The microscopic description is based on both Heim G85 and Thoen 4635. The spores of Alexander 13 are somewhat more heavily ornamented.
An edible species according to Zeller (1982).
33. Lactarius denigricans Verbeken & Karhula
Verbeken, Persoonia 16: 219 (1996). − Type: Tanzania, Lirondo, Songea distr., 30 Jan. 1993, Saarimäki & al. 1467 (H, holotype).
Selected descriptions and icons. Verbeken, Persoonia 16: 219–222 (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 127–128 & Pl. 41–44 (1996d); Karhula & al., Karstenia 38: fig. 11 (1998); De Kesel & al., Guide des champignons comestibles du Bénin: 171 (2002); Härkonen & al., Norrlinia 10: 83 (2003).
Pileus 60–80 mm diam., convex to slightly depressed; margin irregular; pellis mat, slightly wrinkled, pale buff, becoming brownish grey to black when bruised. Lamellae decurrent, unequal with mostly short lamellulae, close, large (8 mm), rather thick, ivory coloured, blackening when bruised; edge entire and concolorous. Stipe 45–60 × 15–20 mm, cylindrical, round to applanate on section, mat, longitudinally wrinkled to grooved, pale buff, becoming brownish grey when bruised. Context in the pileus white, becoming red, then lilac and finally black by the latex; in the stipe white, then yellow, red under the surface; taste mild, smell absent. Latex abundant, transparent-whitish, changing red when exposed, finally black; taste mild. Spore-deposit not observed. – Pl. 20
Basidiospores subglobose to ellipsoid, 6.8–7.7–8.4(–8.7) × 5.6–6.3–7.1 mm (Q = 1.12–1.21–1.32; n = 50); ornamentation amyloid, low, up to 0.5 mm high, composed of irregular knotty warts and fine connective lines, never forming a complete reticulum; plage distinct, with a central amyloid spot. Basidia (38–)40–45(–47) × 7–9(–10) mm, cylindrical to slightly clavate, 4-spored; sterigmata 5–7 × 1–2 mm. Pleurolamprocystidia abundant, emergent, 60–135 × 10–11 mm, cylindrical to narrowly fusiform, rounded with thickened wall (1–2 mm). Pleuropseudocystidia not abundant, cylindrical, 7–8 mm diam.; content brownish oleiferic. Lamellar edge heterogenous, composed of lamprocystidia, basidioles and basidia. Hymenophoral trama composed of sphaerocytes and abundant broad laticiferous hyphae. Pileipellis a lampropalisade; suprapellis 30–60 mm thick; subpellis 40–50 mm thick; elements of suprapellis 15–55 × 7–8(–10) mm, cylindrical, sometimes clavate, sometimes septate, thick-walled (0–1 mm); subpellis thin, pseudoparenchymatous; spherical cells 5–15 mm. Stipitipellis a lamprotrichoderm; 90–140 mm thick; hyphae interwoven and ascending in the suprapellis; no spherical cells; terminal elements cylindrical to slightly tortuous, (10–)25–60(–80) × 6–8(–10) mm, thick-walled (1–2 mm). Clamp-connections absent. – Fig. 57
Ecology. Found in miombo woodland with mainly Uapaca and some Brachystegia, and Sudanian rivulet forest dominated by Uapaca somon.
Distribution. Known from Benin and Tanzania. Widespread but apparently rare.
Revised specimens.
Benin. Atacora Prov., Kota, 17 June 2000, A. De Kesel 2825 (BR 126378–84).
Tanzania. Southern Prov., Songea distr., Lirondo, 30 Jan. 1993, T. Saarimäki 1467 (H, holotype).
Notes. This striking species differs from any other species in the section by its colour-change of context and latex. The microscopic characters make it fit very well in Lactarius section Pseudogymnocarpi: emergent and abundant lamprocystidia, hymenophoral trama composed of sphaerocytes, pileipellis-structure a regular lampropalisade, spore-ornamentation an almost complete reticulum and the plage with a central amyloid spot. There are some resemblances with Lactarius rubroviolascens by the colour-changes, the macroscopic aspect of the pileus and the very emergent and abundant lamprocystidia, but the latter species differs in spore-ornamentation and pileipellis-structure.
Sect. Pseudogymnocarpi Verbeken
Lactarius sect. Pseudogymnocarpi Verbeken, Mycotaxon 66: 376 (1998).
− Type species: Lactarius gymnocarpoides Verbeken.
Key to tropical African species
1. Pleurocystidia abundant and emergent .................................... 2
Pleurocystidia present but inconspicuous, not emergent; stipe and lamellae remarkably yellow to greenish yellow ....................... 40. L. luteopus
2. Spores on average 12 µm long, 9 µm broad; pileus < 30 mm diam., with a striking carmine red colour ....... 39. L. carmineus
Pileus differently coloured, spores smaller ............................................... 3
3. Pileus becoming whitish pruinose when older; gill edge orange or brownish; elements of the pileipellis very elongate, up to 300 mm long ....................................................... 38. L. medusae
Pileus not whitish pruinose; gill edge concolorous; elements of the pileipellis up to 100 mm long ............ 4
4. Stipe white, contrasting with orange pileus ......................................................................................... 5
Stipe pale orange to pale yellow, more concolourous with pileus .................................................. 6
5. Spores elongate, average Q = 1.33–1.60; cap up to 7 cm diam., hymenophoral trama with sphaerocytes .... 36. L. longisporus
Spores ellipsoid, average Q = 1.20–1.27; cap < 3 cm diam., hymenophoral trama lacking sphaerocytes ..... 37. L. pumilus
6. Lamellar edge fertile; ornamentation of the spores composed of an incomplete reticulum without distinct warts .......................... 34. L. gymnocarpoides
Lamellar edge sterile; ornamentation of the spores composed of a partial reticulum and some isolated warts ................................ 35. L. pseudogymnocarpus
34. Lactarius gymnocarpoides Verbeken
Verbeken, Mycotaxon 55: 530 (1995). − Type: Burundi, Rumonge, Nkyamba, 15 March 1994, A. Verbeken 94-150 (BR 57604–83, holotype; GENT, isotype).
Misappl.: Lactarius phlebophyllus ss. Morris (1990).
Selected descriptions and icons. Verbeken, Mycotaxon 55: 530–533 (1995); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 126–127 & Pl. 28–40 (1996d); Karhula & al., Karstenia 38: fig. 5 (1998); De Kesel & al., Guide des champignons comestibles du Bénin: 164 (2002); Härkonen & al., Norrlinia 10: 86 (2003).
Pileus (40–)60–90 mm diam., quite thick, planoconvex to slightly depressed and infundibuliform; margin incurved when young, then irregular to crenulate and even undulate, sometimes grooved; pellis not dehiscent, dry, smooth to finely felty, wrinkled to cracking, greyish orange to orange when young (6B5–6), then rather light orange (5A4 to 5B4–6). Lamellae decurrent, unequal with lamellulae (1 to 3 between 2 lamellae, regular pattern), distant (L+l = 2+3 to 3+6/cm), medium broad ((3–)5–7 mm), thick, brittle, with remarkable venation when older, yellowish white to pale yellow (3A2–3 to 4A2 to 4B4); edge entire, concolorous. Stipe 25–65 × 10–20 mm, cylindrical to fusiform, tapering downwards, exceptionally clavate, dry, smooth, yellowish white to light yellow (4A2–5) or light orange (5A3–5), solid to chambered and broadly fistulous. Context firm, white; taste mild to bitter; smell pleasant. Latex not abundant (very abundant in Verbeken 94-644), fluid, white, unchanging, mild to astringent. Spore-deposit white. – Pl. 21
Chemical reactions. Context salmon pink to greyish pink with FeSO4; pale yellow with KOH; unchanging with HCl and NH4OH. Pileipellis olive green (4C5) with KOH.
Basidiospores ellipsoid, 8.2–8.9–9.3–10.3 × 6.6–7.2–7.9–8.9 mm (Q = 1.07–1.17–1.27–1.43; n = 120); ornamentation amyloid, forming an almost complete reticulum with rather big meshes which become smaller on adaxial side; prominences very low (< 0.2 mm high); plage diffusely amyloid. Basidia 50–70 × 5–13 mm, clavate, 4-spored. Pleurolamprocystidia abundant, 80–130 × 8–12 mm, fusiform, thick-walled (up to 1.5 mm). Pleuropseudocystidia abundant, 3–5(–7) mm, cylindrical, with rounded apex. Lamellar edge fertile; basidia 37–40 x 9 µm; marginal cells also present, slightly thick-walled, sometimes septate, 40–50 x 8–10 µm. Hymenophoral trama composed of sphaerocytes; laticiferous hyphae present but not very conspicuous. Pileipellis a lampropalisade; elements of the suprapellis 20–60 × 4–11 mm, regularly cylindrical, with slightly thickened wall (0.5–1 mm); subpellis pseudoparenchymatous, spherical cells (7–)10–20(–30) mm, with slightly thickened wall. Stipitipellis a trichopalisade, composed of densely interwoven hyphae with terminal elements forming a suprapellis; terminal elements 15–30 × 4–8 mm, cylindrical, sometimes clavate, thin-walled. Clamp-connections absent. – Fig. 58
Ecology. Widespread in tropical Africa, particularly common in Sudanian woodland and drier miombo woodland; also collected in Uapaca bojeri woodland, dry coastal forest, rainforest, riparian forest, semi‑evergreen forest.
Distribution. Known from Benin, Burundi, Democratic Republic of Congo, Guinea, Madagascar, Malawi, Senegal, Tanzania, Zambia and Zimbabwe.
Revised specimens
Benin. Borgou Prov., Wari Maro, 20 Aug. 1997, A. De Kesel 1949 (BR 74940–56); Ibid., 21 Aug. 1997, A. De Kesel 1950 (BR 74941–57); Ibid., 20 June 1998, A. De Kesel 2162 (BR 112673–56); Ibid., 20 Sept. 1998, A. De Kesel 2287 (BR 112780–66); Ibid., 21 Sept. 1998, A. De Kesel 2292 (BR 112785–71); Ibid., Borgou, 22 Sept. 2000, A. De Kesel 2915 (BR 129126–19). Atacora Prov., Kota, 23 Aug. 1997, A. De Kesel 1986 (BR 74838–51); Kounagnigou, 3 Sept. 1997, A. De Kesel 2044 (BR 74877–90); Ibid., 5 Sept. 1997, A. De Kesel 2062 (BR 74895–11); Kouandé, 25 Sept. 2000, A. De Kesel 2941 (BR 129152–45); Bassila, 4 Oct. 2000, A. De Kesel 3001b (BR 129213–09); Ibid. Boukombé, 2 Aug. 1996, A. De Groote 9 (BR 166524–72).
Burundi. Bururi Prov., Nkayamba, near Rumonge, March 1994, A. Verbeken 94-149 (BR 57603–82) & 94-150 (BR 57604–83, holotype); Ibid., April 1992, B. Buyck 4380, 4549 & 4551 (all PC); Ibid., Nov. 1992, B. Buyck 4629, 4630 & 4631 (all PC); Ibid., Dec. 1992, B. Buyck 4747b (PC); Ibid., Feb. 1993, B. Buyck 4937 (PC); Nyamirambo, Rumonge Forest Reserve, March 1994, A. Verbeken 94-397 (BR 57657–39), 94-429 (BR 57661–43) & 94-474 (BR 57681–63); Ruyigi Prov., Ruvubu National Park, April 1994, A. Verbeken 94-644 (BR 57712–94), 94-645 (BR 57713–95) & 94-648 (BR 57715–00) ; Cankuzo Prov., Murango, March 1994, B. Nzigidahera 53 & 54 (all PC).
Democratic Republic of Congo. Shaba Prov., Kanduba, 27 March 1986, J. Schreurs 1529 (BR 8291–46).
Guinea. Fouta Djalon Prov., Lébékéré, 14 July 1988, D. Thoen 7966 (BR 66847–14).
Madagascar. Ranohira, 60 km in direction Tulear, near TV Antenna, 3 Feb. 2000, B. Buyck & al. 00-1162 (PC); Arivonimamo, 30 Jan. 1997, B. Buyck & al. 97-045a & 97-116 (all PC).
Malawi. Southern Prov., Maloson mountain, 30 Dec. 1981, B. Morris 493 (K(M) 38469); Machinga Forest Reserve, 11 Feb. 1993, Kathumba/Chifunti 23950 (K(M) 38467); Southern Prov., Machinga distr., Liwonde Forest Reserve, 22 Jan. 2005, A. Verbeken 05-011 & 05-112 (all GENT); Ibid., 23 Jan. 2005, A. Verbeken 05-036 (GENT); Northern Prov., Mzimba distr., Perekezi Forest Reserve, 28 Jan. 2005, A. Verbeken 05-097 & 05-106 (all GENT); Ibid., 29 Jan. 2005, A. Verbeken 05-115, 05-116 & 05-144 (all GENT); Mzimba distr., 31 Jan. 2005, A. Verbeken 05-184/187 & 05-191(all GENT).
Senegal. Basse Casamance Prov., National Park of Basse Casamance, 22 Sept. 1986, D. Thoen s.n. (BR 66214–60).
Tanzania. Southern Highlands Prov., Njombe distr., Mbalali, 1 Feb.1993, T. Saarimäki & al. 1508 (H).
Zambia. Copperbelt Prov., Mwekera forest, between Kitwe and Ndola, 21 Jan. 1991, B. Buyck 3491 (PC); Northern Prov., North Luangwa National Park, 25 Jan. 1995, D. Shah‑Smith 163 (K(M) 35509); farm Gibsons, 16 Jan. 1996, B. Buyck & G. Eyssartier 96-061 (PC); road Mpongwe-Lusaka, Mikati, 18 Jan. 1996, B. Buyck & G. Eyssartier 96-098 (PC).
Zimbabwe. Midlands Prov., Mvuma, Central Estates, Beacon Hill section, 19 Feb. 1996, C. Sharp 455/96 (GENT); Ibid., 9 Jan. 1997, C. Sharp 530/97 (BR 60407–73); Ibid., 26 Jan. 1999, A. Verbeken 99-010 & 99-016 (all GENT); Mashonaland East Prov., Bromley, Liemba farm, 1 Feb. 1999, A. Verbeken 99-063 (GENT); Manicaland Prov., along Mutare-Bvumba road, near Prince of Wales Viewpoint, 11 Feb. 1999, A. Verbeken 99-172 & 99-173 (all GENT); Manicaland Prov., Chipinge, Busi farm, around Scott’s house, 12 Feb. 1999, A. Verbeken 99-183 (GENT); Ibid., 12 Feb. 1999, A. Verbeken 99-200 & 99-202 (all GENT).
Notes. The macroscopic description is based on notes from Verbeken and Buyck. The microscopic description is based on Buyck 4549, completed with Buyck 4747 and Buyck 4380.
The species is most closely related to Lactarius longisporus, from which it differs by the coloured stipe and the more ellipsoid spores which are ornamented with a looser reticulum with bigger meshes. Furthermore, the stipitipellis consists of thin-walled terminal elements whereas the terminal elements in the stipitipellis of Lactarius longisporus are thick-walled.
Lactarius gymnocarpoides is an edible species, locally well appreciated (Buyck 1994, De Kesel & al. 2002, Yorou & De Kesel 2002, Härkonen & al. 2003). It can be found in large quantities, i.e. up to 100kg/ha/y in savannah woodlands with Uapaca togoensis or Isoberlinia tomentosa (Yorou & al. 2002).
35. Lactarius pseudogymnocarpus Verbeken
Verbeken, Mycotaxon 55: 523 (1995). − Democratic Republic of Congo, Binga, M. Goossens-Fontana 899 (BR 12324–05, holotype).
Misappl.: Lactarius gymnocarpus ss. Heim (1955) pro parte, Pegler (1977).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: Pl. 2, fig. 1a,c–e (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 13, fig. 1a,c (1955); Pegler, Kew Bull., Addit. Ser. 6: 571–572 (1977, as "L. gymnocarpus"); Verbeken, Mycotaxon 55: 523–527 (1995); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 120–122 & Pl. 20–25 (1996d).
Pileus 70–140 mm diam., planoconvex and depressed to infundibuliform; margin crenulate and very irregularly lobed when older, striate to grooved (up to 1/3 of the radius), with very fine veins between the grooves; pellis not dehiscent, dry, finely tomentose, with a fine reticulum of gelatinous, sinuous veins, pale yellow, yellowish orange, bright orange to apricot, almost carrot red becoming brownish orange or with brownish or more yellowish spots. Lamellae decurrent and extending in longitudinal veins on the stipe, unequal with lamellulae of three lengths, distant (L+l = 4/cm), broad (up to 12 mm), thick, white to cream (3A3), with brownish or greyish purple spots; edge entire and concolorous. Stipe 45–100 × 10–20 mm, cylindrical, sometimes tapering downwards, somewhat rugose, whitish, pale orange to ochraceous pink downwards, with a brownish orange tomentose base. Context firm, very thin near the outer part of the pileus, white to cream, with brown spots in older specimens; taste mild. Latex not abundant, transparent to white; taste mild. Spore-deposit white. – Pl. 22
Chemical reactions. Context immediately brown with pyrogalic acid; soon greyish pink with FeSO4; intensely blue green with gaiac; slowly chocolate brown with phenol; pink with sulfovanilline; unchanging with aniline, naphthol, NH3, SF. Hymenophoral trama violently pink with sulfovanilline. (According to Heim 1955a).
Basidiospores ellipsoid, (7.1–)7.7–8.4–9.1–9.9 × 6.0–6.7–7.0–7.6 mm (Q = 1.08–1.26–1.29–1.43; n = 50); ornamentation amyloid, composed of a partial reticulum; warts connected by irregular connective lines, some isolated, up to 0.5 mm high; plage weakly amyloid. Basidia 45–80 × 8–10 mm, cylindrical to clavate, 4-spored, slightly thick-walled; content granular. Pleurolamprocystidia abundant, 50–80(–100) × 5–7 mm. Pleuropseudocystidia scarce. Lamellar edge sterile; marginal cells 15–37 × 4–7 mm, cylindrical, tapering or obtuse, often septate, thick-walled (up to 1.0 mm). Hymenophoral trama composed of sphaerocytes; laticiferous hyphae present. Pileipellis a lampropalisade; elements of the suprapellis 30–60 × 5–7 mm, regularly cylindrical, obtuse, thick-walled (1–2 mm); subpellis pseudoparenchymatous; spherical cells 10(–20) mm, with slightly thickened wall. Stipitipellis a lamprotrichoderm; elements of the suprapellis 20–60 × 5–10 mm, cylindrical, obtuse to slightly tapering upwards, thick-walled (1–2 mm). Clamp-connections absent. – Fig. 59
Ecology. Found in lowland rainforest with Isoberlinia, drier Guineo-Congolian rainforest, Sudanian savannah woodland with Isoberlinia.
Distribution. Known from Benin, Democratic Republic of Congo and Tanzania.
Revised specimens
Benin. Borgou Prov., Wari Maro, 20 June 1998, A. De Kesel 2164 (BR 112675–58).
Democratic Republic of Congo. Equateur Prov., Binga, date unknown, M. Goossens-Fontana 899 (BR 12324–05, holotype); Ibid., June 1942, M. Goossens-Fontana 3001 (BR 12323–04); Kivu Prov., Irangi, April 1972, J. Rammeloo Z244 (GENT).
Tanzania. Tanga Prov., Amani Forest, E. Usambara Mts., Lushoto, 15 April 1968, D. Pegler T497 (K(M) 159502).
Notes. The macroscopic description is primarily based on the description of Pegler (1977) completed with fieldnotes from Goossens-Fontana and Rammeloo on the specimens from Democratic Republic of Congo . There is a serious contradiction concerning the taste and smell. Pegler (1977) describes the taste as "mild" and the smell as "none", while Heim mentions a "very acrid" taste and "acrid and pepper-like" smell, based on observations made by Goossens-Fontana. Interpreting organoleptic characters noted by Goossens-Fontana must be done with caution, since her taste was strongly affected by the use of kinin (Heinemann, pers. comm.). The microscopic description is based on Goossens-Fontana 899 & 3001. The spores from Pegler T497 measure 7.4–8.6–9.7 × 5.9–6.7–7.5 mm (Q = 1.17–1.28–1.45).
Heim (1955a) noticed that there are differences in venation of the pileus and taste of context and latex, between what he calls "la forme guinéenne" and the collections of Goossens-Fontana. The remarkable differences in spore-dimensions and pileipellis-structure, however, make it clear that both "forms" concern different species. Lactarius gymnocarpus has smaller spores (7.1–8.2 × 5.9–6.4 mm) and the elements of the suprapellis of the pileus and the stipe are very irregularly shaped, varying from narrowly cylindrical to broadly fusiform or almost rounded, often tortuous, with thickened wall (1–3 mm). Lactarius pseudogymnocarpus has longer and wider spores (8.4–9.1 × 6.7–7.0 mm) and the elements of the suprapellis of the pileus and the stipe are regularly cylindrical, obtuse, thick-walled (1–2 mm). Despite the macroscopic resemblance of those two species (aspect and colour of the pileus), important microscopic differences, especially the presence of true cystidia in Lactarius pseudogymnocarpus, and the lack of true cystidia in L. gymnocarpus, make them belong to different sections.
36. Lactarius longisporus Verbeken
Verbeken, Mycotaxon 55: 527 (1995). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 1 April 1994, A. Verbeken 94-557 (BR 57699–81, holotype; GENT, isotype).
Misappl.: Lactarius gymnocarpus ss. Buyck & Nzigidahera (1996).
Selected descriptions and icons. Verbeken, Mycotaxon 55: 527–530 (1995); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 124–126 & Pl. 32–37 (1996d).
Pileus 30–75 mm diam., quite thick (20–45 mm), convex to planoconcave and slightly depressed to depressed, when old infundibuliform and irregular; margin incurved when young, then slightly crenulate and sometimes irregularly grooved when older; pellis not dehiscent, dry, mat, smooth to finely felty, wrinkled to cracked, pale orange to greyish orange (5A4 to 6A5–6 to 6B5–6), sometimes greyish red to brownish orange. Lamellae decurrent, unequal with lamellulae [3(–5) between 2 lamellae, regular pattern], very distant (L+l = 2+4 to 3+5/cm), medium broad (4–8 mm), thick, very brittle, with remarkable venation when older, white to cream or yellowish; edge entire, concolorous (pale yellow in Buyck 3583). Stipe 25–50 × 8–12 mm, cylindrical, slender, dry, smooth to very finely felty, white, chambered to hollow. Context firm, white to cream; taste mild to bitter; smell Cantharellus-like or absent. Latex not abundant, white, unchanging; taste mild to bitter. Spore-deposit white. – Pl. 23
Chemical reactions. Context salmon pink (sometimes nihil) with FeSO4; unchanging with HCl and NH4OH. Pileipellis light brown to greyish brown with KOH (Verbeken 94-520).
Basidiospores ellipsoid to strongly elongate, 8.4–9.1–11.4–12.5 × 6.1–6.6–7.6–8.4 mm (Q = 1.22–1.33–1.60–1.80, n = 320); ornamentation amyloid, < 0.1 mm high, composed of a dense and regular reticulum (without isolated warts); plage distinct, amyloid. Basidia (48–)54–60(–80) × (8–)9–10.5(–12) mm, cylindrical to clavate, 4-spored. Pleurolamprocystidia quite abundant, emergent, (55–)70–80(–90) × 8–12 mm, cylindrical to narrowly fusiform, with slightly thickened wall (to 1 mm); content slightly granular. Pleuropseudocystidia present, undistinct, cylindrical, sometimes capitate; content oleiferic. Lamellar edge fertile. Hymenophoral trama composed of sphaerocytes; laticiferous hyphae present but inconspicuous. Pileipellis a lampropalisade; elements of the suprapellis (10–)35–50(–120) × 4–10 mm, cylindrical, tortuous, sometimes branched and septate, thick-walled (1–2 mm); subpellis pseudoparenchymatous; spherical cells (5–)10–25(–35) mm, thick-walled (0.5–1.5 mm). Stipitipellis a lamprotrichopalisade, hyphae interwoven and ascending in suprapellis; spherical cells scarce; terminal elements very tortuous, sometimes with corkscrew-like curls, 15–45 × 3–4 mm, slightly thick-walled; some terminal elements cylindrical, 25–45 × 4–6 mm; slightly thick-walled (< 1 mm). Clamp-connections absent. – Fig. 60
Ecology. Growing in woodlands dominated by ectomycorrhizal trees.
Distribution. Known from Benin, Burundi, Democratic Republic of Congo, Kenya, Madagascar, Malawi, Zambia and Zimbabwe.
Revised specimens
Benin. Atacora Prov., Kota, 29 Sept. 2000, A. De Kesel 2966 (BR 129213–09).
Burundi, Bururi Prov., Nyamirambo, Rumonge Forest Reserve, March 1994, A. Verbeken 94-019 (BR 57569–48), 94-066 (BR 57577–56), 94-078 (BR 57588–67), 94-101 (BR 57595–74), 94-220 (BR 57628–10), 94-277 (BR 57634–16), 94-278 (BR 57635–17), 94-282 (BR 57636–18), 94-348 (BR 57648–30), 94-430 (BR 57662–44), 94-520 (BR 57690–72), 94-521 (BR 57691–73) & 94-522 (BR 57692–74); Ibid., April 1994, A. Verbeken 94-557 (BR 57699–81, holotype; GENT, isotype); Ibid., Dec. 1993, B. Buyck 5234, 5235, 5299 & 5300 (all PC); Nkayamba, near Rumonge, Dec. 1992, B. Buyck 4675 & 4754 (all PC); Ibid., March 1993, B. Buyck 5044, 5057 & 5139 (all PC); Rubindi, between Kigwena and Nyanza, 16 Apr. 1978, J. Lambinon 78/169 (LG).
Kenya. Coast Prov., Kwale distr., Arabuko-Sokoke Forest Reserve, plot 1, Aug. 1994, M.H. Ivory IV934 (BR 49370–94); Ibid., plot 3, Nov. 1994, M.H. Ivory IV5025 (BR 49369–93).
Democratic Republic of Congo. Shaba Prov., Ruashi, Jan. 1949, D. Soyer 303 (BR 4905–55); Kipopo, Dec. 1959, M.C. Schmitz-Levecq 185 (BR 12298–76).
Madagascar. Ranohira, 60 km in direction Tulear, near TV Antenna, 3 Feb. 2000, B. Buyck & al. 00-1163 (PC); Ambotantihely, N of Ankazobe, sentier botanique, 15 Feb. 2000, B. Buyck & al. 00-1519 (PC).
Malawi. Northern Prov., Mzimba distr., Perekezi forest reserve, 28 Jan. 2005, A. Verbeken 05-101 (GENT); Ibid., 29 Jan. 2005, A. Verbeken 05-119, 05-120 & 05-130 (all GENT); Ibid., 29 Jan. 2005, F. Noe 05/12 (GENT); Ntanga forest reserve, 30 Jan. 2005, A. Verbeken 05-164 (GENT).
Zambia. Copperbelt Prov., Chati-forest, near Kitwe, Dec. 1990, B. Buyck 3135, 3136, 3201, 3202 & 3209 (all PC); Luapala Prov., Samfya, Dec. 1990, B. Buyck 3256, 3278 & 3279 (all PC); Mansa, Kabunda mission, Jan. 1991, B. Buyck 3351 (PC); Western Prov., Mwinilunga, road to Solwezi, Jan. 1991, B. Buyck 3510, 3513 & 3583 (all PC); farm Gibsons, 16 Jan. 1996, B. Buyck & G. Eyssartier 96-062 (PC); Chibuli, 31 Jan. 1996, B. Buyck & G. Eyssartier 96-251, 96-252 & 96-253 (all PC).
Zimbabwe. Midlands Prov., Mvuma, Central Estates, Baru section, 27 Jan. 1999, A. Verbeken 99-034 (GENT); Midlands Prov., Mvuma, Central Estates, south of Beacon hill ridge, 27 Jan. 1999, A. Verbeken 99-022 (GENT); Ibid., 1 Jan. 1997, C. Sharp 504/97 (BR 60404–70); Ibid., 3 Jan. 1997, C. Sharp 516/97 (BR 60406–72); Ibid., 09 Jan. 1997, C. Sharp 535/97 (BR 60408–74); Ibid., 10 Jan. 1997, Sharp 537/97 (BR 60409–75); Bromley, Liemba farm, 3 Feb. 1999, A. Verbeken 99-098 (GENT); Manicaland Prov., Chipinge, Busi farm, around Scott’s house, 12 Feb. 1999, A. Verbeken 99-197 (GENT).
Notes. The macroscopic description is mainly based on notes from Verbeken, completed with notes from Buyck. The microscopic description is based on Buyck 5139, 4675, 4754, 3135 & 5235.
The species is easily recognized by the orange pileus, contrasting with the white stipe and the elongate spores with the ornamentation forming a dense reticulum. Especially in young condition it shares some macroscopic characters with Lactarius pumilus, from which it differs by the much longer spores with a denser reticulum and by the hymenophoral trama, here composed of sphaerocytes, whereas composed of hyphae in L. pumilus.
In some specimens (Buyck 4754 & 3351) small, irregularly diverticulate elements are observed in the hymenium. It is not clear whether those are cystidia or deformed basidioles.
This species is locally eaten (e.g. Nzigidahera 1993, Verbeken & al. 2000).
37. Lactarius pumilus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 205 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 1 April 1994, A. Verbeken 94-559 (BR 57702–84, isotype GENT).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 122–123 & Pl. 26–28 (1996d); Karhula & al., Karstenia 38: fig. 9 (1998); Härkonen & al., Norrlinia 10: 90 (2003).
Pileus 10–30 mm diam., firm, fleshy, planoconvex and depressed, rather irregular; margin incurved, irregularly lobate, then crenulate; pellis dry, finely felty, pale orange, greyish orange to brownish orange or reddish blond (5A3 to 5B3 to 5C3). Lamellae slightly decurrent, unequal with 1–3 lamellulae between two lamellae (regular pattern, lamellulae mostly very short), moderately distant (L+l = 4+9/cm), elastic, paper-like, pale yellow to light yellow (4A3–4); edge entire, darker. Stipe 10–25 × 5–10 mm, short cylindrical, dry, finely felty, pale orange to orange white (5A3–4) in upper half, more whitish near the base, firm, solid, sometimes subfistulous. Context firm, white, slightly yellowing when cut or bruised; smell not remarkable; taste mild. Latex scarce, white. Spore-deposit not observed. – Pl. 24
Chemical reactions. Context pale orange with FeSO4. Pileipellis dark brown (8F3) with KOH; greyish brown (8D3) with NH4OH.
Basidiospores sometimes subglobose, sometimes elongate, mostly ellipsoid, 7.2–8.5–8.9–9.9(–10.3) × 5.9–6.7–7.4–8.0(–8.3) mm (Q = 1.12–1.20–1.27–1.45, n = 40); ornamentation amyloid, composed of narrow, low ridges, forming an incomplete to almost complete reticulum, without isolated warts; plage sometimes with central amyloid spot. Basidia (30–)40–45(–50) × 8–11 mm, subclavate, 4-spored, sometimes 2-spored; content guttate. Pleurolamprocystidia very abundant, emergent, 50–80 × 5–8(–10) mm, cylindrical to subcylindrical, rounded at the apex, rarely constricted and tapering upwards, thick-walled, hyaline or with needle-like content. Pleuropseudocystidia scarce, not emergent, 3–5 mm diam., cylindrical, rounded at the apex; content oleiferic. Lamellar edge sterile; marginal cells 10–25 × 3–5 mm, subfusiform to irregular, sometimes branched, hyaline, thin-walled or slightly thick-walled. Hymenophoral trama irregular, composed of narrow, thin-walled, densely interwoven hyphae and abundant lactifers. Pileipellis a lampropalisade; suprapellis composed of narrow, long, cylindrical elements; terminal elements thick-walled, 10–100 × 3–7 mm, sometimes septate; subpellis pseudoparenchymatous; isodiametric cells thin-walled or slightly thick-walled, 5–15(–20) mm diam. Stipitipellis a layer of ascending, narrow, long, cylindrical elements, thin-walled or slightly thick-walled, 10–100(–200) × 3–5 mm, sometimes septate, remarkably corkscrew-like at the end, originating from a layer of hyaline, thin-walled, recumbent hyphae; no sphaerocytes. Clamp-connections absent. – Fig. 61
Ecology. Found in miombo woodland, semi-evergreen forest, riverine woodland, Sudanian savannah woodland with Isoberlina or Afzelia.
Distribution. Known from Benin, Burundi, Kenya, Senegal, Tanzania, Zambia and Zimbabwe.
Revised specimens
Benin. Atacora Prov., Kota, 23 Aug 1997, A. De Kesel 1987 (BR 74839–52); Ibid., 1 Oct 1999, A. De Kesel 2731 (BR 116740–49).
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, 1 April 1994, A. Verbeken 94-559 (BR 57702–84, holotype; GENT, isotype); Nkayamba, just N of Rumonge, 1 March 1993, B. Buyck 5000 (PC).
Kenya. Coastal Prov., Kayatelesi, Shimba Hills, 4 Dec. 2004, A. De Kesel 3977 (BR 166508–56).
Senegal. Basse Casamance, 22 Aug. 1987, D. Thoen 7618 (BR 3155–51).
Zambia. Northern Prov., North Luangwa National Park, Mwaleshi Camp, 20 Jan. 1995, D. Shah‑Smith 156 (K(M) 35501); Copperbelt Prov., Chati forest near Kitwe, 15 Dec. 1990, B. Buyck 3137 (PC).
Zimbabwe. Manicaland Prov., Chipinge, Busi farm, around Scott’s house, 12 Feb. 1999, A. Verbeken 99-198 (GENT); Mashonaland East Prov., Bromley, Liemba farm, 2 Feb. 1999, A. Verbeken 99-070 (GENT); Masvingo Prov., south of Lake Mutirikwe, around Norma Jean’s lodge, 16 Feb. 1999, A. Verbeken 99-222 (GENT).
Notes. The macroscopic description is based on fieldnotes of Buyck; the microscopic description is based on Buyck 5000 and Verbeken 94-559.
This is the only species in this group where no sphaerocytes were observed in the trama. In all other characters the species fits well in this section. The species is characterized by the very small and bright orange basidiomes (contrasting with the white stipe). The pileus does not exceed 3 cm in diam.
This is an edible species (e.g. Karhula & al. 1998).
38. Lactarius medusae Verbeken
Verbeken, Mycotaxon 55: 532 (1995). − Type: Zambia, near Kawambwa, 6 Jan. 1991, B. Buyck 3455 (BR 38489–77, holotype; PC, isotype).
Misappl.: Lactarius vellereus ss. Morris (1984, 1987, 1990) and Williamson (1975, 1976); Lactarius. latifolius ss. Parent & Thoen (1977); Lactarius gymnocarpus ss. Pegler & Piearce (1980).
Selected descriptions and icons. Verbeken, Mycotaxon 55: 532–535 (1995); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 123–124 & Pl. 29–31 (1996d); Karhula & al., Karstenia 38: fig. 10 (1998); Härkonen & al., Norrlinia 10: 89 (2003).
Pileus (63–)70–120 mm diam., moderately thick, planoconcave or applanate to slightly depressed; margin irregularly lobulate in old specimens; pellis not dehiscent, velutinous, mat, dry, pruinose, concentrically wrinkled, dark orange or pinkish brown, pale yellow to yellowish white (4A2–3) when dried. Lamellae subdecurrent, decurrent or adnate, distant (L+l = 4–5/cm), medium broad to broad (3–9 mm), rather thick, cream; edge entire, brown or orange. Stipe 40–65 × 12–27 mm, central to eccentric, irregularly cylindrical, slightly tapering downwards, smooth, pruinose and wrinkled at the base, reddish blond to brownish orange (5C4), darker towards the base. Context brittle, more solid in the stipe, yellowish white to white, orange under the pileipellis; taste mild; smell pleasant, slightly acid. Latex white, changing violet on the context; taste mild. Spore-deposit not observed. – Pl. 25 & 26
Basidiospores ellipsoid, 7.5–8.5–9.2–10.3 × 5.3–6.6–7.1–7.8 mm (Q = 1.15–1.28–1.31–1.42; n = 60); ornamentation amyloid, composed of a complete and dense reticulum; warts never isolated, up to 0.3 mm high; connective lines narrow; plage distinct, centrally amyloid. Basidia 55–65(–85) × 7–9(–11) mm, cylindrical to slightly clavate, 4-spored; content granular. Pleurolamprocystidia abundant, emergent up to 35 mm, 60–80 × 5–6,5 mm, cylindrical, with thickened wall (up to 2 mm). Pleuropseudocystidia not abundant, 3–5 mm, cylindrical, obtuse. Lamellar edge sterile; marginal cells 45–70 × 4–7 mm, cylindrical to slightly clavate; apex obtuse; wall thickened up to 1 mm. Hymenophoral trama composed of sphaerocytes; laticiferous and oleiferic hyphae abundant. Pileipellis a lampropalisade; suprapellis up to 300 mm, elements of the suprapellis extremely long, (20–)80–250 × 3–6 mm, regularly cylindrical, tapering, sometimes branched, with thickened wall (1–2 mm); subpellis pseudoparenchymatous, spherical cells (5–)10–20 mm, with slightly thickened wall. Stipitipellis a lampropalisade, idem as pileipellis, spherical cells less distinct. Clamp-connections absent. – Fig. 62
Ecology. Miombo woodland.
Distribution. Known from Democratic Republic of Congo, Malawi, Tanzania, Zambia and Zimbabwe.
Revised specimens
Democratic Republic of Congo. Shaba Prov., bought by Parent on road to Likasi, March 1973, D. Thoen 5741 (BR 66366–18).
Malawi. Northern Prov., 3 miles E of Mzuzu, Katoto, March 1973, J. Williamson 629 (K(M) 34216); Southern Prov., Mulanje distr., Mulanje mountains, Likabula Valley, 12 March 1973, L. Ryvarden 11507 (K(M) 38472); Makwawa, Zomba distr., March 1980, B. Morris 110a (K(M) 159574); Northern Prov., Mzimba distr., Ntanga Forest Reserve, 30 Jan. 2005, A. Verbeken 05-145, 05-149 & 05-153 (all GENT); Ibid., Mtanga tanga forest, near Chicangana village, 19 Feb. 2005, F. Noe 05/400-411 (all GENT); Ibid., Perekezi Forest Reserve, 29 Jan. 2005, A. Verbeken 05-114 (GENT); Ibid., Mzimba distr., 31 Jan. 2005, A. Verbeken 05-181 (GENT).
Tanzania. Southern Highlands Prov., Njombe distr., Mbalali, 1 Feb. 1993, T. Saarimäki & al. 1502 (H); Iranga distr., Sao Hill Sawmills, 29 March 1991, T. Saarimäki & al. 763 (H); Iranga distr., Mafinga, 6 Feb. 1993, T. Saarimäki & al. 1589 (H).
Zambia. Luapala Prov., near Kawambwa, 6 Jan. 1991, B. Buyck 3455 (BR 38489–77, holotype; PC, isotype); Northern Prov., North Luangwa National Park, Mwaleshi Camp, 15 Feb. 1995, D. Shah‑Smith 236 (K(M) 35505); road Mpongwe to Lusaka, 18 Jan. 1996, B. Buyck & G. Eyssartier 96-099 (PC); Copperbelt Prov. Kitwe, 21 Dec. 1978, G. Piearce FP600/1c (K(M) 38494).
Zimbabwe. Manicaland, Vuma, Chinziwa Mountain, 24 Jan. 2006, C. Sharp 1788 (GENT); n.l., Feb. 1999, D. Arora 99-112 (SFSU).
Notes. The macroscopic description is mainly based on fieldnotes of Saarimäki & al. The microscopic description is based on Buyck 3455.
Small, irregularly diverticulate elements are observed in the hymenium (Buyck 3455). It is not clear whether those are cystidia or deformed basidioles.
The species is easily recognized by the pruinose and velutinous pileus, the coloured lamella edge, and microscopically by the very long terminal elements in the pileipellis.
Lactarius medusae is a well appreciated edible milk cap (e.g. Karhula & al. 1998, Morris 1984).
39. Lactarius carmineus Verbeken & Walleyn
Verbeken & al., Syst. Geogr. Pl. 70: 190 (2000). – Type: Zimbabwe, Mashonaland East, Bromley, Liemba farm, 3 Feb. 1999, A. Verbeken 99-099 (GENT, holotype).
Selected descriptions and icons. Verbeken & al., Syst. Geogr. Pl. 70: 190–192 (2000).
Pileus 13–30 mm diam., planoconvex to almost applanate, with an undeep depression and often an irregular papilla; pellis velvety in very young specimens, later still a bit velvety in center, but rugose towards margin, carmine (10F8, “violet brown”) with paler margin (10E8) in young material, later reddish brown to carmine (9E8 to 8D8), brownish orange (6C4) towards margin; margin striate almost up to center, sulcate with deep and broad grooves. Lamellae very distant (L+l = 14–16/half of the pileus), sometimes forked, with abundant lamellulae (short ones, without regular pattern), decurrent in older specimens, adnate in younger material, pale orange (5A3), with irregular venations in between. Stipe 10–30 × 3–6 mm, irregularly cylindric, smooth, brownish orange, greyish orange, pale orange (6B6–7 to 6C6–7 to 6B4 or 6A3), very brittle. Context very brittle, thin-fleshed in pileus, with abundant chambers in the stipe, whitish; taste mild; smell not remarkable. Latex watery white, unchanging with KOH. Spore-deposit not observed. – Pl. 26
Chemical reactions. Context slightly greyish with FeSO4, unchanging with gaiac.
Basidiospores ellipsoid, 11–12.1–13.4 × 8.1–9.0–9.8 µm (Q = 1.24–1.34–1.50, n = 20); ornamentation amyloid, composed of irregular, mostly elongated warts of 0.2 µm high, aligned or connected and forming a very incomplete but extremely dense reticulum; plage not amyloid. Basidia 55–70 × 12–15 µm, subclavate, 4-spored, thin- or thick-walled (1–2 µm); some 1-spored or deformed basidia present. Pleurolamprocystidia abundant, emergent, 100–120 × 10–12 µm, very thick-walled (2–3 µm), lanceolate. Leptocystidia (deformed basidioles) present, 20–30 × 6–8 µm, thin-walled, hyaline, with diverticulate knobs. Pleuropseudocystidia scarce, not emergent. Hymenophoral trama cellular. Lamellar edge sterile; marginal cells very irregularly shaped, with diverticulate knobs, hyaline, thin-walled. Pileipellis a lampropalisade, 100–150 µm thick; terminal elements cylindric or slightly irregular, with rounded apex, 25–80 × 5–10 µm, thick-walled, often septate; subpellis composed of rather small (5–10 µm diam.) and thin-walled globose cells. Stipitipellis a lamprotrichoderm; terminal elements slightly to clearly thick-walled, 20–80 × 4–8 µm. Clamp-connections absent. – Fig. 63
Ecology. Drier Zambezian miombo woodland.
Distribution. Only known from Zimbabwe and Malawi.
Malawi. Northern Prov., Mzimba distr., Ntanga Forest Reserve, 30 Jan. 2005, A. Verbeken 05-146 (GENT).
Zimbabwe. Mashonaland East Prov., Bromley, Liemba farm, Carters estates, 3 Feb. 1999, A. Verbeken 99-099 (GENT, holotype).
Notes. The species is recognized by the striking carmine to reddish brown cap-colour (especially in young specimens) and the large spores, some of the largest known in African Lactarius species.
In the Malawian material we observed some mature basidia at the lamella edge.
40. Lactarius luteopus Verbeken
Verbeken, Mycotaxon 55: 536 (1995). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, A. Verbeken 94-463 (BR 57669–51, holotype; GENT, isotype).
Misappl.: Lactarius caperatus ss. Morris (1990).
Selected descriptions and icons. Verbeken, Mycotaxon 55: 536–539 (1995); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 119–120 & Pl. 17–19 (1996d); Karhula & al., Karstenia 38: fig. 1 (1998); De Kesel & al., Guide des champignons comestibles du Bénin: 172 (2002); Härkonen & al., Norrlinia 10: 88 (2003).
Pileus 20–60(–100) mm diam., rather thin (1–3 mm), planoconvex and slightly depressed to infundibuliform and irregularly so, sometimes with a small papilla; margin slightly incurved when young, then irregular to crenulate and even undulate; pellis not dehiscent, dry, smooth, finely wrinkled, especially towards the margin, sometimes cracking, pale yellow to orange yellow (3A4–5, 4A4–8) or deep orange to greyish orange (5A6, 5B5–6); margin paler yellow when young. Lamellae decurrent, unequal with lamellulae (1 to 3 between 2 lamellae, regular pattern), distant (L+l = 2+3 to 5+15/cm), medium broad [4–7(–11) mm], thick, very brittle, with remarkable venation when older, vivid yellow when young (4A8), rather pale yellow (4A4) or pale orange (5A5) to even yellowish white (3A2) when older; edge entire, concolorous. Stipe 10–35(–45) × 6–15(–20) mm, shortly cylindrical, often tapering downwards, dry, smooth, vividly yellow (23A7–8) to greenish yellow (1A7–8), when paler then always yellow at the base, firm, becoming fistulous. Context fragile, white, vividly yellow around the cavities in the stipe; taste mild, pleasant; smell pleasant, Cantharellus-like. Latex not abundant, white, unchanging, mild. Spore-deposit white. – Pl. 27
Chemical reactions. Context immediately orange or faintly so with FeSO4, unchanging with HCl, KOH, NH4OH.
Basidiospores ellipsoid to elongate, 7.3–8.3–8.9–9.9 × 5.8–6.3–6.7–7.3 mm (Q = 1.17–1.26–1.41–1.55; n = 180); ornamentation amyloid, forming an incomplete to almost complete, irregular reticulum; isolated warts also present but rare; prominences very low (< 0.2 mm high); plage centrally amyloid. Basidia 45–60 × 7–10 mm, cylindrical, (2)4-spored. Pleurocystidia very scarce, not very emergent, cylindrical to narrowly fusiform, often septate, 55–70 × 7–10 mm, with a slightly thickened wall; content granular. Pleuropseudocystidia not abundant, 3–5 mm, cylindrical, obtuse. Lamellar edge fertile. Hymenophoral trama composed of sphaerocytes; laticiferous hyphae present but not very conspicuous. Pileipellis a lampropalisade; elements of the suprapellis 15–70 × 3–6 mm, cylindrical, sometimes branched, sometimes septate, slightly tapering, densely interwoven, with slightly thickened wall, subpellis pseudoparenchymatous. Stipitipellis a lampropalisade; elements of the suprapellis 25–55 × 4–8 mm, cylindrical, sometimes slightly clavate, thick-walled (up to 1.5 mm); subpellis pseudoparenchymatous. Clamp-connections absent. – Fig. 64
Ecology. Zambezian miombo woodland, Sudanian woodland. Found e.g. under Isoberlinia, Uapaca, Brachystegia.
Distribution. Known from Benin, Burundi, Malawi, Tanzania, Zambia and Zimbabwe.
Revised specimens
Benin. Atacora Prov., Kota, 23 Aug. 1997, A. De Kesel 1988 (BR 74840–53); Ibid., 29 Aug. 1997, A. De Kesel 2018 (BR 74864–77); Ibid. 7 Jun 2002, A. De Kesel 3310 (BR 152185–89); Ibid. 18 Jun. 2004, A. De Kesel 3694 (BR 157125–82); Donga Prov., Bassila, 4 Oct. 2000, A. De Kesel 3001 (BR 129213–09); Ibid. 25 Jun. 2002, A. De Kesel 3456 (BR 152259–66); Ibid. 17 Jun 2004, A. De Kesel 3665a (BR 157032–86) & 3665b (BR 157034–88); Kikélé, 13 Jun. 2002, A. De Kesel 3396 (BR 152106–10); Pénésoulou, 8 Oct. 2002, A. De Kesel 3543 (BR 152028–29); Borgou Prov., Doguè, 10 Oct. 2001, A. De Kesel 3210 (BR 149716–45); Wako, 11 Sep. 2001, A. De Kesel 3136 (BR 149795–27); Wari Maro, 20 Jun. 1998, A. De Kesel 2159 (BR 112670–53); Ibid., A. De Kesel 2160 (BR 112671–54); Ibid., 21 Sept. 1998, A. De Kesel 2290 (BR 112783–69); Ibid., 22 Sept. 2000, A. De Kesel 2914 (BR 112783–69).
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, March 1994, A. Verbeken 94-347 (BR 57647–29), 94-428 (BR 57660–42), 94-463 (BR 57669–51, holotype; GENT, isotype) & 94-464 (BR 57670–52); Nkayamba, near Rumonge, Dec. 1992, B. Buyck 4807 (PC); Mugara, 18 Nov. 1978, J. Rammeloo 5785 (BR 8768–38).
Malawi. Southern Prov., Makwawa, Zomba, April 1991, B. Buyck 3976 (PC); Zomba distr., near R.C. Cathedrale, 9 Jan. 1974, J. Williamson 768 (K(M) 38484); Central Prov., Lilongwe distr., Bunda, 26 Jan. 1972, J. Williamson 581 (K(M) 38475); Southern Prov., Machinga distr., Liwonde Forest Reserve, 23 Jan. 2005, A. Verbeken 05-047 (GENT); Northern Prov., Mzimba distr., Perekezi Forest Reserve, 28 Jan. 2005, A. Verbeken 05-081 & 05-100(all GENT); Ibid., 29 Jan. 2005, A. Verbeken 05-117 & 05-118 (all GENT); Mzimba distr., Ntanga Forest Reserve, 30 Jan. 2005, A. Verbeken 05-148 (GENT); Ibid., 31 Jan. 2005, A. Verbeken 05-174 (GENT).
Tanzania. Southern Prov., Songea distr., Mawa, 27 Jan. 1993, T. Saarimäki & al. 1415 & 1424 (all H); Songea distr., Hanga, Nyamagoma, 27 Jan. 1993, T. Saarimäki & al. 1432 (H); Southern Highlands Prov., Njombe distr., Ngaramira, 31 Jan. 1993, T. Saarimäki & al. 1486 (H); Mbalali, 1 Feb. 1993, T. Saarimäki & al. 1504 (H); Lake Prov., Bitarambo distr., Bwanga Research Station, 4 Jan. 1973, D.L. Ebbels F14 (K(M) 38488).
Zambia. Luapala Prov., Samfya, Dec. 1990, B. Buyck 3252, 3270, 3272, 3281, 3282 & 3285 (all PC); Mansa, Kabunda mission, Dec. 1990, B. Buyck 3286 & 3287 (all PC); Copperbelt Prov., Misaka Forest Reserve, Dec. 1982, G. Piearce FP731/3 (K(M) 38499); road Mpongwe-Lusaka, Mikati, 18 Jan. 1996, Buyck & Eyssartier 96-097 (PC); farm Gibsons, 20 Jan. 1996, B. Buyck & G. Eyssartier 96-145 (PC); Lusaka, near airport, 11 Feb. 1996, B. Buyck & G. Eyssartier 96-350 (PC).
Zimbabwe. Midlands Prov., Mvuma, Central Estates, along road to Kwekwe, 15 Feb. 1999, A. Verbeken 99-217 (GENT); Mashonaland Central Prov., Mazowe Botanical Reserve, 16 Feb. 1988, L. Ryvarden 25091 (GENT).
Notes. The macroscopic description was based on fieldnotes from Buyck and Verbeken. The microscopic description was adapted from Buyck 3281 & 3286.
A 5-spored basidium was observed in Buyck 3281.
In Rammeloo 5785 small, irregularly diverticulate elements are observed in the hymenium. It is not clear whether those are cystidia or deformed basidioles.
The species is easily recognized in the field by the vividly yellow stipe and vividly yellow young lamellae. It differs from the other species in the section by the scarce, not emergent true cystidia, but is nevertheless classified in Lactarius section Pseudogymnocarpi given the other microscopic and macroscopic characters.
This is an edible species (e.g. De Kesel & al. 2002, Yorou & al. 2002, Karhula & al. 1998).
Lactarius sect. Rugati (Pacioni & Lalli) Verbeken, Mycotaxon 66: 372 (1998). Lactarius subsect. Rugati Lalli & Pacioni, Mycotaxon 44: 190 (1992).
− Type species: Lactarius rugatus Kühner & Romagn.
Key to tropical African species
1. Pileipellis with orange tinges, rather tomentose, not veined or wrinkled, at most slightly wrinkled; stipe short compared to pileus diameter ........................................................................................ 2
Pileipellis with reddish brown tinges, finely but distinctly veined and wrinkled or cracking; stipe longer (3–8 cm) ......... 3
2. Lamellae pure white and contrasting with the orange to yellowish orange stipe and pileus ..... 41. L. volemoides
Lamellae orange ................................ 42. L. pseudovolemus
3. Lamellae dense, cap often cracking ................................................................... 45. L. xerampelinus
Lamellae (moderately) spaced, not cracking ........................................... 4
4. Pileus orange-red to brown; spores 9 × 6.5–7 mm; warts small, mostly elongate and up to 0.1 mm high; elements of stipitipellis (20–)50–100 ´ 4–6 mm, thick-walled, extremely long and tortuous at the base of the stipe ....... 43. L. rubiginosus
Pileus dark brown when young, then paler and more reddish, sometimes with ochraceous spots; spores smaller, 7.9–8.4 ´ 5.9–6.3 mm, warts more rounded, up to 0.2 mm high; elements of the stipitipellis 10–40 ´ 2–4 mm, thin-walled, only thick-walled at the base ........ 44. L. kivuensis
41. Lactarius volemoides Karhula
Karhula & al., Karstenia 38: 53 (1998). − Type: Tanzania, Mbeya distr., Mbozi, 24 March 1991, T. Saarimäki & al. 705 (H, holotype).
Selected descriptions and icons. Karhula & al., Karstenia 38: 53–55, fig. 6 (1998); Verbeken & al., Syst. Geogr. Pl. 70: 211 (2000); Härkonen & al., Norrlinia 10: 92 (2003).
Pileus 90 mm diam., undulate, slightly depressed; pellis finely velvety, soft, dry, wrinkled and radially wrinkled at the margin, with an irregular surface, deep orange to brownish orange (6B8 to 6C8) in the center, a bit paler and orange (5A6) near margin, orange white to pale orange (5A2–3) at margin; margin irregularly widely crenulate. Lamellae decurrent with tooth, very distant (L+l = 22/half a pileus), rather thick, brittle, pure white; edge entire, concolorous. Stipe 60 × 30 mm, irregularly cylindric, tapering; pellis smooth, glabrous, dry, light yellow to pale orange (4A4 to 5A3), unicolorous except for the upper zone (zone of the lamellae teeth) where the stipe is pure white as the lamellae (a very distinct separation). Context thick and firm but very thin at the margin of the pileus, firm and solid in the stipe, white, unchanging; taste agreeable; smell herb-like, agreeable. Latex rather abundant, white, pale brownish red when drying on the lamellae; taste mild. Spore-deposit not observed. – Pl. 28
Chemical reactions. Context slowly blueing with gaiac, salmon with FeSO4.
Basidiospores ellipsoid to elongate, 8.5–9.5–10.5 × 6.0–7.0–8.5 µm (Q = 1.15–1.40–1.50); ornamentation amyloid, composed of narrow and low ridges, up to 0.2 µm high, forming a very incomplete reticulum, some very short ridges or irregular elongate warts present; plage centrally amyloid. Basidia 55–75 × 9–11 µm, narrowly cylidric, long and slender, 4-spored; content guttate. Pleurocystidia absent. Pleuropseudocystidia not abundant, not emergent, 3–6 µm diam., cylindric with rounded apex, with refractive oil-like and guttate content. Lamellar edge sterile; marginal cells cylindric, 25–40 × 4–7 µm, hyaline, thin-walled. Hymenophoral trama mixed, composed of hyaline hyphae, sphaerocytes (forming distinct rosettes) and abundant lactifers. Pileipellis a lampropalisade; suprapellis 40–50 µm thick, composed of erected hairs which are 10–50 × 3–6 µm, subcylindric to subfusiform, tapering upwards, often septate; subpellis 60–80 µm, composed of globose cells of 5–30 µm diam. with a slightly thickened wall. Clamp-connections absent. – Fig. 65
Ecology. Found in drier Zambezian miombo woodland, Sudanian riverine forest with Uapaca somon.
Distribution. Known from Benin, Tanzania and Zimbabwe.
Revised specimens
Benin. Atacora Prov., Kota, 29 Aug 1997, A. De Kesel 2019 (BR 74865–78).
Tanzania. Southern Highlands Prov., Mbeya distr., Mbozi, 24 March 1991, T. Saarimäki & al. 705 (H, holotype); Ibid., Mbeya distr., Ipembe Hill, 28 March 1991, T. Saarimäki & al. 749 (H).
Zimbabwe: Midlands Prov., Mvuma, Central Estates, Beacon Hill section, leg. R. Walleyn 27 Jan. 1999, A. Verbeken 99-020 (GENT).
Notes. This species is eaten in Tanzania (e.g. Karhula & al. 1998).
42. Lactarius pseudovolemus R. Heim
Heim, Candollea 7: 378 (1938). – Type: Madagascar, S of Maningory, 7 Dec. 1934, R. Heim G61 (PC, holotype).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 49–51 (1938); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 109–110 & Pl. 1–3 (1996d).
Pileus 50–60 mm diam., firm, fleshy, thick, convex, then deeply depressed; margin largely incurved; pellis not dehiscent, finely tomentose-granular, dry, orange-brown near the margin, dark brownish red in the centre. Lamellae adnate to decurrent, unequal with lamellulae of different lengths, distant (L+l = 36/total), rather narrow (1.5–3.5 mm), thick, brittle, orange; edge sharp, entire, concolorous. Stipe 30–35 × 8–9 mm, cylindrical, finely tomentose-granular, orange-brown, slightly paler than the pileus, with tough mycelium towards the base, solid; base slightly rooting. Context cream, unchanging, becoming blackish to brownish in alcohol; smell not particular; taste not particular. Latex abundant, white, unchanging; taste mild, a bit astringent. Spore-deposit white. (According to Heim 1938b).
Chemical reactions. Context slowly blue-green with gaiac; slowly lilac with gaiacol; slowly purple with pyramidol.
Basidiospores ellipsoid, sometimes elongate, 7.0–8.0–9.0 × 5.4–6.0–6.6(–7) mm (Q = 1.24–1.33–1.42, n = 30); ornamentation amyloid, composed of elongated warts, aligned or connected by fine connective lines, seldom isolated, forming an almost complete reticulum, up to 0.2 mm high; plage inamyloid. Basidia 45–50 × 9–10 mm, cylindrical to subclavate, 4-spored, sometimes 2-spored; sterigmata 3–4 × 1–2 mm. Pleurocystidia absent. Pleuropseudocystidia abundant, slightly emergent, (2–)4–7 mm diam., cylindrical, tapering upwards; content oleiferic. Hymenophoral trama with very abundant lactifers. Pileipellis a palisade; suprapellis 40–50 mm thick; subpellis 50–70 mm thick; terminal elements of suprapellis 20–40(–50) × 3–7 mm, cylindrical, 2- or 3-septate, obtuse, with granular content, slightly thick-walled; subpellis pseudoparenchymatous, composed of slightly thick-walled, isodiametric cells, 5–15(–20) mm. Stipitipellis with more slender, long terminal elements, 15–70 × 3–5 mm, cylindrical, obtuse, thin-walled. Clamp-connections absent. – Fig. 66
Ecology. Malagasy lowland rainforest.
Distribution. Only known from the type locality.
Revised specimens
Madagascar. Betsimisarka Prov., south of Maningory, 7 Dec. 1934, R. Heim G61 (PC, holotype).
Notes. I traced two exsiccata with the number G61 in PC. One of them (renumbered here as G61bis) consists of small, immature basidiomes, conserved in alcohol and representing most probably Lactarius annulatoangustifolius. The type of L. pseudovolemus consists of one half basidiome in very bad condition. As the material was not in a good condition I did not succeed in studying the lamellar edge and the hymenophoral trama.
Lactarius pseudovolemus differs from both other African Rugati by the tomentose pileipellis without veins, the smaller pileus, the subiculum and rooting stipe. A central amyloid spot on the plage is not observed in this species, whereas it occurs in Lactarius rubiginosus and L. kivuensis.
The two Tanzanian collections described by Karhula in Härkönen & al. (1995) as Lactarius aff. pseudovolemus were later described as a distinct species, L. volemoides.
43. Lactarius kivuensis Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 202 (1996). − Type: Democratic Republic of Congo, Kivu, Irangi, April 1972, J. Rammeloo Z310 (GENT, holotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 111–112 & Pl. 6–7 (1996d).
Pileus 20–78 mm diam., firm, regular, planoconvex to applanate, slightly depressed when young, irregularly waving and deeper depressed when older; margin first involute then slightly incurved to straight, undulate and crenulate when older, slightly grooved when old; pellis not dehiscent, mat, dry, concentrically wrinkled, with small veins which are already present on the margin of young specimens, dark brown (7E6) when young, then paler and more reddish (7D6–7), sometimes with ochraceous spots, especially in the centre. Lamellae adnate to slightly decurrent, unequal with lamellulae of 2 different lengths (1–3 lamellulae between 2 lamellae), rather distant (L+l = 4+4 to 5+3/cm), thick, brittle, yellowish white (3A2) to light yellow (4A4–6); edge concolorous, entire. Stipe 30–55 × 6–20 mm, cylindrical to subcylindrical, tapering downwards, sometimes slightly curved at the base, mat, dry, smooth but powdery at the base, light orange to greyish orange when young, especially near the apex paler when old, greyish white at the base, solid. Context rather firm, white, unchanging; smell not particular; taste mild. Latex abundant, watery, white, becoming brownish grey after a long period of drying; taste mild. Spore-deposit not observed. – Pl. 28
Chemical reactions. Context unchanging with anilated H2O, phenol, phenolaniline, NH4OH and TL4. Pileipellis ochraceous with NH4OH.
Basidiospores ellipsoid, sometimes elongate, 6.8–7.9–8.8–9.8 × 5.1–5.9–6.4–6.8(–7.0) mm (Q = 1.17–1.30–1.38–1.57, n = 100); ornamentation composed of irregularly shaped and sized warts, connected or aligned, seldom isolated, forming an incomplete to almost complete reticulum, up to 0.2 mm high; plage inamyloid or with central amyloid spot. Basidia 38–52 × 7–9 mm, cylindrical to subcylindrical, long and slender, 4-spored, hyaline; sterigmata 5–7 × 1–2(–3) mm. Pleurocystidia absent. Pleuropseudocystidia rather scarce, sometimes emergent, 3–5 mm diam., cylindrical to tortuous; content oleiferic. Lamellar edge sterile; marginal cells 15–20(–35) × 3–6 mm, cylindrical to subfusiform, long and slender, thin-walled, hyaline. Hymenophoral trama composed of rather small sphaerocytes and abundant lactifers. Pileipellis a palisade; suprapellis 30–80 mm thick; subpellis 40–60 mm thick; elements of the suprapellis cylindrical to subcylindrical, long and slender, 25–70 × 3–5(–6) mm, sometimes septate, thin-walled; subpellis composed of isodiametric cells, 5–15 mm diam., thin-walled. Stipitipellis a palisade; suprapellis 40–50 mm thick; subpellis 20–25 mm thick; elements of the suprapellis cylindrical to subcylindrical, 10–40 × 2–4 mm, thin-walled, septate, becoming longer towards the base of the stipe; slightly thick-walled elements present at the base. Clamp-connections absent. – Fig. 67
Ecology. Guineo‑Congolian rainforest.
Distribution. Only known from the type locality.
Revised specimens
Democratic Republic of Congo. Kivu Prov., near the river Luhoho, April 1972, J. Rammeloo Z233 (GENT) & Z310 (GENT, holotype), Z341, Z366 & Z416 (all GENT); Ibid., May 1972, J. Rammeloo Z439 & Z491 (all GENT).
Notes. The macroscopic description is based on Rammeloo Z233 & Z310, the microscopic description on all the specimens studied.
This species is very closely related to Lactarius rubiginosus from which it differs macroscopically by the darker pileus in young specimens, the less unicolorous pileus in older specimens and the latex which becomes greyish after drying. Microscopically, the species differs in spore-ornamentation and spore-dimensions. The stipitipellis in Lactarius kivuensis is a palisade with mainly thin-walled elements, whereas the stipitipellis of L. rubiginosus lacks the layer of isodiametric cells and consists mainly of thick-walled cells which become extremely long and more tortuous at the base of the stipe.
44. Lactarius rubiginosus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 207 (1996). – Type: Zambia, Chati‑forest, near Kitwe, 14 Jan. 1991, B. Buyck 3466 (BR 38488–76, holotype; PC, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 110–111 & Pl. 4–5 (1996d).
Pileus 40–110 mm diam., firm, planoconvex to applanate, slightly depressed in the centre; margin involute when young, then incurved to straight; pellis not dehiscent, dry, finely and completely wrinkled to veined, orange-red to brown. Lamellae subdecurrent to decurrent with a small tooth, unequal with lamellulae of different lengths (1–3 lamellulae between two lamellae, mostly very short), distant (L+l = 3 to 5/cm), broad (4–7 mm), rather thick, yellowish orange; edge entire, concolorous. Stipe (35–)55–82 × 16–22 mm, cylindrical, long, dry, paler than the pileus. Context very firm; taste mild; smell not remarkable. Latex abundant, white, watery; taste mild. Spore-deposit not observed. – Pl. 28
Chemical reactions. Context green with FeSO4.
Basidiospores ellipsoid, sometimes elongate, 8.2–9.1–9.4–10.4 × (5.6–)5.9–6.6–6.9–7.5 mm (Q = 1.24–1.36–1.38–1.55, n = 60); ornamentation amyloid, composed of rounded to elongated warts, aligned or connected by fine connective lines, forming an incomplete to almost complete reticulum, 0.1 mm high; plage sometimes with central amyloid spot. Basidia 55–65 × 7–9 mm, cylindric to subclavate, long and slender, 4-spored; sterigmata 4–7 × 1–2(–3) mm. Pleurocystidia absent. Pleuropseudocystidia rather abundant, seldom emergent, 3–5 mm diam., cylindrical to tortuous; content oleiferic to guttate. Lamellar edge sterile; marginal cells 15–25 × 3–5 mm, cylindrical, with rounded apex, sometimes septate, thin-walled, hyaline. Hymenophoral trama composed of sphaerocytes and abundant lactifers. Pileipellis a palisade; suprapellis 40–60 mm thick; subpellis 40–50 mm thick; terminal elements of the suprapellis 15–40(–60) × 3–4(–6) mm, cylindrical, long, slender, with rounded apex, often septate, thin-walled; subpellis composed of isodiametric cells, 5–10(–20) mm diam., thin-walled. Stipitipellis a trichoderm to lamprotrichoderm; suprapellis 50–80 mm thick; subpellis 10–20 mm thick; terminal elements (20–)50–100 × 4–6 mm, cylindrical, with rounded or tapering apex, often septate, thick-walled, becoming extremely long and more tortuous at the base of the stipe. Clamp-connections absent. – Fig. 68
Ecology. Zambezian miombo woodland.
Distribution. So far, only known from Tanzania and Zambia.
Revised specimens
Tanzania. Morogoro Prov., 23 Dec. 1992, E. Munyanziza 2-18 a,b,c (all WAG).
Zambia. Copperbelt Prov., Chati‑forest, near Kitwe, 14 Jan. 1991, B. Buyck 3466 (BR 38488–76, holotype; PC, isotype); Ibid., B. Buyck 3473 (PC); Western Prov., Mwinilunga, road to Solwezi, km 50, B. Buyck 3535 (PC); Luapala Prov., near Mansa, secundary school, 7 Dec. 1978, A.C. Rose 7813 (K(M) 38502); n.l., Feb. 1996, B. Buyck & G. Eyssartier 96-292b (PC).
Notes. The macroscopic description is based on the fieldnotes of Buyck; the microscopic description is based on Buyck 3466, 3473 & 3535.
45. Lactarius xerampelinus Karhula & Verbeken
Karhula & al., Karstenia 38: 59 (1998). − Type: Tanzania, Tabora distr., Lulanguru, 14 Dec. 1991, T. Saarimäki & al. 1116 (H, holotype; GENT, isotype).
Selected descriptions and icons. Karhula & al., Karstenia 38: fig. 15 (1998); Härkonen & al., Norrlinia 10: 93 (2003).
Pileus 50–120 mm diam., at first convex with central depression, then depressed to infundibuliform; margin in old specimens irregular, wavy; surface dark reddish brown, mat, at margin pruinose, in old specimens cracking and finely areolate. Lamellae shortly decurrent, subdistant with several lamellulae, fairly thick, up to 5 mm broad, pale cream; margin entire. Stipe 20–60 × 10–30 mm, variable in shape, cylindrical to tapering downwards, pale ochraceous, mat, weakly scrobiculate. Context in cap up to 13 mm thick, firm, solid in stipe, white, turning ochraceous when bruised; taste mild, pleasant; smell fresh, fruity. Latex white, abundant. Spore print pale cream. – Pl. 29
Spores ellipsoid, (7.1–)7.5–8.1–8.7–10 × 5.0–5.9–6.0–7.0 µm (Q = 1.29–1.35–1.48–1.67, n = 152) ; ornamentation amyloid, composed of very low (up to 0.2 µm high), mostly elongated warts and short ridges, sometimes aligned, often isolated; plage centrally amyloid. Basidia 60–70 × 7–9 µm, 4-spored. Pleurocystidia absent. Pleuropseudocystidia abundant, 3–5 µm diam. Lamellar edge sterile; marginal cells thin-walled, cylindrical to fusiform, often tapering upwards, 70–80 × 10 µm. Hymenophoral trama composed of sphaerocytes; lactifers 4 µm diam. Pileipellis a trichopalisade, composed of elongate, thin-walled cells forming chains; terminal elements 20–35 × 6–10 µm, clavate or cylindrical. Stipitipellis trichoderm-like, composed of repent, thin-walled hyphae, locally ascending. Clamp connections absent. – Fig. 69
Ecology. Zambezian miombo woodland.
Distribution. Known from Tanzania, Zambia and Zimbabwe.
Revised specimens
Tanzania. Western Prov., Tabora distr., Lulanguru, 14 Dec. 1991, T. Saarimäki & al. 1116 (H, holotype; GENT, isotype); Ibid., Urambo market, 13 Dec. 1991, T. Saarimäki & al. 1090 (H.)
Zambia. Miputu, 22 Jan. 1996, B. Buyck & G. Eyssartier 96-152 (PC); road Lusaka-Kapiri-Mposhi, bought from road stall, 4 Feb. 1996, B. Buyck & G. Eyssartier 96-34 (PC).
Zimbabwe. Mashonaland North, Hogwe Bush Camp, Omay C.L./Matusadona, 8 Feb. 1996, C. Sharp 418/96 (GENT); Midlands Prov., Mvuma, Lovedale Ranch, 29 Jan. 1998, C. Sharp 897-98 (GENT); Manicaland Prov. (Eastern Highlands), Stapleford J. Meikle, 4 Feb. 1999, A. De Kesel 2394 (BR 112564–44).
Notes. This species is locally eaten in the miombo area (e.g. Karhula & al. 1998, Verbeken & al. 2000).
Lactarius sect. Polysphaerophori Singer, Beih. Sydowia 7: 106 (1973).
Lactarius sect. Gymnocarpi R. Heim ex Verbeken, Mycotaxon 66: 374 (1998).
− Type species: Lactarius veraecrucis Singer.
Note. The placement of the African “Gymnocarpi” in L. sect. Polysphaerophori Singer, a section raised for South American species, was already suggested by Montoya et al. (2007) based on the similarity of L. luteopus and L. veraecrucis, the type species of this section.
Key to tropical African species
1. Pileus yellowish orange, orange, brownish orange, sometimes more ochraceous when older ....... 3
Pileus not orange or reddish, yellowish white, buff, brownish ........................................................ 2
2. Pileus pale, cream, yellowish white, tomentose; lamellae moderately distant; spore-ornamentation up to 1 µm high, subreticulate to reticulate ............................................................ 51. L. foetens
Pileus buff to brownish, strongly pruinose to granular; spore-ornamentation up to 0.2 µm high, consisting of warts which are often aligned, but never forming a reticulum ......... 52. L. brunnescens
3. Terminal elements of the pileipellis very thick-walled (up to 3 µm), often 10–15 mm broad; lamellae distant, decurrent with tooth ........ 46. L. gymnocarpus
Terminal elements less thick-walled, in general <10 µm broad; lamellae mostly crowded to moderately distant (if distant, then terminal elements of pileipellis < 8µm broad) ................. 4
4. Stipe white .............................................. 49. L. tanzanicus
Stipe yellow to orange or brownish orange, usually brightly coloured ............................................ 5
5. Spore ornamentation very low, at most 0.2 µm high, consisting of elongated warts which are not connected; plage not amyloid; stipe yellow to pale orange or brownish orange, with distinct pure white zone at top ......... 48. L. albocinctus
Spore ornamentation up to 0.5 µm high, subreticulate and with distinct centrally amyloid spot; stipe sometimes also with whitish zone on top .................................... 6
6. Spores ellipsoid, 7.3–8.4 ´ 5.5–6.0 mm; average Q = 1.20–1.50 ........................... 47. L. flammans
Spores elongate, 8.2–10.3 ´ 5.2–6.3 mm; average Q = 1.40–1.75 .................. 50. L. goossensiae
Species descriptionS
46. Lactarius gymnocarpus R. Heim ex Singer
Singer, Pap. Michigan Acad. Sci. 32: 107 (1946). – Lactarius gymnocarpus R. Heim, Boissiera 7: 273 (1943), nom. prov. – Type: Liberia, near Nengbe, 9 April 1939, G.W. Harley 56 (FH, lectotype).
Lactarius gymnocarpus R. Heim ex R. Heim, Bull. Jard. Bot. Etat Bxl 25: 24 (1955), illegit. – Type: not selected.
Misappl.: Lactarius goossensiae Beeli ss. Beeli (1928) pro parte.
Excl.: Lactarius gymnocarpus ss. Pegler (1977, = L. pseudogymnocarpus); L. gymnocarpus ss. Heim pro parte (1955, = L. pseudogymnocarpus), see also under notes.
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: Pl. 2, fig. 1b (1955); Verbeken, Mycotaxon 55: 520–523 (1995); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 113–115 & Pl. 8–13 (1996d); Thoen & Ba, New Phytol. 113: Pl. 1, fig. 2e (1989).
Pileus 65–80(–150) mm diam., quite thick, first planoconvex then planoconcave to slightly depressed and depressed; margin crenulate; pellis not dehiscent, dry, finely tomentose, with small and sometimes anastomosing veins, yellowish orange to golden yellow (5B7 to 4A7), ochraceous when older (5C7 to 5D7). Lamellae decurrent, extending in longitudinal veins on the stipe, unequal with lamellulae of different lengths, distant (L+l = 4/cm), medium broad (4–6 mm), thick, yellowish white to pale yellow (3A3), sometimes staining brown; edge concolorous. Stipe 40–45 × 6–14 mm, cylindrical, slightly tapering downwards, dry, finely tomentose, with longitudinal veins at the apex, concolorous, paler at the apex when young, darker when old, white tomentum at the base, solid and soon fistulous. Context firm, brittle, white, irregularly browning; taste mild to sweet; smell slightly unpleasant (“Valeriana-like” according to Heim, 1943). Latex rather abundant, watery to waxy, white, unchanging, mild, ochraceous brown on drying. Spore-deposit white. – Pl. 30
Chemical reactions. Context in pileus intense blue-green with FeSO4, in stipe darker (almost black) with FeSO4: immediately yellow with olive-shade with NH4OH; greyish brown with phenol 2%; unchanging with anilated H2O, phenolaniline, I2, NaOH and TL4. Lamellae very slowly reddening with anilated H2O, and phenolaniline, intense ochraceous with NaOH.
Basidiospores ellipsoid, 6.3–7.1–8.2–8.7 × 5.3–5.9–6.4–7.3 mm (Q = 1.14–1.21–1.29–1.42; n = 100); ornamentation amyloid, composed of irregular warts and fine connective lines, never forming a reticulum; some warts isolated, up to 0.5 mm high; plage sometimes amyloid in the centre. Basidia 45–55(–75) × 7–9 mm, cylindrical to slightly clavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia very numerous, emergent and distinct, 3–5(–7) mm diam., cylindrical, tapering upwards, sometimes constricted; content oleiferic with granules. Lamellar edge sterile; marginal cells 35–60 × 4–6 mm, cylindrical, slightly clavate or tapering upwards, often septate; wall thickened up to 0.5 mm. Hymenophoral trama composed of sphaerocytes; laticiferous hyphae present. Pileipellis a lampropalisade; elements of the suprapellis 20–100 × 5–15 mm, very irregularly shaped, varying from narrowly cylindrical to broadly fusiform or almost rounded, often tortuous, with thickened wall (1–3 mm); subpellis cellular; spherical cells (10–)15–30 mm, with slightly thickened wall. Stipitipellis a lampropalisade; elements of the suprapellis 30–120 × 10–25 mm, very irregularly shaped, often branched, thick-walled (3–5 mm); subpellis pseudoparenchymatous; spherical cells 10–25 mm diam., with slightly thickened wall (1 mm). Clamp-connections absent. – Fig. 70
Ecology. Collected in rainforest, lowland rainforest, dry coastal forest, riverine forest. The ectomycorrhizal relationship with Uapaca guineensis was fully established by Thoen & Ba (1989: 555, fig. 4b). The same authors considered Afzelia africana as a possible host tree. Anthonota crassifolia is listed as putative host tree in Thoen & Ducousso (1989).
Distribution. Widespread in Central and West Africa. Known from Cameroon, Democratic Republic of Congo, Gabon, Guinea, Ivory Coast, Liberia and Senegal. Also reported from Burkina Faso (Sanon & al. 1997, not revised).
Lactarius gymnocarpus is an edible species, eaten by local populations (e.g. Heim 1955b; Rammeloo, in herb.; Van Dijk & al., 2003).
Revised specimens
Cameroon. South Western Prov., Korup National Park, near Mundema, Jan. 1989, R. Watling 21470 & 21475 (all E); Ibid., 24 March 1991, R. Watling 24154 (E); Ibid., Ireba-Inene camp, 14 April 1997, M.E. Bechem K1183 (K(M) 91496); Ibid., trail to chimpanzee camp, 11 April 1997, P. Roberts K1074 (K(M) 914998); Ibid., trail from Mana Bridge to transect P, 15 Apr. 1997, P. Roberts K1166 (K(M) 50404); forest near Amugreta, Ebolowa, 16 Apr. 1946, R. Heim Q43 (PC).
Democratic Republic of Congo. Equateur Prov., Eala, June 1923, M. Goossens-Fontana 213 (BR 7712–49); Ibid. Jan. 1926, M. Goossens-Fontana 547 (BR 12326–07); Binga, 11 Dec. 1929, M. Goossens-Fontana 915 (BR 12314–92); Kivu Prov., Irangi, near river Luhoho, March 1972, J. Rammeloo Z201 (GENT); Ibid., Apr. 1972, J. Rammeloo Z286 (GENT); Ibid., May 1972, J. Rammeloo Z490 (GENT).
Gabon. Research Station of Ipassa-Makokou, 8 March 2005, M. Nguele 3 (BR 164550–38); Ibid., 13 March 2005, P. Mbazza 1 (BR 164540–28); Ibid., 16 March 2005, S. Dibaluka Mpulusu 26 (BR 164549–37); Ibid., border of Ivindo river, 21 March 2005, J. Degreef 308 (BR 164538–26).
Guinea. Fouta Djalon Prov., Saala waterfalls, north of Labbé, 4 July 1988, D. Thoen 7894 (BR 66457–12); forest of Upper Guinea, April 1939, R. Heim s.n. (PC); Nord de Tountourou, July 1988, D. Thoen 8066 (BR 66478–33); savane arborée de Lola, 3 Aug. 2000, A. Bâ 329 (GENT).
Ivory Coast. West Prov., Tiapleu, Tonkoui, Apr. 1934, R. Heim B30 (PC).
Liberia. Near Nengbe, April 1939, G.W. Harley 56 (FH, lectotype).
Senegal. Basse Casamance Prov., National Park of Basse Casamance, Aug. 1986, D. Thoen 7605 (BR 66435–87) & 7646 (BR 3160–56); Ibid., 22 Sep. 1986, D. Thoen s.n. (BR 66215–61); Ibid., 21 Sep. 1987, D. Thoen 7881 (BR 66454–09); Ibid., July 1988, D. Thoen 7996 (BR 66471–26).
Notes. The macroscopic description is based on Rammeloo Z286 and Thoen 7605 & 7646 and completed with the descriptions of Heim (1943) and Singer (1946); macrochemical characters are from Rammeloo Z286. The microscopic description is based on Thoen 7894, 7605 & 7646, Watling 24154 and Heim’s collections.
Lactarius gymnocarpus, invalidly described by Heim in 1943, was validated by Singer (1946) but later again by Heim (1955) with a superfluous Latin diagnosis and emended with data from the collections of Goossens-Fontana from Democratic Republic of Congo, which belong to Lactarius pseudogymnocarpus (Verbeken 1995).
The species is easily recognized by its pileipellis-structure: such thick-walled and very irregularly shaped terminal elements are only observed in Lactarius gymnocarpus and L. tanzanicus, but in the latter species they do not exceed 10 mm width.
The name Lactarius gymnocarpus was erroneously used by several authors: Buyck (1994b) and Buyck & Nzigidahera (1996) (= L. longisporus); Heim (1955) (= L. pseudogymnocarpus); Nzigidahera (1993) (based on Buyck 4380 = L. gymnocarpoides); Pegler (1977) (= L. pseudogymnocarpus); Pegler & Piearce (1980), Piearce (1981) (= Lactarius sp.). Citations of Lactarius gymnocarpus in Morris (1984, 1987, 1990), Chipompha (1994), Maghembe & Redhead (1980, in a pine plantation in Tanzania), and Sanon & al. (1997) are also considered as doubtful.
The photograph of Rammeloo Z490 was used for the stamp of "40k" (first issued 22 Jan. 1979, as Lactarius phlebonemus) in former Zaire (Democratic Republic of Congo).
47. Lactarius flammans Verbeken
Verbeken, Mycotaxon 55: 539 (1995). − Type: Zambia, Copperbelt Prov., near Kitwe, Mwekera‑Forest, 21 Jan. 1991, B. Buyck 3492 (BR 38486–74, holotype; PC, isotype).
Selected descriptions and icons. Verbeken, Mycotaxon 55: 537, 539–540 (1995), Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 116–117 & Pl. 14–15 (1996d); De Kesel & al., Guide des champignons comestibles du Bénin: 163 (2002).
Pileus 40–80 mm diam., firm, solid, planoconvex to applanate and depressed, irregularly wrinkled; pellis not dehiscent, wrinkling concentrically, mat, smooth, deep orange (5A7–8, 6A7–8 to 6B7–8), becoming paler in the centre when older; margin always distinctly yellowish to cream, which gives the pileus a two-zoned aspect. Lamellae decurrent, unequal with lamellulae (1 to 3 between 2 lamellae), distant (L+l = 3+2/cm), medium broad (2–4(–6) mm), pale. Stipe 20–35 × 10–20 mm, cylindrical to tapering downwards, firm, smooth, deep orange (5A7–8, 6A7–8 to 6B7–8), paler towards the base. Context white. Latex not observed. Spore-deposit white. – Pl. 30
Basidiospores ellipsoid, sometimes elongate, 6.5–7.3–8.4–9.3 × 4.9–5.5–6.0–6.5 mm (Q = 1.20–1.32–1.39–1.50; n = 60); ornamentation amyloid, forming an incomplete and irregular reticulum with distinct and sometimes also isolated warts; ridges irregular, some very fine; prominences low (< 0.5 mm high); plage sometimes faintly amyloid. Basidia 50–65 × 8–10 mm, cylindrical to slightly clavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia abundant, 4–8 mm, cylindrical to tortuous, obtuse, moniliform or irregular at the apex; content granular to oleiferic. Lamellar edge sterile; marginal cells 25–50 × 3–5 mm, cylindrical to fusiform, obtuse, often septate, with slightly thickened wall. Hymenophoral trama composed of sphaerocytes; laticiferous hyphae abundant. Pileipellis a lampropalisade; elements of the suprapellis 35–130 × 4–5(–7) mm, cylindrical to slightly tortuous, obtuse, sometimes septate, thick-walled (up to 1 mm); subpellis pseudoparenchymatous, spherical cells 5–15 mm, slightly thick-walled. Stipitipellis a lampropalisade; elements of the suprapellis 30–80 × 4–7 mm, cylindrical to tortuous, obtuse or tapering upwards, thick-walled (up to 1.5 mm); subpellis not distinct, pseudoparenchymatous. Clamp-connections absent. – Fig. 71
Ecology. Found in miombo woodland (under Brachystegia, Julbernardia) and Sudanian Isoberlinia (under Isoberlinia, Afzelia) dominated woodland.
Distribution. Known from Benin, Malawi and Zambia.
Revised specimens
Benin. Atacora Prov., Kounagnigou, 3 Sept. 1997, V. Antonín B97-151 (BR 101193–22); Atacora Prov., Koussou, 05 July 1998, A. De Kesel 2253 (BR 112761–47); Atacora Prov., Kouandé, 25 Sept. 2000, A. De Kesel 2936 (BR 129147–40); Atacora Prov., Tchétou, 02 Oct. 2000, A. De Kesel 2979 (BR 129191–84); Atacora Prov., Bassila, 04 Oct 2000, A. De Kesel 2999 (BR 129211–07); Borgou Prov., Wari Maro, 15 June 2000, A. De Kesel 2805 (BR 126358–64); Borgou Prov., Kpéssou-Samari, 23 June 2000, A. De Kesel 2860 (BR 126413–22).
Malawi. Northern Prov., Chinteche, 2 Jan. 1978, E. Tybaert 137 (GENT).
Zambia. Luapala Prov., near Kawambwa, 6 Jan. 1991, B. Buyck 3436 & 3437 (all PC); Copperbelt Prov., Mwekera-forest, near Kitwe, 21 Jan. 1991, B. Buyck 3492 (BR 38486–74, holotype; PC, isotype).
Notes. The macroscopic description is based on Buyck 3437, the microscopic description is based on Buyck 3437 & 3492.
Macroscopically, the species is well recognized by the strongly wrinkled and two-zoned pileus: yellow to cream at the margin and orange in the centre.
In the collections from Benin, some additional macroscopical characters, lacking in the original description are observed: the latex is white (De Kesel 2253), mild and abundant (De Kesel 2936 & 2999), with a nut-like, weakly aromatic taste (De Kesel 2253 & 2999). In one collection (De Kesel 2253) the context becomes slightly greyish.
48. Lactarius albocinctus Verbeken
Verbeken & al., Syst. Geogr. Pl. 70: 182 (2000). − Type: Zimbabwe, along Birchenough-Masvingo road, Chikuku area, near Mbuyanchanda school, leg. R. Walleyn 13 Feb. 1999, A. Verbeken 99-211 (GENT, holotype).
Selected descriptions and icons. Verbeken & al., Syst. Geogr. Pl. 70: 182–184 (2000).
Pileus 30–120 mm diam., convex and slightly depressed in the centre to almost applanate and undeeply depressed or V-shaped, becoming a bit irregular; margin first straight, then waving and irregularly widely crenulate; pellis dry, minutely felty, soft, dry, wrinkled and radially wrinkled at margin, usually not cracking, only in older specimens slightly cracked near the margin, orange (6A7, 6B7 to 6C7), locally greyish orange (5A4 to 5B4), brownish orange (6B8, 6C8, 5A8 to 5B8) in the center, paler outwards, light orange (5A6) near margin, locally orange white (5A2–3) at margin. Lamellae decurrent with tooth, very distant (L+l = 22/half a pileus, L+l = 5–6/cm halfway radius), with abundant lamellulae, thick but narrow, brittle, pure white, staining brown because of the latex; edge entire, concolorous. Stipe 20–60 × 13–32 mm, robust, shortly cylindric, a bit irregular, sometimes tapering; pellis smooth, glabrous to softly felty, dry, unicolorous, light yellow (4A4–5), pale orange (5A3), brownish orange (6B7 to 6C7), but pure white in the very well-defined zone of the decurrent teeth of the lamellae. Context thick and firm in pileus center but very thin near margin, firm and solid in the stipe, whitish, unchanging,; smell agreeable, aromatic, slightly Crustaceae-like; taste mild, agreeable. Latex rather abundant, white, changing brownish red to brown when drying, unchanging with KOH; taste mild. – Pl. 31
Chemical reactions. Context slowly staining blue or unchanging with gaiac, salmon with FeSO4.
Basidiospores broadly ellipsoid to ellipsoid, 7.7–8.5–8.6–9.7 × 5.6–6.4–6.5–7.2 µm (Q = 1.12–1.33–1.48, n = 40); ornamentation amyloid, composed of very low (< 0.2 µm) warts and ridges, forming a dense but incomplete and indistinct reticulum; plage not amyloid. Basidia 65–100 × 8–10 µm, narrowly cylindric, very long and narrow, 4-spored, with guttate content; sterigmata 5–12 × 1–2 µm. Pleurocystidia absent. Lamellar edge sterile; marginal cells 30–50 × 4–6 µm, subcylindric, mostly with tapering apex, slightly thick-walled. Hymenophoral trama mixed, composed of small globose cells and narrow, hyaline hyphae; lactifers not abundant. Pileipellis a lampropalisade, 120–150 µm thick; terminal elements 20–100 × 5–12 µm, some very long and narrowly cylindric, hair-shaped with a more or less acute apex, some much shorter, rather clavate or irregular, with rounded apex, with very thick walls and sometimes septate; subpellis cellular, composed of rather small globose cells of 5–10 µm diam. Stipitipellis a lamprotrichoderm; terminal elements mostly hair-shaped, also with very thick-walled but more irregularly shaped, often branching; subpellis composed of hyaline, multiseptate hyphae; globose cells or swollen elements almost lacking. Clamp-connections absent. – Fig. 72
Ecology. Found in drier Zambezian miombo woodland dominated by Brachystegia spiciformis on sandy soil.
Distribution. So far, only known from Zimbabwe.
Revised specimens
Zimbabwe. Midlands Prov., Mvuma, Mtao forest, 28 Jan. 1999, A. Verbeken 99-042 (GENT); Masvingo Prov., along Birchenough-Masvingo road, Chikuku area, near Mbuyanchanda school, leg. R. Walleyn 13 Feb. 1999, A. Verbeken 99-211 (GENT, holotype).
Notes. In the field this firm species is characterized by its orange colours, its pale distant gills which turn brown when bruised and a narrow but distinct whitish zone at the stipe apex.
Lactarius albocinctus is closely related to L. flammans Verbeken, from which it differs especially by the hardly wrinkled and uniformly coloured cap and the lower spore ornamentation. The pinkish reaction with FeSO4 and the hardly wrinkled cap are characters atypical for Lactarius section Polysphaerophori, but common in Lactarius section Rugati. As we consider the pileipellis structure the most important feature, we provisionally place Lactarius albocintus in L. section Polysphaerophori.
49. Lactarius tanzanicus Karhula & Verbeken
Karhula & al., Karstenia 38: 50 (1998). − Type: Tanzania, Kilwa distr., Mavuji, 18 Jan. 1993, T. Saarimäki & al. 1277 (H, holotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 115–116 & Pl. 16 (1996d); Karhula & al., Karstenia 38: 50–53, fig. 2 (1998); Härkonen & al., Norrlinia 10: 91 (2003).
Pileus 60–130 mm diam., convex to slightly depressed; edge wavy and uneven; pellis bright orange. Lamellae decurrent, cream with brown spots, fairly thick, 7 mm broad, spaced with several lamellulae (L+l = 5–8/cm). Stipe 30–80 × 15–30 mm, cylindrical, striate, white or cream, turning brown when bruised. Context white, hard, brittle; smell faintly like Russula xerampelina; taste mild. Latex white. Spore-deposit not observed. (According to Karhula & al. 1998). – Pl. 31
Basidiospores 6.9–7.4–8.0 × 5.2–5.8–6.3 mm, ellipsoid (Q = 1.19–1.28–1.37, n = 20); ornamentation very fine with small warts and few fine connective lines; plage inamyloid. Basidia 60–65 × 7–8 mm, cylindrical to slightly clavate, 4-spored, often thick-walled at the base. Pleuropseudocystidia not abundant, 3–5 mm, cylindrical, obtuse. Pleurocystidia absent. Lamellar edge sterile; marginal cells cylindrical and obtuse to tapering and almost pointed. Hymenophoral trama composed of sphaerocytes; laticiferous hyphae present. Pileipellis a lampropalisade; elements of the suprapellis 20–100 × 5–10 mm, clavate to irregularly cylindrical, some narrowly cylindrical, often branched, thick-walled (1–3 mm); subpellis pseudoparenchymatous with spherical cells of 10–25 mm diam. Stipitipellis a lampropalisade; elements of the suprapellis 30–45 × 8–10(–15) mm, broadly clavate to lanceolate and cylindrical, thick-walled; subpellis pseudoparenchymatous. Clamp-connections absent. – Fig. 73
Ecology. Miombo woodland, under Brachystegia spp.
Distribution. Only known from Tanzania.
Revised specimens
Tanzania. Marryi, 18 Jan. 1993, Saarimäki & al. 1277 (H, holotype); Morogoro Prov. Morogoro distr., 17 Dec. 1992, E. Munyanziza 1-19-20 (WAG); Ibid., 29 Dec. 1992, E. Munyanziza 3-19-20 (WAG).
Notes. This species differs from Lactarius gymnocarpus by the white or whitish stipe and the clearly denser lamellae. The elements from the suprapellis of the pileipellis are less irregular than those of Lactarius gymnocarpus, and somewhat narrower (up to 10 mm). Furthermore, the elements of the stipitipellis are up to 45 mm long, whereas in Lactarius gymnocarpus they can be 120 mm long.
Locally eaten in Tanzania (Härkonen & al. 2003).
50. Lactarius goossensiae Beeli
Beeli, Bull. Soc. Roy. Bot. Belg. 60: 165 (1928). – Type: Democratic Republic of Congo, Eala, July 1923, M. Goossens-Fontana 268 (BR 7713–50, lectotype).
Selected descriptions and icons. Verbeken, Mycotaxon 55: 540 (1995, as “sp. 1”); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 117–118 & Pl. 16–17 (1996d).
Pileus convex to infundibuliform, thin, brownish orange. Lamellae adnate to decurrent, cream. Stipe firm, yellowish orange. Context firm, taste acrid. Spore-deposit white. (According to Beeli 1928).
Basidiospores (7.9–)8.2–9.1–10.3 × 5.2–5.8–6.3, elongate to extremely elongate (Q = 1.38–1.57–1.75, n = 20); ornamentation an incomplete reticulum with distinct warts. Pleurocystidia absent. Pleuropseudocystidia rather abundant, very emergent, cylindrical to flexuous, obtuse to tapering, 7–15 mm diam. Hymenophoral trama composed of sphaerocytes; laticiferous hyphae present. Pileipellis a lampropalisade; elements of the suprapellis 10–45 × 4–6 mm, narrowly cylindrical, thick-walled (up to 1 mm); subpellis pseudoparenchymatous. Stipitipellis a lampropalisade; elements of the suprapellis 10–45 × 4–6 mm, narrowly cylindrical, thick-walled (up to 1 mm); subpellis pseudoparenchymatous. Clamp-connections absent. – Fig. 74
Ecology. Found in swamp forest and woodland with Uapaca somon.
Distribution. Known from the Democratic Republic of Congo and Guinea.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Eala, July 1923, M. Goossens-Fontana 268 (BR 7713–50, lectotype).
Guinea. Savanne arborée de Lola, 3 Aug. 2000, A. Bâ 320 (GENT)
Notes. Heim (1955) mentioned that Goossens-Fontana 213 and Goossens-Fontana 268 both described by Beeli as Lactarius goossensiae, probably represent two different species. Goossens-Fontana 213 represents indeed Lactarius gymnocarpus (so does Goossens-Fontana 547, which is described by Beeli (1928) as "probably L. goossensiae"). Therefore, Goossens-Fontana 268 was selected as lectotype for L. goossensiae (Verbeken 1996). The species differs from Lactarius gymnocarpus by the absence of the short, broad and irregularly shaped elements in the pileipellis, by the absence of the browning reaction of the lamellae and by the longer and more elongate spores.
51. Lactarius foetens Verbeken & Van Rooij
Van Rooij & al., Nova Hedwigia 77: 230 (2003). − Type: Benin, Atacora Prov., Bassila, 21 June 2000, A. De Kesel 2840 (BR 126393–02, holotype; GENT, isotype).
Selected descriptions and icons. Van Rooij & al., Nova Hedwigia 77: 230–231, 242 (2003).
Pileus 65 mm diam., planoconvex to distinctly umbilicate; pellis strongly wrinkled, mat and felty, yellowish white (3A2 to 4A2); margin slightly involute and wrinkled. Stipe 29 × 11–16 mm, cylindric, slender, slightly tapering upwards; pellis tomentose, whitish but staining brownish (5D4–7). Lamellae broadly adnate to emarginate with decurrent tooth, sometimes forked at the margin of the pileus or anastomosing, distant (L+l = 3/cm) and thin (to 6 mm); lamellulae with regular pattern (1–3/L), white to cream (4A3), turning dark brownish to greyish when dried; edge entire and concolorous. Context very thin in the pileus, marrow-like and solid in the stipe, cream to pale yellow, turning brownish; taste very agreeable, reminding of Boletus edulis but stronger; smell weak and unremarkable when fresh, very unpleasant and strongly fish-like when dried. Latex moderate, not very liquid, white. Spore-deposit not observed. – Pl. 32
Basidiospores ellipsoid, sometimes subglobose or elongate (6.0–)6.1–7.1–7.2–8.2(–8.4) × 4.8–5.6–5.8–6.5 µm (Q = 1.07–1.24–1.31–1.46, n = 120); ornamentation amyloid, composed of irregular, subspherical to subconical warts, up to 1 µm high, aligned and connected by fine connective lines, forming a distinct and incomplete to complete reticulum, warts seldom isolated; plage often centrally to almost totally amyloid. Basidia 55–73 × 7–10 µm, subclavate, with oil-like content, thin-walled, 4-spored (seldom 2-spored). Pleurocystidia absent. Pleuropseudocystidia 10–15 µm diam., regularly cylindric with rounded, sometimes subcapitate apex; content needle-like and granular, sometimes with oil-like droplets. Lamellar edge sterile; marginal cells 15–23 × 2–5 µm, subclavate to irregularly cylindric, sometimes branched, thin-walled, hyaline, sometimes with oil-like droplets. Hymenophoral trama cellular, with abundant lactifers. Pileipellis a lampropalisade, composed of a distinct pseudoparenchymatic layer (subpellis) covered with distinctly developed tufts of hair-shaped thick-walled elements (discontinuous suprapellis); elements of suprapellis 25–125(–170) × 3–5 µm, cylindric, hair-shaped, sometimes tapering towards the apex, septate, with thickened walls (0.5–1 µm); subpellis pseudoparenchymatous, with spherical cells (10–)15–25(–30)µm, sometimes with thickened walls. Stipitipellis idem, without developed pseudoparenchymatic layer, terminal elements usually longer and thin-walled (<0.5 µm). Clamp-connections absent. – Fig. 75
Ecology. Found in the Guineo-sudanian transition zone, exclusively in forest galeries with Berlinia grandiflora and Uapaca somon.
Distribution. Known from Benin and Togo.
Revised specimens.
Benin. Donga Prov., Bassila, 21 June 2000, A. De Kesel 2840 (BR 126393–02, holotype; GENT, isotype); Ibid. 28 Jun. 2002, A. De Kesel 3486 (BR 152042–43); Ibid. 7 Oct. 2002, A. De Kesel 3526 (BR 152007–08); Atacora Prov., Kota falls, 18 Jun. 2004, A. De Kesel 3688 (BR 157117–74).
Togo. Central Prov., Forêt Classée d’Alédjo, 11 Jul. 2007, A. De Kesel 4283 (BR 163675–36).
52. Lactarius brunnescens Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 199 (1996). − Type: Burundi, Nkayamba hill, 15 March 1994, A. Verbeken 94-159 (BR 57613–92, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 131–133 & Pl. 52–53 (1996d).
Pileus 60–150 mm diam., firm, fleshy, convex to applanate and slightly depressed; margin first incurved, then straight, slightly crenulate to irregular, not striate; pellis not dehiscent, dry, mat, slightly felty, sometimes pruinose, strongly wrinkling towards the margin, buff (4B3–4), yellowish brown (5E6–8 to 5F6–8), brownish orange (6C4–7) to dark brown (6E6–7), darker when bruised. Lamellae adnate to slightly decurrent, unequal with lamellulae of different lengths (3–5 lamellae between 2 lamellae), very crowded (L+l = 4+13/cm), thin, narrow (2–3 mm), brittle, sometimes with anastomosing veins, whitish to cream, dark brown when bruised; edge entire, concolorous. Stipe 25–50 × 18–22 mm, subcylindrical to cylindrical, sometimes tapering downwards, dry, slightly felty, concolorous, paler towards the apex, firm, solid. Context firm, whitish to cream, browning when cut or when bruised; taste mild; smell disagreeable, sometimes slightly fish-like. Latex very abundant (not in A. Verbeken 94-159), white to greyish orange (5B5), changing to dark brown (8F8); taste mild. Spore-deposit whitish to cream. – Pl. 33 & 34
Chemical reactions. FeSO4: strongly green to blue-green on pellis, nihil to very faint on context.
Basidiospores ellipsoid, 7.2–8.2–8.5–9.0 × 5.9–6.8–7.2–7.9 mm (Q = 1.12–1.17–1.22–1.32; n = 80); ornamentation amyloid, weak, composed of very low (< 0.2 mm high) and small irregular warts, aligned or connected by fine connective lines; plage sometimes faintly amyloid in the centre. Basidia 45–52 × 7–9 mm, cylindrical to subcylindrical, 4-spored; content guttate. Pleurocystidia absent. Pleuropseudocystidia rather scarce, 4–5 mm diam., cylindrical, rounded at the apex, emergent up to 15 mm; content oleiferic. Lamellar edge sterile; marginal cells 25–55 × 4–6 mm, cylindrical, rounded at the apex, septate, sometimes branched; wall slightly or not thickened. Hymenophoral trama irregular, composed of densely interwoven hyaline hyphae of 3–6 mm diam.; lactifers present. Pileipellis a lampropalisade, locally a lamprotrichoderm with inflated elements at the base; elements of the suprapellis 25–80 × 4–7 mm, cylindrical, with rounded apex, septate, sometimes slightly incrustated, with slightly thickened wall, sometimes originating from isodiametric or irregular cells; subpellis pseudoparenchymatous, not well developed. Stipitipellis a trichoderm to lamprotrichoderm; elements of the upper layer 15–40 × 4–7 mm, cylindrical, rounded at the apex, septate, often heavily incrustated, wall slightly thickened. Clamp-connections absent. – Fig. 76
Ecology. Zambezian miombo woodland. Found under Brachystegia spp., Julbernardia globiflora, Uapaca kirkiana, Marquesia macroura.
Distribution. Known from Burundi, Malawi, Zambia and Zimbabwe.
Revised specimens
Burundi. Bururi Prov., near Rumonge, Nkayamba hill, 22 Feb. 1993, B. Buyck 4956 (PC); Ibid., 15 March 1994, A. Verbeken 94-159 (BR 57613–92, holotype; GENT, isotype); Cankuzo Prov., Murango, 3 March 1994, B. Nzigidahera 63 (PC).
Malawi. Southern Prov., Zomba, Feb. 1991, B. Morris 91-116 (PC); Northern Prov., Mzimba distr., Perekezi Forest Reserve, 28 Jan. 2005, A. Verbeken 05-083 (GENT); Ibid., 28 Janv. 2005, F. Noe 05/09 (GENT); Ibid., 29 Jan. 2005, A. Verbeken 05-133 & 05-138 (all GENT); Ibid., 30 Jan. 2005, A. Verbeken 05-172 (GENT); Ibid., 31 Jan. 2005, A. Verbeken 05-180 & 05-182 (all GENT).
Zambia. Luapala Prov., Mansa, Kabunda mission, 3 Jan. 1991, B. Buyck 3345, 3346 & 3350 (all PC); near Kawambwa, 6 Jan. 1991, B. Buyck 3432 & 3433 (all PC); near Mansa, secundary school, 6 Dec. 1978, A.C. Rose 7805 (K(M) 38501); Northern Prov., Mpika distr., North Luangwa National Park, 10 Jan. 1995, D. Shah‑Smith 98 (K(M) 35503); Copperbelt Prov., Kitwe State Ranch, 6 Jan. 1982, G. Piearce FP708/7 (K(M) 38497).
Zimbabwe. Manicaland Prov., Eastern Highlands, Mguzu, 5 Feb. 1999, Gansemans in A. De Kesel 2741 (BR 115759–38).
Notes. The description is based on Verbeken 94-158 & 94-159 and Buyck 3432.
Sect. Phlebonemi R. Heim ex Verbeken
Lactarius sect. Phlebomeni R. Heim ex Verbeken, Mycotaxon 66: 378 (1998).
Lactarius subsect. Luteoli Pacioni & Lalli, Mycotaxon 44: 190 (1992).
− Type species: Lactarius phlebonemus R. Heim & Gooss.-Font.
Notes. The distinction between this section and L. sect. Polysphaerophori is artificially based on spore characteristics but maintained for practical reasons. Species with capitate terminal pileipellis elements, in Africa only represented by L. nonpiscis, might form a natural group (L. subsect. Luteoli).
Key to tropical African species
1. Elements of the pileipellis capitate, spherical at the apex ..................................... 57. L. nonpiscis
Elements of the pileipellis not capitate ................................................................................................. 2
2. Pileus zonate, brownish ochraceous, then becoming paler to cream-coloured 56. L. pisciodorus
Pileus never zonate, never becoming very pale ................................................................................. 3
3. Pleurocystidia present and abundant ............................................................................ 55. L. arsenei
Pleurocystidia absent .............................................................................................................................. 4
4. Pileus with remarkable gelified sinuose veins, very strongly wrinkling, yellow ochre with orange centre to brownish ochre or dark brown; latex changing ochre to light brown, then dark brown; connective lines in the spore-ornamentation rare, although present in almost every spore; terminal elements of the pileipellis strongly thick-walled (walls up to 1 µm thick) .................. 53. L. phlebonemus
Pileus without veins, almost smooth, felty, vivid ochraceous orange; latex white, unchanging, context changing brown; connective lines in the spore-ornamentation very rare, present in only a few spores; terminal elements of the pileipellis only slighty thick-walled (walls up to 0.5 µm thick) ............. 54. L. angustus
Species descriptionS
53. Lactarius phlebonemus R. Heim & Gooss.‑Font.
Heim, Bull. Jard. Bot. Etat Bxl 25: 38 (1955). – Type: Democratic Republic of Congo, Binga, Aug. 1942, M. Goossens-Fontana 2044 (BR 7727–64, lectotype).
Invalidly described infraspecific forms without taxonomic significance: f. brunneus a R. Heim, Ibid.: 39; f. brunneus ß R. Heim, Ibid.: 40; f. ocraceus R. Heim, Ibid.: 43 (see also notes).
Excl.: Lactarius phlebonemus ss. Morris (1990) and Pegler & Young (1982), = L. tenellus.
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: Pl. 3, fig. 2ab & Pl. 4, fig. 2 (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 14, fig. 3–4 (1955); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 130–131 & Pl. 45–51 (1996d); Verbeken & al., Syst. Geogr. Pl. 70: 199–201 (2000).
Pileus 130 mm diam., very irregularly shaped, slightly depressed in the center; pellis smooth, chamois-leather-like, concentrically wrinkled at the extreme margin, not unicolorous, greyish brown to light brown (6DE3–4), locally brownish orange (6C3) or greyish orange (5B3); margin irregularly waving. Lamellae decurrent with tooth, moderately distant (L+l = 8/cm halfway radius), very thick, 5 mm broad, with abundant lamellulae of many different lengths (no regular pattern, but with abundant very short ones), cream, yellowish white (3A2); edge pale brownish. Stipe 70 × 55 mm, very irregular; pellis chamois-leather-like and smooth, concolorous with pileus. Context extremely thick-fleshy, 15 mm thick and rather firm in pileus, solid in stipe, whitish but staining pale brown (5D4); taste mild, a bit rancid, reminding of nuts, not very agreeable; smell chemical, like paint. Latex abundant, watery and pale brown right from the beginning, like half coffee, half milk or pale mokka, later (after drying) chocolate brown, unchanging with KOH, pale brown on white tissue. Spore-deposit not observed. – Pl. 35
Chemical reactions. Context immediately blue with gaiac, immediately blue-green with FeSO4.
Basidiospores ellipsoid, (6.0–)6.7–7.2–7.8–8.4(–9.2) × 5.1–5.9–6.4–7.4 mm (Q = 1.12–1.21–1.23–1.30; n = 90); ornamentation amyloid, composed of conical to subconical warts, 0.5 mm high, isolated or sometimes connected by fine connective lines; plage inamyloid. Basidia 45–52 × 7–10 mm, cylindrical to subcylindrical, 4-spored. Pleurocystidia absent. Pleuropseudocystidia very abundant, 3–5 mm diam., cylindrical to irregularly tortuous or tapering downwards, with rounded apex, emergent up to 20 mm; content oleiferic. Lamellar edge sterile; marginal cells 30–70 × 4–6 mm, cylindrical, apex rounded, sometimes tapering and fusiform, sometimes remarkably swollen (up to 9 mm). Hymenophoral trama irregular; composed of interwoven, hyaline hyphae, 3–5 mm diam.; lactifers present. Pileipellis a lampropalisade; elements of suprapellis 15–100 × 5–10 mm, cylindrical, hair-shaped, septate; wall thickened (1 mm); subpellis pseudoparenchymatous, isodiametric cells 10–40 mm, sometimes slightly thick-walled. Stipitipellis a lampropalisade, similar to pileipellis but elements of suprapellis shorter, up to 50 mm long, and subpellis sometimes more rudimentary. Clamp-connections absent. – Fig. 77
Ecology. Found in lowland rainforest, drier Guineo‑Congolian rainforest and drier Zambezian miombo woodland.
Distribution. Known from Cameroon, Democratic Republic of Congo and Zimbabwe.
Revised specimens
Cameroon. South Western Prov., Korup National Park, near Mundema, transect P19, 26 March 1991, R. Watling 24172 (E); Ibid., transect P13, 24 March 1991, R. Watling 25220 (E).
Democratic Republic of Congo. Equateur region, Binga, Dec. 1931, M. Goossens-Fontana 956a (BR 7722–59) & 957a (BR 7723–60); Ibid., Aug 1942, M. Goossens-Fontana 2044 (BR 7727–64, lectotype).
Zimbabwe. Midlands Prov., Mvuma, Central Estates, hill northeast of Beacon Hill ridge, leg. R. Walleyn 6 Feb. 1999, A. Verbeken 99-133 (GENT).
Notes. The macroscopical description given here is based on the Zimbabwean collection. The microscopical description is based on an extended review of the collections of Goossens-Fontana, completed with some data of the more recently collected material.
The diagnosis of this species was based on three collections, provisionally named “f. ochraceus”, “f. brunneus alpha” and “f. brunneus beta”) (Heim l.c.). One of these collections, Goossens-Fontana 956 (“f. ochraceus”) has been erroneously indicated as holotype by Verbeken (1998a). It is in a rather poor condition and consists of three incomplete basidiomes but only one of them represents L. phlebonemus (now labelled as Goossens-Fontana 956a). Considering the bad condition of collection Goossens-Fontana 956, it is not possible to verify if the coloured plates of this collection, which show pale basidiocarps (Heim 1955a & 1955b, as “f. ochraceus”), represent Lactarius phlebonemus or the second species mixed with it. Both other orignal collections, Goossens-Fontana 957 and 2044, as well as the material from Zimbabwe and Cameroon consist of specimens with dark brown cap and stipe, making it likely that the ochraceous colours described for “f. ochraceus” are based on the species erroneously mixed with Goossens-Fontana 956. To avoid further confusion Goossens-Fontana 2044 was proposed as lectotype (Verbeken & al. 2000).
54. Lactarius angustus R. Heim & Gooss.‑Font.
Heim, Bull. Jard. Bot. Etat Bxl 25: 67 (1955). – Type: Democratic Republic of Congo, Binga, April 1928, M. Goossens-Fontana 713 (BR 7716–53, holotype).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 67-69 & Pl. 3, fig. 3a–b (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 15, fig. 7 (1955); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 133 & Pl. 55–56 (1996d).
Pileus 50–80 mm diam., convex to applanate, then broadly depressed in the centre, irregularly curved; margin finely incurved, irregularly crenulate; pellis almost smooth, very finely felty, felty when dry, unicolorous, vivid ochraceous orange. Lamellae almost free when young, decurrent when older, very dense, white, brittle. Stipe 40–65 × 10–18 mm, cylindrical, subbulbous, concolorous with the pileus, firm, solid. Context granulose, white, becoming brown when exposed; taste acrid and bitter; smell nauseous. Latex very abundant, white; taste "acrid". Spore-deposit white. (According to Heim 1955a). – Pl. 36
Basidiospores ellipsoid, sometimes subglobose, 6.5–7.6–8.6(–9.0) × 5.2–6.3–7.4 mm (Q = 1.07–1.22–1.48, n = 30); ornamentation amyloid, composed of subspherical to cylindrical rounded warts, up to 0.75 mm high, isolated, very rarely connected by a fine line; plage inamyloid. Basidia 38–53 × 6–8 mm, cylindrical, 4-spored. Pleurocystidia absent. Pleuropseudocystidia abundant, 4–6 mm diam., cylindrical to tortuous, rounded at the apex, sometimes branched; content oleiferic and guttate. Lamellar edge sterile; marginal cells 12–42 × 3–6(–8) mm, irregularly cylindrical to fusiform, often septate, sometimes branched, hyaline, thin-walled, sometimes slightly thick-walled. Hymenophoral trama irregular, composed of hyaline hyphae and abundant lactifers. Pileipellis a lampropalisade; terminal elements 10–50 × 5–8(–10) mm, short, subclavate to cylindrical, rounded at the apex, often septate; wall slightly thickened to thickened (up to 0,5 mm); subpellis pseudoparenchymatous, isodiametric cells 10–20 mm. Stipitipellis as pileipellis but with longer and more irregular terminal elements which have up to 1,5 mm thickened walls. Clamp-connections absent. – Fig. 78
Ecology. Collected in drier Guineo‑Congolian rainforest with Gilbertiodendron dewevrei.
Distribution. Only known from the type locality.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Binga, April 1928, M. Goossens-Fontana 713 (BR 7716–53, holotype).
Notes. The type consists of about 15 basidiomes, macroscopically in rather good condition; for the microscopic study however it is difficult to make the collapsed elements inflate.
Heim, Candollea 7: 380 (1938, as "arseni"). – Type: Madagascar, West of Ivoloina, road to Ambatondrazaka, West of Tamatave, 10 July 1937, R. Heim s.n. (PC, holotype).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 61–62 (1938); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 135–136 & Pl. 60–63 (1996d).
Pileus 40 mm diam., thick, convex to applanate, then slightly depressed; margin involute; pellis not dehiscent, finely velvety, brownish ochraceous. Lamellae adnate, unequal with numerous lamellulae, crowded, narrow (up to 1.5 mm), thick, cream; edge fimbriate. Stipe 30 × 7 mm, cylindrical to fusiform, concolorous with the pileus, spotted, solid. Context firm, brittle, white, ochraceous orange when conserved in alcohol; smell fish-like; taste mild. Latex abundant, white, unchanging; taste mild. Spore-deposit white (?). (According to Heim 1938b).
Basidiospores subglobose to ellipsoid, (6.6–)7.2–7.7–8.2(–8.6) × (5.8–)6.1–6.6–7.2(–7.7) mm (Q = 1.08–1.17–1.26, n = 30); ornamentation amyloid, composed of conical, obtuse or pointed warts, mostly isolated, sometimes connected by fine connective lines; plage inamyloid. Basidia 45–55 × 9–13 mm, subclavate to subventricose, 4-spored; content slightly granular. Pleurocystidia abundant, not emergent, 30–50 × 5–10 mm, cylindrical to subclavate, 2- or 3-septate, with rounded apex; wall slightly thickened. Pleuropseudocystidia abundant, 5–7 mm diam, cylindrical to tortuous; content oleiferic. Lamellar edge sterile; marginal cells 30–50 × 4–7(–10) mm, cylindrical, subclavate or subfusiform, 2- or 3-septate; wall slightly thickened. Hymenophoral trama heteromerous; composed of sphaerocytes and narrow, subparallel, hyaline hyphae; lactifers abundant. Pileipellis a lampropalisade; terminal elements in suprapellis cylindrical to subcylindrical, 20–90 × 5–10 mm, septate, thick-walled with granular content; subpellis pseudoparenchymatous, composed of isodiametric to irregular cells which are 10–30 mm, with slightly thickened wall; with a layer of brownish extracellular incrustations in the lower part of the subpellis. Stipitipellis similar; cylindrical terminal elements more numerous. Clamp-connections absent. – Fig. 79
Ecology. Collected in Malagasy lowland rainforest.
Distribution. Only known from the type locality.
Revised specimens
Madagascar. Betsimisaraka Prov., W of Ivoloina, road to Ambatondrazaka, W of Tamatave, leg. Arsène Botozanany 10 July 1937, R. Heim s.n. (PC, holotype).
Notes. The type-specimen consists of one basidiome in bad condition.
56. Lactarius pisciodorus R. Heim
Heim, Candollea 7: 380 (1938). – Type: Madagascar, Nosy‑Mangabe (Ile Marosse), 30 Dec. 1934, R. Heim J52 (PC, holotype).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 58–60 & Pl. 1e (1938); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 136 & Pl. 64 (1996d).
Pileus 30–40 mm diam., rather thick, soon depressed, subinfundibuliform; margin slightly grooved, involute, then straight; pellis not dehiscent, smooth, finely felty, zonate with some darker circular lines, brownish ochraceous, then becoming paler, cream to orange, darker in the centre. Lamellae subdecurrent, unequal with lamellulae of different lengths, crowded, very narrow (up to 1.5 mm), rather thick, cream; edge crenulate, concolorous. Stipe 35–40 × 7–10 mm, fusiform, long, robust, finely felty, not hirsute at the base, pinkish ochraceous or straw, more pinkish towards the base, solid. Context firm, thick, white, unchanging, brownish orange when conserved in alcohol; smell fish-like; taste acrid. Latex abundant in the pileus, not abundant in the stipe, white; taste acrid. Spore-deposit not observed. (According to Heim 1938b). – Pl. 36
Chemical reactions. FeSO4: pinkish lilac. Gaiac: positive, intense. Gaiacol: positive, intense. Pyramidon: lilac. (According to Heim 1938b).
Basidiospores subglobose to ellipsoid, 6.8–8.3 × 6.8–7.7 mm; ornamentation composed of rounded to irregularly shaped warts, up to 0.4 mm high, sometimes aligned, sometimes connected, mostly isolated; plage inamyloid. Basidia 52–55 × 8–9 mm, long, fusiform, 4-spored. Cystidia or pseudocystidia not observed. Pileipellis a trichoderm, composed of undulating, varicose, arising terminal elements, 2–5 mm diam., hyaline, rarely septate, tapering upwards, with slightly thickened walls (up to 0.7 mm). Stipitipellis idem. Clamp-connections absent. – Fig. 80
Ecology. Malagasy lowland rainforest.
Distribution. Only known from the type locality.
Revised specimens.
Madagascar. Nosy-Mangabé (Ile Marosse), 30 Dec. 1934, R. Heim J52 (PC, holotype).
Notes. The macroscopic and microscopic description are based completely on the notes from Heim. The type-specimen consists of one quarter of a basidiome and is in very bad condition. A few spores could be observed, but most of them had collapsed. The basidia I noticed were shrunk and very small, but there are pseudocystidia present in the hymenium. Pileipellis and stipitipellis could not be observed.
57. Lactarius nonpiscis Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 204 (1996). – Type: Zambia, Chati‑forest, near Kitwe, 18 Dec. 1990, B. Buyck 3171 (GENT, holotype; PC, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 133–134 & Pl. 57–59 (1996d); Verbeken & al., Syst. Geogr. Pl. 70: 196–198 (2000).
Pileus 13–26 mm diam., firm, convex to applanate with papilla or slightly depressed, mostly asymmetric, sometimes slightly irregular; margin undulate to crenulate; pellis strongly wrinkled, with small veins, very rugulose, mat, felty, with an almost pruinose aspect, cream, yellowish white to pale yellow (3A2–4A2–3), sometimes greyish orange (5B3–5), brownish orange (5C4–5), staining purplish brown. Lamellae narrowly adnate to slightly decurrent, unequal with 1–7 lamellulae between 2 lamellae, rather distant (L+l = 7–10/cm), thin, rather brittle, paper-like, narrow, 2 mm broad, yellowish white (3A2) to orange grey (5B2–3) with a weak lilac tinge in it, concolorous with stipe; edge entire, concolorous or a bit more yellow-orange, soon staining brownish. Stipe 20–40 × 3–7(–10) mm, irregularly subcylindrical, sometimes curved near the base, sometimes a bit broader near the base and tapering upwards, slightly eccentric, locally applanate or indented, cream or orange-grey (5B2–3 to 5C2–3), finely felty, greyish and strongly felty near the base, staining purplish brown, soon spotted when touched (darker brownish, blackish violet spots), chambered to fistulous. Context white, dirty pale brown to pale brown; taste mild, agreeable, nut-like; smell first chemical then fish-like, distinctly fish-like, even in young and very fresh basidiomes. Latex moderately abundant, watery white, staining purplish brown to blackish violet in contact with flesh, unchanging on white tissue, unchanging with KOH; taste mild. Spore-deposit not observed. – Pl. 36
Chemical reactions. Context staining purplish brown, dark blue green with gaiac, dark blue-grey with FeSO4, unchanging with KOH.
Basidiospores ellipsoid, 8.0–8.7–9.2–10.0 × 6.1–6.6–6.8–7.3 mm (Q = 1.21–1.31–1.36–1.49; n = 60); ornamentation amyloid, composed of warts, up to 1,5 mm high, conical, slightly subspherical to truncate, mainly isolated, seldom aligned or connected; plage mostly inamyloid, sometimes slightly amyloid in the centre. Basidia 40–55 × 9–11 mm, cylindrical to subclavate, 4-spored; content guttate. Pleurocystidia absent. Pleuropseudocystidia very abundant, cylindrical to tortuous, seldom branched, 4–5(–6) mm diam.; content oleiferic to guttate. Lamellar edge sterile; marginal cells 30–40(–60) × 4–8 mm, cylindrical, subfusiform, 2–3(4)-septate, slightly thick-walled. Hymenophoral trama irregular, composed of thin-walled hyaline hyphae and abundant lactifers. Pileipellis a palisade to a lampropalisade; elements of suprapellis 40–80(–100) × 4–7 mm, long, slender, septate, slightly thick-walled, very often capitate; subpellis composed of isodiametric cells, 10–30 mm, thin-walled. Stipitipellis a trichoderm to lamprotrichoderm, composed of interwoven, hyaline hyphae and ascending slightly thick-walled hyphae with the same capitate apex as in the pileipellis; ascending hyphae becoming very long and more thick-walled towards the base of the stipe. Clamp-connections absent. – Fig. 81
Ecology. Zambezian miombo woodland, found under Brachystegia, Uapaca and Isoberlinia.
Distribution. Known from Malawi, Zambia and Zimbabwe.
Revised specimens
Malawi. Northern Prov., Mzimba distr., Ntanga Forest Reserve, 30 Jan. 2005, A. Verbeken 05-169 (GENT).
Zambia. Copperbelt Prov., Chati‑forest, near Kitwe, 18 Dec. 1990, B. Buyck 3171 (GENT, holotype; PC, isotype); Luapala Prov., Samfya, 28 Dec. 1990, B. Buyck 3265bis (PC); Ibid., 29 Dec. 1990, B. Buyck 3266bis, 3267 & 3268 (all PC); Mansa, Kabunda mission, 31 Dec. 1990, B. Buyck 3305 & 3306 (all PC).
Zimbabwe. Midlands Prov., Mvuma, Central Estates, Beacon Hill section, 26 Jan. 1999, A. Verbeken 99-003 (GENT); Mvuma, Central Estates, along road to Kwekwe, 15 Feb. 1999, A. Verbeken 99-218 (GENT); Mashonaland West Prov., on top of the Great Dyke, just off the highway from Harare to Lake Kariba, 4 Feb. 1999, D. Arora 99-78 (SFSU).
Notes. This striking species is well characterized by the remarkably capitate elements in its pilei-pellis, combined with the small and pale, purplish brown staining basidiomes with a fishy smell.
Lactarius sect. Chromospermi Verbeken, Mycotaxon 77: 439 (2001).
− Type species: Lactarius chromospermus Pegler.
Species description
58. Lactarius chromospermus Pegler
Pegler, Kew Bull. 37: 269 (1982). – Type: : Zambia, Luapala Prov., Mansa, 7 Dec. 1978, A.C. Rose 7811 (K(M) 159499, holotype).
Selected descriptions and icons. Munyanziza, Miombo trees and mycorrhizae: Pl. 5c (1994, black/white); Buyck & Verbeken, Mycotaxon 46: 429–442 (1995); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 202–204 & Pl. 199–207 (1996d); Buyck, Pagine Micol. 12: 39, fig. 2 (1999).
Pileus 45–140(–170) mm diam., very firm, fleshy, planoconvex to applanate and slightly depressed, subumbilicate to umbilicate; margin inflexed to involute when young, straight or sometimes revolute and irregular in older specimens; pellis dehiscent only near the extreme margin, smooth, dry, greasy when humid, finely areolate when old, cream, pale yellow to greyish yellow and greyish orange (4A4–5, 4B4–5, to 5B3), seldom entirely this colour, shaded with reddish brown to greyish buff (5A2, 6C5–6 to 6D5–6) or even locally dark brown (7C2). Lamellae subfree to narrowly adnate, arcuate to subventricose, easily separable from the pileus-trama, dense (L+l = 3+4 to 6+13/cm), unequal with 1–3(–5) lamellulae between two lamellae, (2–)6–10 mm broad, thin, paper-like, greyish orange (5B4) when young, dirty brown when bruised, dark brown (6F5–6 to 9E4–5 or 9F4–5) at maturity; edge entire, slightly paler. Stipe 25–80 × 10–30 mm, subcylindrical, firm, solid to fistulous, cream to pale yellowish (3A2 to 4A2), whitish pruinose in the upper half, leaving finger-prints when bruised, turning brownish grey near the stipe-base. Context (3–)6–12 mm thick in the pileus, whitish, strongly greying when cut, especially in the stipe-cortex, turning brownish in the central part of the stipe when older and when bruised; taste mild or bitter to faintly acrid, not particularly unpleasant; smell strong and unpleasant, more or less spermatic. Latex scarce to quite abundant, white, yellowish on drying; taste mild to slightly acrid. Spore-deposit dark brown (7F3–5) when fresh, fading to chestnut brown (5F7–8). – Pl. 37 & 38
Basidiospores ellipsoid, (7.6–)8.0–8.7–9.0–9.7 × (6.8–)7.0–7.6–8.8–8.6 mm (Q = 1.04–1.12–1.14–1.22; n = 120); ornamentation strongly amyloid, composed of a partial, very irregular reticulum; ridges < 1(1.5) mm high; a few isolated warts present; meshes smaller on the adaxial side; plage distinct, inamyloid. Basidia 48–64 × 10–13 mm, clavate to ventricose, sometimes with slightly thickened wall, 4-spored; sterigmata 4–7 × 2–3 mm. Pleurocystidia abundant, (30–)62–83 × 8.8–13(–14) mm, cylindrical to fusiform or ventricose, near the apex often with irregular, knotty protuberances, thin-walled; content slightly granular to needle-like. Pleuropseudocystidia (2–)3–5 mm diam., cylindrical to irregularly tortuous, obtuse, sometimes forked, often slightly subterminally restricted; content densely oleiferic, sometimes granular. Lamellar edge sterile, marginal cells small, cylindrical to clavate, 0–1(2)-septate. Hymenophoral trama irregular, composed of delicate, hyaline hyphae of 3–6 mm diam.; sphaerocytes only near the pileus trama; oleiferic hyphae very abundant, (2–)2.5–5 mm diam., sometimes branched; laticiferous hyphae abundant, frequently branched, 5–8 mm diam. Pileipellis one-layered, 50–100 mm thick, with a gelatinous matrix at least when young, composed of densely interwoven, recumbent to ascending hyphae and the extremities of laticiferous and oleiferous hyphae; terminal cells 20–55 × 3–6 mm, cylindrical, unbranched, obtuse, sometimes slightly tortuous; pileocystidia scarce and obscure, 43–80 × 3–6 mm, capitulate or tapering upwards, with poor, needle-like content. Stipitipellis resembling the pileipellis, with more laticiferous hyphae; caulocystidia numerous, 30–40 × 2–4 mm, capitulate. Ectomycorrhizal sheath with distinct laticiferous hyphae and numerous myceliocystidia measuring 45–70 × 2–4 mm. Clamp-connections absent. – Fig. 82
Ecology. Presumably only mycorrhizal with Brachystegia spp. (B. utilis, B. microphylla, B. spiciformis) in Zambezian miombo woodlands.
Distribution. Known from Burundi, Tanzania (e.g.: Munyanziza 1994: Morogoro distr., Uluguru Mts.), Zambia and Zimbabwe. Not common.
Revised specimens
Burundi. Bururi Prov. Nkayamba, just N of Rumonge, March 1992, B. Buyck 4237, 4247, 4264, 4265 & 4305 (all PC); Ibid., April 1992, B. Buyck 4370, 4384 & 4436 (all PC); Ibid., Nov. 1994, B. Buyck 4555, 4556, 4626, 4627 & 4628 (all PC); Ibid., Dec. 1992, B. Buyck 4739, 4741 & 4742 (all PC); Ibid., Jan. 1993, B. Buyck 4919 (PC); Ibid., March 1993, A. Verbeken 94-148 (BR 57602–81); Nyamirambo, Rumonge Forest Reserve, March 1993, B. Buyck 5134 (PC); Ibid., March 1994, A. Verbeken 94-387 (BR 57656–38), 94-467 (BR 57673–55) & 94-468 (BR 57674–56); Ibid., April 1994, A. Verbeken 94-555 (BR 57697–79).
Tanzania. Southern Highlands Prov., Njombe distr., between Sengele and Masaulva, 2 Feb. 1993, T. Saarimäki & al. 1539 (H); Morogoro, Morogoro distr., 20 Dec. 1992, E. Munyanziza 1-8 (WAG); Ibid., 25 April 1993, E. Munyanziza 6-9 (WAG); Ibid., 2 May 1993, E. Munyanziza 7-11 (WAG).
Zambia. Luapala Prov., near Mansa, 7 Dec. 1978, A.C. Rose 7811 (K(M) 159499, holotype).
Zimbabwe. Midlands Prov., Mvuma, Central Estates, Beacon Hill section, 21 Jan. 1997, C. Sharp 569-97 (BR 60410–76); Manicaland Prov., 15.5 km peg along Mutare-Bvumba road, 11 Feb. 1999, A. Verbeken 99-174 (GENT).
Notes. When comparing dimensions of spores, basidia, and cystidia of the type with those provided by Pegler (1983), some important differences are observed, in particular concerning spores and basidia However, the dimensions of the revised specimens agree well with the measurements obtained from examination of the type (Buyck & Verbeken 1995).
Sect. Piperites (Fr. ex J. Kickx f.) Burl. s.l.
Lactarius sect. Piperites (Fr. ex J. Kickx f.) Burl., Mem. Torrey Bot. Club 14: 14 (1908).
Lactarius subgenus Piperites (Fr. ex J. Kickx f.) Kauffman, Agaricaceae of Michigan I : 88 (1918).
incl. Lactarius sect. Afrobarbati Verbeken, Mycotaxon 77: 439 (2001).
− Type species: Lactarius torminosus (Schaeff.: Fr.) Pers.
Notes. Species with a cutis to ixocutis are very well-represented in the northern hemisphere and classified into many sections and subsections L. subgenus Piperites. In Africa species with an ixocutis without differentiated elements are poorly represented (3 spp. described so far). We prefer to classify them here provisionally in a broad concept of L. sect. Piperites s.l. although a close relation to the temperate species is uncertain.
Key to tropical African species
1. Pileus with strongly hairy margin ................................................................................ 59. L. barbatus
Pileus with smooth margin ..................................................................................................................... 2
2. Stipe paler than the pileus, scrobiculate; taste mild to bitter; spore ornamentation composed of mainly isolated warts .............................................................................. 60. L. afroscrobiculatus
Stipe concolorous with pileus, not scrobiculate; taste burning acrid; spore ornamentation composed of mainly aligned warts ................................................................... 61. L. acrissimus
Species descriptionS
59. Lactarius barbatus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 198 (1996). – Type: Burundi, Nkayamba hill, 4 Dec. 1992, B. Buyck 4683 (BR 38485–73, holotype; PC, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 199–200 & Pl. 195–198 (1996d).
Pileus 48–110 mm diam., firm, fleshy, convex and deeply depressed to applanate and depressed; margin involute, strongly hairy; pellis dehiscent up to the centre, viscid, hairy, cream (3A2 to 4A2–3), with reddish brown spots. Lamellae adnate, unequal with lamellulae of different lenghts [1–2(3) lamellulae between two lamellae], rather distant, broad (3–5 mm), ivory-colour, with yellowish brown spots; edge concolorous, entire. Stipe 20–40 × 9–22 mm, cylindrical, smooth, whitish to cream, locally with yellowish to reddish brown spots, chambered to fistulous. Context white, changing orange in stipe when cut or bruised; taste mild, slightly bitter, like immature fruits; smell idem, Cantharellus-like. Latex not abundant, white. Spore-deposit not observed. – Pl. 38
Chemical reactions. Context slightly orange with FeSO4, reddish brown in stipe.
Basidiospores ellipsoid, 9.1–10.2–10.3–11.5 × 7.2–8.0–8.1–8.9(–9.2) mm (Q = 1.14–1.25–1.28–1.41; n = 40); ornamentation amyloid, composed of a coarse and regular reticulum, with small meshes; ridges up to 1.5 mm high; plage distally amyloid or almost totally amyloid. Basidia 40–50(–55) × 13–15 mm, subcylindrical to ventrose, 4-spored; content guttate. Pleurocystidia absent. Pleuropseudocystidia very abundant, emergent, 4–6 mm diam., cylindrical to tortuous, often branched, with rounded apex; content oleiferic and needle-like. Lamellar edge sterile; marginal cells cylindrical, slender, with rounded apex, 15–25(–30) × 3–5 mm, thin-walled, hyaline. Hymenophoral trama almost completely composed of lactifers; some sphaerocytes present. Pileipellis between a cutis and an ixocutis, composed of parallel hyphae which are long and slender, 2–5 mm diam., septate, thin-walled with guttate content; subpellis composed of hyaline hyphae which are more compact, 4–6 mm, very often septate and becoming broader near the septae. Stipitipellis a cutis, similar to pileipellis. Clamp-connections absent. – Fig. 83
Ecology. Wetter Zambezian miombo woodland, e.g. under Brachystegia microphylla, B. boehmii. Fructification presumably restricted to the first weeks of the rainy saison.
Distribution. Known from Burundi and Zambia.
Revised specimens
Burundi. Bururi Prov., Nkayamba, Rumonge, 14 Dec. 1991, B. Buyck 4025 (PC); Ibid., 18 Dec. 1991, B. Buyck 4072 (PC); Ibid., 4 Dec. 1992, B. Buyck 4644 (PC) & 4683 (BR 38485–73, holotype; PC, isotype).
Zambia. Luapala Prov., Mansa, Kwanfumwe, 2 Jan. 1991, B. Buyck 3312 (PC); Ibid., Kabunda mission, 3 Jan. 1991, B. Buyck 3352, 3359 & 3360 (all PC).
60. Lactarius acrissimus Verbeken & Van Rooij
Van Rooij & al., Nova Hedwigia 77: 225 (2003). − Type: Benin, Borgou Prov., Wari Maro, 20 June 1998, A. De Kesel 2161 (BR 112672–55, holotype; GENT, isotype).
Selected descriptions and icons. Van Rooij & al., Nova Hedwigia 77: 225, 240 (2003).
Pileus 31–50 mm diam., convex and depressed with regular margin; cream, darker in the center, dirty white at margin. Lamellae broadly adnate to decurrent, anastomosing near both the stipe and the margin, moderately distant (L+l = 13/cm); lamellulae (1l/L) of irregular length, cream (4A3). Stipe (17–)36–44 × (12–)18–23 mm, firm, thick and slightly clavate; surface completely finely tomentose, locally with a pinkish tinge. Context thick in pileus, solid in stipe, cream, sometimes with pale orange tinge; taste burning acrid. Latex not observed. Spore-deposit not observed. – Pl. 39
Chemical reactions. Context unchanging with NH4OH, HCL, KOH, HNO3, FeSO4 and phenol.
Basidiospores subglobose to broadly ellipsoid, sometimes globose or ellipsoid, 6.8–7.4–7.9–8.0 × 5.6–6.5–7.0–7.9 µm (Q = 1.00–1.10–1.16–1.25, n = 80); ornamentation amyloid, composed of aligned warts (up to 1 µm high), connected with fine lines forming a partial or complete reticulum, with some isolated warts present; plage inamyloid. Basidia (50)60–80 × 4–16 µm, cylindric to clavate, 4-spored, seldom 2-spored, often with oil-like content. Pleurocystidia absent. Pleuropseudocystidia not abundant, mostly not emergent, 5–13 µm diam., subclavate, with rounded apex, seldom subutriform, thin-walled; content granular, needle-like, sometimes refractive and oil-like. Lamellar edge fertile. Hymenophoral trama cellular with abundant lactifers. Pileipellis an ixocutis to (locally) ixotrichoderm, up to 190–250 µm thick, filamentous, hyphae slightly interwoven, hyaline, regularly cylindric, often septate, sometimes branched, 2–7 µm diam.; lactiferous hyphae present. Stipitipellis similar to pileipellis. Clamp-connections absent. – Fig. 84
Ecology. Found in savannah woodland dominated by Uapaca togoensis.
Distribution. Only known from the type locality.
Revised specimens
Benin. Borgou Prov., Wari Maro, 20 June 1998, A. De Kesel 2161 (BR 112672–55, holotype; GENT, isotype).
Notes. This species is well characterised by the presence of a conspicuous ixocutis to ixotrichoderm structure in the pileipellis and stipitipellis, rare in African species but typical for Lactarius subgenus Piperites.
61. Lactarius afroscrobiculatus Verbeken & Van Rooij
Van Rooij & al., Nova Hedwigia 77: 226 (2003). − Type: Benin, Borgou Prov., Wari Maro, 20 Sept. 1998, A. De Kesel 2280 (BR 112773–59, holotype; GENT, isotype).
Selected descriptions and icons. Van Rooij & al., Nova Hedwigia 77: 226-227, 241 (2003).
Pileus 64–108 mm diam., planoconcave to concave; pellis sticky and smooth, light yellow to yellowish brown (4A4–5 to 5C8–7); margin regular when young, undulate when older. Stipe 12–36 mm × 15–33 mm, short, thick, firm, cylindric to clavate; pellis scrobiculate, pale white. Lamellae decurrent, very dense (L+l = 13–16/cm), sometimes with anastomosing venation at the pileus margin; with one lamellula between two lamellae, very short, light yellow or darker (4A4–5), turning brownish yellow when bruised. Context very firm, thick in pileus, chambered to fistulous in stipe; taste first mild then bitter, like Lactarius subdulcis, faint resin-like; smell often somewhat fish-like. Latex not observed. – Pl. 39
Basidiospores broadly ellipsoid, sometimes subglobose or ellipsoid, 6.4–7.3–7.9–8.8 × 5.3–6.1–6.6–7.4 µm (Q = 1.06–1.17–1.21–1.35, n = 80); ornamentation amyloid, composed of irregular warts, up to 1 µm high, subconical to subspherical, mostly isolated, seldom connected with fine lines or aligned; plage slightly amyloid. Basidia 45–53 × 5–14 µm, subclavate, sterigmata long (7–12 × 2–3 µm) and sharp, thin-walled to slightly thickened, 4-spored; content seldom with oil-like droplets. Pleuromacrocystidia abundant, (43–)60–90 × 10–15 µm, subcylindrical, slightly tapering upwards or slightly clavate, slightly emergent; content needle-like and granular, slightly thick-walled. Pleuropseudocystidia very rare, 5–8 µm diam., irregularly cylindric, with refractive content. Lamellar edge sterile; marginal cells 12–21 × 2–6 µm, clavate to irregulary cylindric, sometimes branched, thin-walled, hyaline. Hymenophoral trama mainly cellular, with some hyaline hyphae and lactifers present. Pileipellis a distinct ixocutis, 120 µm thick, composed of densely interwoven recumbent or ascending hyphae, 2–5 µm diam., regulary cylindric, sometimes slightly tortuous, septate, sometimes branched, sometimes with little knobs. Stipitipellis an ixocutis to ixotrichoderm. Clamp-connections absent. – Fig. 85
Ecology. The material has been collected in extensively pastured savannah woodlands dominated by Burkea africana and Isoberlinia doka, as well as in savannahs dominated by Uapaca togoensis and/or Detarium sp.
Distribution. Only known from Benin.
Revised specimens
Benin. Borgou Prov., Wari Maro, 20 Sept. 1998, A. De Kesel 2280 (BR 112773–59, holotype; GENT, isotype); Ibid., 25 Sept. 1998, A. De Kesel 2337 (BR 112830–19); Ibid., 13 Sept. 2001, A. De Kesel 3160 (BR 149827–59); Ibid., 9 Oct. 2001, A. De Kesel 3189 (BR 149713–42); Ibid., 23 Sept. 2001, A. De Kesel 3236 (BR 149752–81); Borgou Prov., Doguè, 10 Oct. 2001, A. De Kesel 3216 (BR 149694–23).
Togo. Centrale Prov., Parc National Fazao, 16 Jun 2007, A. De Kesel 4387 (BR 163718–79); Forêt Classée d’Alédjo, 29 May 2008, A. De Kesel 4577 (BR 163477–32); Ibid., 31 May 2008, A. De Kesel 4599 (BR 163478–33).
Notes. The species is macroscopically well characterised by the sticky cap and scrobiculate stipe; microscopically by the conspicuous ixocutis structure, a common character in temperate species, but in tropical Africa only known for Lactarius afroscrobiculatus. Abundant pleurocystidia, scrobiculate stipe, brown staining gills and an ixocutis are typical for Lactarius subgenus Piperites (Fr. ex J. Kickx. f.) Kauffman. In subgenus Piperites the spore ornamentation is most often reticulate, but this species has spores with distinctly isolated and rounded warts, sometimes connected with fine connective lines. The spore ornamentation, the very scarce to sometimes absent pleuropseudocystidia and the knobby excrescences in the pileipellis are characters pleeding for a classification in Lactarius subgenus Russulopsis Verbeken, section Russulopsidei Verbeken. As the pale colours and especially the scrobiculate stipe are not known in this section, we consider the classification of this species as provisory uncertain.
Remarkably, for some specimens, even young and fresh ones, no latex could be observed. Microscopically, pseudocystidia could not be observed in all specimens, but their presence in some basidiocarps convinced us that this species has to be considered as a true Lactarius species. The same phenomenon was also observed in Lactarius ruvubuensis Verbeken, where basidiocarps without latex and without pseudocystidia occur together with basidiocarps with latex and pseudocystidia.
Lactarius sect. Amari Verbeken, Mycotaxon 77: 441 (2001).
− Type species: Lactarius subamarus Verbeken.
Key to tropical African species
1. Spores on average 9.5–10 × 8.5 µm; taste mild and variable (farinaceous, raphanoid); lamellae free to adnexed, rather spaced ....... 62. L. subamarus
Spores average 7.0 × 5.7 µm; taste acrid and bitter; lamellae deeply decurrent, crowded ........ 63. L. amarus
Species descriptionS
62. Lactarius subamarus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 209 (1996). – Type: Burundi, Mugara, 21-11-1978, J. Rammeloo 5850 (BR 8771–41, holotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 197–198 & Pl. 191–194 (1996d); Verbeken & al., Syst. Geogr. Pl. 70: 206–207 (2000); Verbeken, Micol. Veget. Medit. 16: fig. 5 (2001).
Pileus (10–)20–30(–40) mm diam., convex then applanate, depressed or distinctly umbilicate; margin regularly to slightly crenulate, shortly and slightly striate to slightly grooved; pellis not dehiscent, smooth, glabrous, with concentrical zones, especially when dry, yellowish white to pale yellow (4A2–5) and pale orange, pale dirty orange-yellow (5B5) or buff, a bit darker in the center, hygrophaneous (becoming paler when drying out, especially in the middle). Lamellae free to adnexed, unequal with lamellulae of different lenghts, rather distant (5+3/cm), moderately thick, somewhat paper-like, rather broad (2.5–5 mm) and rounded, first pale yellow (4A4–5), then more yellowish orange, pale orange (5A3–4); edge entire, concolorous. Stipe 20–35 × 1–5 mm, cylindrical, with a bulb-like structure at the base; pellis smooth, mat, sometimes whitish pruinose, sometimes longitudinally grooved, paler than the pileus, yellowish white to pale yellow (5A2–3), sometimes whitish and hirsute at the base. Context thin-fleshy, chambered to fistulous in the stipe, whitish to pale yellow (4A3), unchanging, unchanging with FeSO4; smell not particular, agreeable, sweet; taste mild at first, then a bit menthol-like, acrid, sweet and refreshing, with a component of cedar-wood. Latex not abundant, transparent to white, unchanging; taste not particular. Spore-deposit white. – Pl. 40
Basidiospores subglobose to ellipsoid, sometimes globose, 8.0–9.5–10.2–11.2 × 7.7–8.3–8.5–9.3 mm (Q = 1.01–1.13–1.20–1.29; n = 100); ornamentation amyloid, composed of ridges up to 1.5 mm high, often with smaller branches, and some isolated, irregularly shaped warts; ornamentation very dense although never forming a complete reticulum; wall verrucose; plage distally amyloid. Basidia 50–60(–70) × 10–11 mm, cylindrical to slightly clavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia very abundant, cylindrical to irregularly tortuous or even cork-screw-like and often branched. Lamellar edge sterile; marginal cells 15–25(–45) × 7–11 mm; cheilopseudocystidia abundant. Hymenophoral trama subregular, composed of hyaline hyphae and laticiferous hyphae; sphaerocytes lacking. Pileipellis a cutis to trichoderm, 100–150 mm thick, filamentous, with interwoven hyaline, regularly cylindrical hyphae which are often septate, 3–5 mm diam., sometimes branched with slightly thickened wall; laticiferous hyphae present. Stipitipellis as pileipellis. Clamp-connections absent, but similar structures observed in pileipellis. – Fig. 86
Ecology. Zambezian miombo woodland. In Zimbabwe Lactarius subamarus was found most often on naked sandy soils while in Burundi it was mostly observed among mosses (Leucobryum).
Distribution. Restricted to the Zambezian miombo woodland area; known from Burundi, Democratic Republic of Congo, Tanzania and Zimbabwe.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, 9 May 1993, B. Buyck 5104 (PC); Ibid., 19 Nov. 1993, B. Buyck 5192 (PC); Ibid., 26 Nov. 1993, B. Buyck 5226 & 5228 (all PC); Ibid., 26 Nov. 1993, B. Buyck 5231 & 5233 (all PC); Ibid., 3 Dec. 1993, B. Buyck 5288, 5290-5293 (all PC); Mugara, 21 Nov . 1978, J. Rammeloo 5850 (BR 8771–41, holotype); Ibid., 24 Nov. 1978, J. Rammeloo 5907 (BR 8772–42); Ibid., 2 Dec. 1978, J. Rammeloo 6004 (BR 18875–57); Ibid., 6 Dec. 1978, J. Rammeloo 6102 (BR 18879–61).
Democratic Republic of Congo. Shaba region, Luiswishi, 14 Dec. 1971, D. Thoen 5118 (BR 8765–35).
Tanzania. Morogoro Prov., Morogoro distr., 26 April 1993, Munyanziza 6-4 (WAG).
Zimbabwe. Manicaland Prov., Chipinge, Busi farm, around Scott’s house, 12 Feb. 1999, A. Verbeken 99-184 (GENT); Mashonaland East Prov., Bromley, Liemba farm, 3 Feb. 1999, A. Verbeken 99-110 (GENT); Midlands Prov., Central Estates, Baru section, 27 Jan. 1999, A. Verbeken 99-037b (GENT); Midlands Prov., Mvuma, Central Estates, south of Beacon Hill ridge, A. Verbeken 99-021 (GENT).
Notes. Lactarius subamarus differs from L. amarus R. Heim (only known from type) in the significantly larger spores, the free to adnexed lamellae and in never having a "bitter and acrid" taste. Its taste seems variable or at least difficult to define: often mild, but "particular" in Buyck 5192, "farinaceous to raphanoid" in Rammeloo 5907, "slightly acrid to raphanoid" in Rammeloo 6102, "farinaceous" in Rammeloo 6004, “acrid, but refreshing and menthol-like” (observations by the author).
Heim, Candollea 7: 379 (1938). – Type: Madagascar, near Manompana, 16 Dec. 1934, R. Heim I56 (PC, holotype).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 56–58 (1938); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 196–197 & Pl. 190 (1996d).
Pileus 40–45 mm diam., infundibuliform, irregular; margin undulate; pellis not dehiscent, smooth, not viscid, regularly concentrically zoned, yellowish orange. Lamellae deeply decurrent, unequal with a lot of very short lamellulae, crowded, narrow (1.5 mm), rather thick, cream, yellow when older. Stipe 20 × 10 mm, short, tapering downwards, clearly but shortly rooting, smooth, hirsute at the base, cream. Context firm, fairly elastic, cream, 4 mm thick in the centre of the pileus, translucid near the margin, fistulous in the stipe; smell not particular. Latex white, unchanging; taste acrid and bitter. Spore-deposit white. (According to Heim 1938b).
Chemical reactions. Context lilac with pyramidon, immediately blue green with gaiac, immediately purple with gaiacol. (According to Heim 1938b).
Basidiospores ellipsoid, (6.0–)6.5–7.0–7.6(–8.3) × (4.5–)5.2–5.7–6.2(–6.9) mm (Q = 1.08–1.24–1.42; n = 35); ornamentation amyloid, composed of irregularly shaped warts and ridges, never forming a reticulum, up to 1 mm high; wall slightly verrucose; plage faintly distally amyloid. Basidia 30–35 × 10–12 mm, cylindrical to clavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia abundant, 3–5 mm diam., cylindrical, narrow, with rounded or mucronate apex. Lamellar edge sterile; marginal cells 10–25 × 5–8 mm, short, cylindrical, sometimes septate. Hymenophoral trama subregular, composed of hyaline hyphae (2–5 mm diam.) and laticiferous hyphae; no sphaerocytes present. Pileipellis a cutis to trichoderm, 100–150 mm thick, filamentous, composed of interwoven hyphae; hyphae long, cylindrical, 3–5 mm diam., often septate, sometimes branched, slightly thick-walled. Stipitipellis as pileipellis; hyphae becoming more thick-walled at the base of the stipe. Clamp-connections absent. – Fig. 87
Ecology. Found in Malagasy lowland rainforest.
Distribution. Only known from the type locality.
Revised specimens
Madagascar. Betsimisaraka Prov., near Manompana, Dec. 1934, R. Heim I56 (PC, holotype).
Notes. The type consists of one half basidiome and is preserved in alcohol. This makes it difficult to observe the spores in Melzer's reagent.
Lactarius sect. Russularia Fr. ex Burl., Mem. Torrey Bot. Club 14 : 14 (1908).
− Type species: Lactarius subdulcis (Pers.: Fr.) Gray.
Species description
64. Lactarius hepaticus Plowr.
Boud., Icon. Mycol. 4: 28 (1905). – Type: not selected.
Selected descriptions and icons. Heilmann-Clausen & al., Fungi of Northern Europe 2: 188–189 (1998); Basso, Fungi Europaei 7: 524–528 (1999); Van der Westhuizen & Eicker, Field guide mushrooms southern Africa: 52–53 (1994).
Ecology. Ectomycorrhizal with introduced Pinus spp.
Distribution. Known from Madagascar. Also present in pine plantations of the South African Republic (Van der Westhuizen & Eicker 1994).
Lactarius sect. Nigrescentes Verbeken, Persoonia 17: 379 (2001).
− Type species: Lactarius melanogalus R. Heim.
Notes. This section contains the blackening species of L. subgenus Plinthogalus, and is maintained for practical reasons.
Key to tropical African species
1. Spores ellipsoid, Q = 1.25–1.7 ............................................................................................................... 2
Spores globose to subglobose, rarely ellipsoid, Q = 1–1.2 .............................................................. 3
2. Spores entirely winged; pileus and stipe greyish yellow to yellowish brown; pileus without papilla, not veined ................................................................................................................. 67. L. orientalis
Spores partially winged, with conical warts and lower ridges present; pileus and stipe dark brown; pileus with a distinct papilla ............................................................................. 68. L. griseogalus
3. Spore ornamentation 1–2 mm high; spores 7.3–7.5 ´ 6.5–6.7 mm; latex finally bluish black ........... 65. L. melanogalus
Spore ornamentation up to 1(1.3) mm high; spores 7.9–8.3 ´ 7.3–7.6 mm; latex finally buff to cream .......... 66. L. baliophaeus
Species descriptionS
65. Lactarius melanogalus R. Heim ex R. Heim
Heim, Bull. Jard. Bot. Etat Bxl 25: 46 (1955). – Type: Democratic Republic of Congo, Binga, M. Goossens-Fontana 979 (BR 7724–61, lectotype).
Excl.: Lactarius melanogalus ss. Nzigidahera (1993), = both L. baliophaeus and L. orientalis.
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 46–50 & Pl. 5, fig. 1–2 (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 14, fig. 6 (1955); Pegler & Young (1982: 33); Verbeken, Persoonia 16: 211–214 (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 171–173 & Pl. 139–141 (1996d).
Pileus (25–)30–40(–55) mm diam., thin, first applanate, then slightly depressed; margin irregular, undulate, not incurved; pellis not dehiscent, smooth, mat, dry, strongly radially wrinkled, olivaceous ochraceous brown to dark brown (5E4–6, 5F4–6, 6E6–7 to 6F6–7) with black spots. Lamellae adnexed to adnate with decurrent tooth, unequal with lamellulae of different lengths, rather dense, 2–3 mm broad, thick, greasy, greyish, ochraceous, cream, with black spots; edge slightly crenular, sometimes darker. Stipe (15–)25–40(–70) × 4–10 mm, cylindrical, curved near the base, smooth, longitudinally grooved, firm, dark cream and greyish brown (4E6–8 to 4F6–8 to 6C3–4 or 6D3–4), with black spots. Context thin, first translucid to cream, then greyish pink, finally blackening; taste mild to acrid; smell not remarkable, sometimes sweet. Latex abundant, first watery, then cream to greyish, finally bluish black. Spore-deposit not observed. – Pl. 40
Chemical reactions. Context intense blue-green (both Goossens-Fontana 979 and B. Buyck 1356) with gaiac; reddish orange to greyish red (± 7B7 to 7C7) with formaline; greyish yellow (4B4) with NH4OH and reddish, then brown (7E8) with aniline.
Basidiospores globose to subglobose, sometimes ellipsoid, 6.7–7.3–7.5–8.2 × 6.0–6.5–6.7–7.2 mm (Q = 1.04–1.11–1.13–1.20; n = 60); ornamentation amyloid, composed of high ridges (1–2 mm), forming a winged reticulum, without isolated warts, denser on adaxial side; plage inamyloid. Basidia 30–45 × 9–11 mm, cylindrical to narrowly utriform, 4-spored. Pleurocystidia scarce, not emergent, 33–43 × 6–8 mm, cylindrical to fusiform, sometimes irregular, with slightly thickened wall. Pleuropseudocystidia scarce; content dark brown. Lamellar edge sterile; marginal cells 18–28 × 4–6 mm, cylindrical to fusiform and tapering upwards; wall slightly thickened; content brown. Hymenophoral trama irregular, composed of hyaline, thin-walled hyphae and abundant laticiferous hyphae with remarkable brown content. Pileipellis a hymeniderm; elements of the suprapellis 20–40 × 7–13 mm, thin-walled, subglobose to slightly clavate or clavate, with brown intracellular pigmentation; subpellis pseudoparenchymatuous, thin. Stipitipellis a hymeniderm; elements of the suprapellis (10–)15–30(–35) × 3–6(–10) mm, cylindrical, slender, sometimes fusiform or irregular, with brown intracellular pigment; subpellis pseudoparenchymatous. Clamp-connections absent. – Fig. 88
Ecology. Found in rainforest, dry coastal forest, drier Guineo-Congolian rainforest, wetter Zambezian miombo woodland.
Distribution. Known from Benin, Cameroon, Democratic Republic of Congo, Ghana, Guinea, Ivory Coast (Heim 1955b), Gabon (Heim 1955b) and Zambia.
Revised specimens
Benin. Atacora Prov., Kota, 26 Aug. 1997, A. De Kesel 2003 (BR 74849–62); Ibid., 30 Aug. 1997, A. De Kesel 2021 (BR 74867–80).
Cameroon. South Western Prov., near Mundema, Korup National Park, transect P, 1984, I. Alexander "6" (E); Ibid., plot 10, 23 March 1991, R. Watling 23140 (E); Ibid., plot 15-18, 23 March 1991, R. Watling 23141 (E).
Democratic Republic of Congo. Equateur Prov., Binga, Dec. 1934, M. Goossens-Fontana 979 (BR 7724–61, lectotype); Tshopo Prov., 5 km NNE of Batiabongena, 8 April 1984, B. Buyck 1356 (BR 18877–59).
Ghana. Cape Coast Prov., Jukwa road, May 1973, A.C. Rose 7306 (K(M) 38464).
Guinea. Forêt humide de Ziama, 12 April 2000, A. Bâ 350 (GENT).
Zambia. Luapala Prov., Mansa, Kwanfumwe, 2 Jan. 1991, B. Buyck 3332 (PC).
Notes. Heim first used the name Lactarius melanogalus (1943) for a specimen from Ivory Coast, without latin diagnosis. In 1955 he redescribed the species, latin diagnosis included, and cited specimens from Ivory Coast, Cameroon and Democratic Republic of Congo . He did not indicate a type. As Heim A87 and Heim “Q43” could not be traced at PC anno 1993, Goossens-Fontana 979 was designated as lectotype (Verbeken 1996). This collection consists of two basidiomes in good condition. In fact, there is a specimen labeled Heim “Q43” at PC which represents Lactarius gymnocarpus and also cited by Heim. As both species are not likely to be confused, we suppose that there were two different collections with the same number. The one representing L. melanogalus could not be traced at PC.
The macroscopic description is based on the description by Heim (1955b) and completed with fieldnotes on Buyck 3332 & 1356. The microscopic description is based on Goossens-Fontana 979, supplemented with spore-measurements of Buyck 1356 & 3332.
Following the description by Heim, the colour of the pileus is ochraceous to olive-brown and the illustration of Heim A87 shows indeed a rather pale mushroom. Goossens-Fontana 979 is mentioned (Heim, 1955a) to be dark brown ("bistre sombre"), which is also the colour of the specimens of Buyck and Watling. The water-colour of Heim A87, which shows a very young specimen, proves that the pileus is ochraceous when young and becomes darker with age.
The taste of latex and context is said to be very acrid in the specimens collected by Heim, very bitter in Goossens-Fontana 979, very acrid (slowly) in Buyck 3332, mild in Buyck 1356. There is a strong and sweetish smell in Buyck 1356, a fish-smell in Heim Q43 and a smell of “Russula pectinata” (Foetentinae) in Heim A87.
The presence of aberrant spores is mentioned by Heim (1955b, fig. 15). Those spores can have the same size as the normal spores, but they have strongly amyloid droplet-like, globose warts, very irregular in size and number, and there is no sign of the normal reticulate ornamentation at all. We observed such spores in Goossens-Fontana 979 and in the specimens of Cameroon, although they are certainly not characteristic for this species. Similar aberrant spores were observed in collections of other African (and even European, e.g. Lactarius uvidus) Lactarius and Russula species. An explication for their presence is unknown.
66. Lactarius baliophaeus Pegler
Pegler, Kew Bull. 23: 237 (1969). – Type: Ghana, Tafo, 29 July 1955, M. Holden GC66 (K(M) 159497, holotype).
Selected descriptions and icons. Pegler, Kew Bull. 23: 237–239 (1969); Verbeken, Persoonia 16: 215–219 & Pl. 1 (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 174–175 & Pl. 145–147 (1996d); De Kesel & al., Guide des champignons comestibles du Bénin: 171 (2002).
Pileus 30–70(–90) mm diam., planoconvex, applanate to slightly depressed, infundibuliform when older; margin slightly incurved when young, then crenulate to undulate, striate when dry; pellis not dehiscent, quite thick, smooth, dry, mat, greyish yellow to brownish orange (4A3–5 to 4B3–5 to 5C4) to dark blond and yellowish brown (5D4–5 to 5E4–5), staining dark brown and black. Lamellae broadly adnate to decurrent, crowded (4+12 to 3+6/cm), unequal with lamellulae (3–5(–7) between 2 lamellae, regular pattern), thin, paper-like, broad (3–)5–7 mm, cream, pale greyish, staining reddish then black; edge entire, sometimes blackish brown. Stipe central to eccentric, (15–)25–50 × 5–10 mm, cylindrical, clavate towards the base, greyish yellow to brownish orange or dark blond, sometimes dark grey, staining black and dark brown, firm, smooth, dry. Context firm, white to cream, changing immediately to orange-red, greyish red and finally black; taste mild, sometimes slightly bitter (Verbeken 94-283) or even first mild, then acrid (Verbeken 94-438). Latex very abundant, water-like, transparently brownish, soon blood-red and then changing to buff and cream or brownish, taste mild (acrid in Verbeken 94-438). Spore-deposit cream (IIa ss. Romagnesi 1967). – Pl. 41 & 42
Chemical reactions. Context unchanging with FeSO4, HCl, NH4OH. Pileipellis intense red with NaOH.
Basidiospores globose to subglobose, 7.0–7.9–8.3–9.0 × 6.5–7.3–7.6–8.2 mm (Q = 1.01–1.07–1.09–1.15, n = 80); ornamentation strongly amyloid, composed of ridges, up to 1(–1.3) mm high, forming a complete reticulum; isolated warts very scarce; plage distally amyloid. Basidia 38–48 × 10–12 mm, clavate to utriform, 4-spored; sterigmata 3–6 × 1–2 mm; content granular or guttate. Pleurocystidia scarce to abundant, not emergent, often arising deep in the hymenium, 40–55 × 9–11, fusiform, wall slightly thickened and brown pigmented. Pleuropseudocystidia (2–)3–6 mm diam., cylindrical, rarely tortuous, with rounded, tapering or mucronate apex; content oleiferic, yellowish brown; sometimes emergent, quite abundant. Lamellar edge sterile; marginal cells 21–40 × (2–)3–6 mm, cylindrical to tortuous or fusiform, with rounded, mucronate or tapering apex, sometimes septate, sometimes slightly thick-walled, hyaline. Hymenophoral trama irregular, composed of thin-walled, hyaline hyphae and rather abundant lactifers with a brown content. Pileipellis a hymeniderm; elements of the suprapellis 10–25 × 3–5 mm, cylindrical, slender, some fusiform, thin-walled, with brown intracellular pigment; subpellis pseudoparenchymatous. Stipitipellis hyphae interwoven and ascending in suprapellis, no spherical cells, terminal elements cylindrical to slightly tortuous, 20–30 × 3–5 mm, thin-walled, with brown intracellular pigment; lactifers in the under layer abundant. Clamp-connections absent. – Fig. 89
Ecology. Found in various forest types, such as miombo woodland with Brachystegia spp., semi-evergreen forest, Guineo-Congolian rainforest, Sudanian savannah woodland with dominance of Burkea africana or Afzelia africana.
Distribution. Widespread in tropical Africa, not uncommon. Known from Benin, Burundi, Ghana, Malawi, Senegal, Zambia and Zimbabwe.
Revised specimens
Benin. Borgou Prov., Wari Maro, 29 June 1998, A. De Kesel 2223 (BR 112732–18) & 2224 (BR 112733–19); Ibid. 16 Jun. 2002, A. De Kesel 3418 (BR 152139–43); Donga Prov., Kikélé, 13 Jun. 2002, A. De Kesel 3390 (BR 152099–03); Pénésoulou, 8 Oct. 2002, A. De Kesel 3538 (BR 152023–24); Atacora Prov., Natitingou, Kota falls, 19 Jun. 2004, A. De Kesel 3705 (BR 157098–55).
Burundi. Bururi Prov, Nyamirambo, Rumonge Forest Reserve, May 1993, B. Buyck 5091, 5092 & 5137 (all PC); March 1994, A. Verbeken 94-012 (BR 57563–42), 94-028 (BR 57571–50), 94-069 (BR 57580–59), 94-283 (BR 57637–19), 94-386 (BR 57654–36), 94-438 (BR 57664–46) & 94-472b (BR 57679–61); Nkayamba, just N of Rumonge, 18 Dec. 1991, B. Buyck 4062 & 4063 (all PC); Ibid., 13 April 1992, B. Buyck 4338 (PC); Ibid., 4 Dec. 1992, B. Buyck 4641 & 4642 (all PC); Ibid., 15 March 1994, A. Verbeken 94-153 (BR 57607–86) & 94-165 (BR 57618–00).
Ghana. Tafo, 29 July 1955, M. Holden GC66 (K(M) 159497, holotype).
Malawi. Northern Prov., Mzimba distr., Ntanga Forest Reserve, 30 Jan. 2005, A. Verbeken 05-155 & 05-156 (all GENT); Ibid., 30 Jan. 2005, F. Noe 05/18 (GENT); Ibid., near Chicangana village, 19 Feb. 2005, F. Noe 05/396 (GENT).
Senegal. Kolda Prov., 25 June 1986, D. Thoen 7662 (BR 3168–64).
Zambia. Luapala Prov., near Kawambwa, 6 Jan 1991, B. Buyck 3446 (PC); farm Gibsons, 16 Jan. 1996, B. Buyck & G. Eyssartier 96-057 (PC); Mpongwe, 25 Jan. 1996, B. Buyck & G. Eyssartier 96-191 (PC); Lusaka, near airport, 11 Feb. 1996, B. Buyck & G. Eyssartier 96-358 (PC).
Zimbabwe. Midlands Prov., Mvuma, paddock 77 Central Estates, 5 Feb. 1999, A. Verbeken 99-125 (GENT); Ibid., Beacon Hill section, 26 Jan. 1999, A. Verbeken 99-007 (GENT); Manicaland Prov., Mutare-Bvumba road near Prince of Wales Viewpoint, 11 Feb. 1999, A. Verbeken 99-181 (GENT); Chipinge, Busi farm, forests around Scott’s house, 12 Feb. 1999, A. Verbeken 99-206 (GENT); Mguzu, 5 Feb. 1999, Gansemans in A. De Kesel 2742 (BR 115757–36); Chipinge, 10 Feb. 1999, A. De Kesel 2445 (BR 112615–95); Mashonaland Central Prov., Harare, Chisipite suburb, Greenberg’s, 12 Jan. 1999, D. Arora 99-2 (SFSU).
Notes. The macroscopic description is compiled from the description of Pegler (1969) and the fieldnotes of Verbeken. The microscopic description is based on Holden GC66, Verbeken 94-153 & 94-283.
The blood-red colour of the latex disappears very quickly and soon the latex becomes pinkish grey and then buff.
67. Lactarius orientalis (Verbeken) Verbeken
Verbeken, Persoonia 17: 380 (2001). − Lactarius baliophaeus var. orientalis Verbeken, Persoonia 16: 219 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 30 March 1994, A. Verbeken 94-472 (BR 57678–60, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Persoonia 16: 218–219 (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 175–176 & Pl. 147–148 (1996d); Verbeken, Persoonia 17: Pl. 2 (2001).
Pileus 30–45 mm diam., planoconvex, applanate to slightly depressed, infundibuliform when older, somewhat irregular and asymmetrical; margin straight, striate when dry; pellis not dehiscent, quite thick, smooth, dry, mat, brownish orange to yellowish brown, staining dark brown and black. Lamellae broadly adnate to decurrent, crowded (L+l = 4+12 to 3+6/cm), unequal with lamellulae (3–5(7) between 2 lamellae, regular pattern), thin, paper-like, 3–5 mm broad, cream, pale greyish, staining reddish then black; edge entire, sometimes blackish brown. Stipe central to eccentric, 25–50 × 10 mm, cylindrical, clavate towards the base, greyish yellow to brownish orange or dark blond, sometimes dark grey, staining black and dark brown, firm, smooth, dry. Context firm, white to cream, changing immediately to orange-red, greyish red and finally black. Latex very abundant, water-like, transparently brownish, soon blood-red and then changing to buff and cream or brownish. Spore-deposit cream (IIa ss. Romagnesi 1967). – Pl. 43
Chemical reactions. Context unchanging with FeSO4, HCl, NH4OH. Pileipellis intense red with NaOH.
Basidiospores ellipsoid, sometimes elongate, 7.4–8.7–9.4–10.3 × 5.8–6.4–7.0–7.4(–7.7) µm (Q = 1.21–1.34–1.36–1.64, n = 40); ornamentation strongly amyloid, composed of ridges, up to 1(–1.3) mm high, forming a complete reticulum; isolated warts or short ridges very scarce; plage slightly distally amyloid. Basidia 35–50 × 10–12 mm, cylincrical to subclavate to slightly utriform, 4-spored; sterigmata 3–6 × 1–2 mm; content granular or guttate. Pleurocystidia scarce to abundant, not emergent, often arising deep in the hymenium, 40–55 × 9–11 µm, fusiform, wall slightly thickened and with brown pigmentation. Pleuropseudocystidia quite abundant, (2–)3–6 mm diam., sometimes emergent, cylindrical, rarely tortuous, with rounded, tapering or mucronate apex; content oleiferic, yellowish to very dark brown. Lamellar edge sterile; marginal cells 21–40 × (2–)3–6 mm, cylindrical to tortuous or fusiform, with rounded, mucronate or tapering apex, sometimes septate, sometimes slightly thick-walled, hyaline. Hymenophoral trama irregular, composed of thin-walled, hyaline hyphae and rather abundant lactifers with a brown content. Pileipellis a hymeniderm; elements of the suprapellis 10–25 × 3–5 mm, cylindrical, slender, some fusiform, thin-walled, with brown intracellular pigment; subpellis pseudoparenchymatous. Stipitipellis hyphae interwoven and ascending in suprapellis, no spherical cells; terminal elements cylindrical to slightly tortuous, 20–30 × 3–5 mm, thin-walled, with brown intracellular pigment; lactifers in the under layer abundant. Clamp-connections absent. – Fig. 90
Ecology. Zambezian miombo woodland, found mainly under Brachystegia spp.
Distribution. Known from Burundi, Zambia and Zimbabwe.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, Jan. 1993, B. Buyck 5232 (PC); Ibid., Nov. 1993, B. Buyck 5229 (PC); Ibid., March 1994, A. Verbeken 94-215 (BR 57623–05), 94-472 (BR 57678–60, holotype; GENT, isotype) & 94-523 (BR 57693–75); Nkayamba, just N of Rumonge, April 1992, B. Buyck 4375 (PC).
Zambia. Copperbelt Prov., Chati-forest, near Kitwe, Dec. 1990, B. Buyck 3115, 3132, 3225, 3227 & 3214 (all PC); Luapala Prov., Mansa, Kwanfumwe, Jan. 1991, B. Buyck 3322 (PC).
Zimbabwe. Midlands Prov., Mvuma, Central Estates, south of Beacon Hill, 27 Jan. 1999, A. Verbeken 99-025 (GENT); Mvuma, Central Estates, Jan. 1995, C. Sharp 325/95 (BR 38492–80); Mashonaland East Prov., Bromley, Liemba farm, 2 Feb. 1999, A. Verbeken 99-071 (GENT).
Notes. Lactarius orientalis differs from L. baliophaeus especially by the spore shape and dimensions. The spores are ellipsoid, 7.4–8.7–9.4–10.3 × 5.8–6.4–7.0–7.4(–7.7) mm (Q = 1.21–1.34–1.36–1.64; n = 60). It is difficult to discriminate Lactarius baliophaeus and L. orientalis in the field, but L. orientalis seems to lack the ochraceous/buff cap colours and the crenulate margin, characteristic for L. baliophaeus.
68. Lactarius griseogalus R. Heim
Heim, Rev. Mycol. (Paris) 32: 204 (1967). – Type: Central African Republic, Bébé savanna, close to Filifi river, 6 Aug. 1966, R. Heim LM2189 (PC, holotype).
Selected descriptions and icons. Heim, Israel J. Bot. 15: 158–162 (1966); Verbeken, Persoonia 16: 214–216 (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 173–174 & Pl. 142–144 (1996d); Verbeken, Persoonia 17: 380–381 (2000).
Pileus up to 36 mm diam., irregular, strongly convex or planoconvex when young, sometimes remaining so but usually becoming widely infundibuliform, first with prominent umbo which is sometimes present in older specimens too; margin lobulate, striate, grooved; surface with dark radial veins prolonging in the intervals between the grooves; pellis very dark brown, almost black, even darker in the center, unicolorous but with very small grey spots (handlens!) giving a powdery aspect. Lamellae broadly adnate, not decurrent, distant (about 28 in total), with lamellulae, rather thick, ivory white, but with some brownish black especially at the edge, becoming completely convered with black dots; edge entire, dark brown. Stipe 30–35 mm long, irregularly to regularly cylindric, curved, flexuose, tapering downwards, slightly eccentric, very dark brown, like the centre of the pileus, with greyish tinge which is absent in a ring-like zone on top and betwee the lamellae, underneath with very thin and irregular veins giving a somewhat bulgy aspect; sometimes slightly strigose at the base. Context very thin and transparent in the pileus, white then cream in the stipe, in some seconds turning red in the pileus, then intensely reddish orange, finally completely black; taste mild, slightly sweet. Latex very abundant, hyaline, then turning grey, with oil-like consistency, dark brown with orange tinge on white paper; taste mild, slightly astringent. (According to Heim 1966).
Chemical reactions. Context immediately and strongly violet with gaiac; pink with pyramidon; strongly reddish purple with gaiacol; purple with phenol; weak greyish green with FeSO4; unchanging with NH3. (According to Heim 1966).
Basidiospores usually ellipsoid, rarely subglobose, 7.0–8.2–8.8–10.0(–10.4) × 5.5–6.0–7.3–7.8 mm (Q = 1.07–1.21–1.36–1.55; n = 20); ornamentation amyloid, forming an almost complete, but somewhat irregular reticulum; ridges up to 1.5 mm high and very acute; amyloid spots present in the meshes; plage not or slightly amyloid. Basidia 35–55 × 7–9 mm, cylindric to subclavate, 4-spored or 2-spored. Pleurolamprocystidia abundant, 50–65 × 7–10 mm, fusiform or irregular, thick-walled (up to 1 mm). Pleuropseudocystidia abundant, 4–7(–10) mm, cylindrical, with rounded and sometimes swollen apex, with dark greyish brown content. Lamellar edge sterile; marginal cells rarely clavate, mostly fusiform, 20–35 x 4–7 µm. Hymenophoral trama mixed; laticiferous hyphae present and abundant, with dark content; small sphaerocytes present. Pileipellis a palisade, 50–70 µm thick; elements of the suprapellis 25–40 × 5–9 mm, mostly fusiform, sometimes subcylindric, thin-walled; subpellis pseudoparenchymatous, with spherical cells 10–20 mm. Clamp-connections absent. – Fig. 91
Ecology. Sudanian woodland, the type was found near Uapaca guineensis.
Distribution. Known from the Central African Republic and Nigeria.
Revised specimens
Central African Republic. Savanna of Bébé, close to Filifi river, 6 Aug. 1966, R. Heim LM2189 (PC, holotype).
Nigeria. Cross River Prov., Obadei Ranch, 24 Apr. 1989, R. Nicholson 179 (K(M) 38486).
Notes. The type consists of one basidiome in a very bad condition.The microscopical description is based on the type and on the material from Nigeria which lacks field notes.
The species differs from Lactarius melanogalus by the ellipsoid spores which have a lower ornamentation.
Sect. Plinthogali (Burl.) Singer s.l.
Lactarius sect. Plinthogali (Burl.) Singer, Ann. Mycol. 40: 123 (1942).
Lactarius [unranked] Plinthogali Burl., Mem. Torrey Bot. Club 14: 83 (1908, as Plinthogalae).
Lactarius sect. Pulchrispermi Verbeken, Mycotaxon 77: 442 (2001).
− Type species: Lactarius plinthogalus (J. Otto) Burl.
Key to tropical African species
1. Carpophore sequestrate (gasteroid, secotioid) ................................................................................... 2
Carpophore with normally developed stipe and gills ......................................................................... 3
2. Fruitbody gasteroid, ochraceous yellow, hypogeous or semihypogeous; spores subglobose, 8–11 µm long; ornamentation highly winged, up to 3(3.5) µm, forming a subcomplete reticulum ........... 69. L. angiocarpus
Fruitbody epigeous, with normally developed stipe; hymenophore sinuate, glebulose to alveolate labyrinthoid .................................. 70. L. dolichocaulis
3. Spore‑ornamentation with a zebroid pattern ....................................................................................... 4
Spore‑ornamentation with a reticulate or spiny pattern ..................................................................... 9
4. Pleurocystidia abundant ......................................................................................................................... 5
Pleurocystidia absent .............................................................................................................................. 6
5. Young cap pale brown to greyish orange; context not turning reddish or pinkish; pleurocystidia emergent, with needle-like content; pileocystidia abundant ............... 71. L. pulchrispermus
Young pileus dark brown; context turning red to orange red when bruised; pileocystida absent ............. 72. L. miniatescens
6. Spores ellipsoid, ornamented with regular, rounded ridges, up to 1 µm high; pileus greyish brown to yellowish brown, 10–45 mm diam.; stipe hollow and fragile....................... 73. L. sulcatulus
Spore ellipsoid to globose, with ornamentation up to 2 µm high ..................................................... 7
7. Pileus > 4 cm diam., fuliginous brown; spore ornamentation consisting of coarse and parallel ridges ....... 76. L. kalospermus
Pileus small, < 2 cm, with yellowish to pale brown colours; spores with a spiny aspect ............. 8
8. Pileus with brown tinges; stipe with distinct hirsute base; spores 8–9.5 × 7–8.5 µm, partially reticulate ...... 74. L. adhaerens
Pileus pale yellow to warm yellow; base of stipe not hirsute; spores 7–8 × 5.5–6.5 µm, not reticulate ....... 75. L. desideratus
9. Context turning pink or reddish when bruised; spore ornamentation winged ............................ 10
Context not turning pink or reddish; spore ornamentation winged or spiny ............................... 12
10. Pleurocystidia present; pileus olivaceous brown ........................................................ 79. L. nudus
Pleurocystidia absent; pileus fuliginous brown to dark brown ..................................................... 11
11. Lamellar edge brownish; spore ornamentation 1(1.5) µm ............................... 77. L. congolensis
Lamellar edge concolorous; spores very highly ornamented with reticulate wings up to 2 µm high, distinctly verrucose between the ridges ................................................... 78. L. melanodermus
12. Lamellae distant ................................................................................................................................... 13
Lamellae (moderately) crowded .......................................................................................................... 14
13. Pileus pale greyish to brownish grey, margin not sulcate ............................. 80. L. rumongensis
Pileus warmer yellowish brown to dark brown, margin sulcate ............................. 81. L. sulcatus
14. Lamella edge coloured (brown) ......................................................................................................... 15
Lamella edge concolourous with lamellae ....................................................................................... 16
15. Spores 7.7–8.1 ´ 7.0–7.5 mm; pileipellis a hymeniderm, terminal elements cylindrical to clavate, up to 12 mm broad ............................................................................................. 82. L. saponaceus
Spores 5.8–6.5 ´ 5.1–5.9 mm; pileipellis a trichoderm to trichopalisade, terminal elements long and slender, up to 5 mm broad ....................................................................... 83. L. pusillisporus
16. Pileus covered with thick radial veins; stipe with a large collar of orange hairs around the base; spore ornamentation spiny(-zebroid); basidia partly 2-spored; pleuropseudocystidia branched and tortuous .......... 74. L. adhaerens
Not with this combination of characters; spore ornamentation lower, spiny or reticulate; terminal part of pleuropseudocystidia cylindric and broad; context often orange in stipe base .... (Pseudolignyoti ad int.) 17
17. Spore‑ornamentation up to 1 µm high, reticulate; species of woodlands .................................. 18
Spore‑ornamentation up to 1.5 mm high, spiny; carpophores slender; species growing in rain forest ..... 20
18. Pileus up to 4 cm diam., young dark brown, soon paler greyish, margin more or less crenulate; young lamellae contrastingly white with cap and stipe (reminding of L. lignyotus); context in stipe base never orange; terminal elements of the pileipellis cylindrical, long and slender, 10–40 ´ 6–10 mm ........ 86. L. tenellus
Pileus larger, remaining brown to dark brown; margin not crenulate; context in base of stipe orange; lamellae soon ochraceous orange ................................................................................ 19
19. Basidiocarp up to 5 cm high; pileus up to 7 cm diam.; terminal elements of the pileipellis obovoid to broadly clavate, 10–20 ´ 8–15 mm; common miombo woodland species, growing in the sparse litter of open patches ........ 84. L. kabansus
Basidiocarp up to 9 cm high; pileus up to 12 cm diam.; terminal elements of the pileipellis subcylindric, subfusiform, never rounded nor clavate, 30–65 × 8–10(12) µm; miombo woodland species preferring shade and thicker leaf litter ............................. 85. L. sciaphilus
20. Pileus cream with dark ochraceous brown squamules; pileus never with a papilla ........ 88. L. pseudolignyotus
Pileus dark brown, finely tomentose; pileus sometimes with a papilla .................... 87. L. acutus
Species descriptionS
69. Lactarius angiocarpus Verbeken & U. Eberh.
Eberhardt & Verbeken, Mycol. Res. 108: 1045 (2004). − Type: Zambia, Copperbelt Prov., about 15 km outside Kitwe off the Kitwe-Ndola highway, 28 Dec. 2000, D. Arora 00-448 (GENT, holotype).
Selected descriptions and icons. Eberhardt & Verbeken, Mycol. Res. 108: 1045–1047, fig. 1
(2004).
Basidiomata 14–26 mm diam., subglobose to irregular. Peridial surface smooth, dry, pale yellowish-tan; bruising reaction absent. Gleba pallid, cream, unchanging, loculate, chambered, with small (up to 1 mm) irregular locules. Stipe absent. Columella absent. Smell not distinctive. Taste mild. Latex abundant, white, unchanging, mild. – Pl. 43
Basidiospores subglobose, 8.1–9.7–11.2 × 7.8–9.3–10.8 µm (Q = 1.00–1.05–1.10, n = 20), mostly symmetric and orthotropic; ornamentation amyloid, highly winged, up to 3(–3.5) µm, forming a subcomplete reticulum, with some rare isolated warts present; hilar appendix 3–4 × 1.5–2 µm, bent; plage absent. Basidia 35–50 × 10–16 µm, 4-spored, hyaline, subcylindrical to subclavate, thin-walled; sterigmata 4–10 µm long. True cystidia absent. Pseudocystidia rare, not or slightly emergent, subcylindric or slightly tortuose, 4–7 µm diam., tapering near apex. Sterile cells (different from basidioles) present in some locules where basidia seem to be lacking or very scarce, hyaline, shortly cylidric to subclavate or irregular, sometimes branching, thin-walled, 13–30 × 5–10 µm. Peridiopellis a trichoderm; terminal elements thin-walled to slightly thick-walled and refringent, cylindric to subclavate, 10–20 × 3–6 µm, hyaline; trichoderm covered by a brownish, incrusted layer up to 25 µm thick; underlaying hyphae interwoven, hyaline, 3–6 (–8) µm diam. Hymenophoral trama 30–60 µm broad, composed of subparallel to interwoven, hyaline hyphae of 2–5 µm diam.; lactifers moderately abundant; some sphaerocytes sometimes present near subhymenium. – Fig. 92
Ecology. Collected in Zambezian miombo woodland.
Distribution. Only known from the type locality.
Revised specimens
Zambia. Copperbelt Prov., about 15 km outside Kitwe off the Kitwe-Ndola highway, along road going into Greystone, 28 Dec. 2000, D. Arora 00-448 (GENT, holotype).
Notes. A very similar taxon has been collected under Caesalpiniaceae on Mount Cameroon (Ludwig Beenken, pers. comm.).
70. Lactarius dolichocaulis (Pegler) Verbeken & U. Eberh.
Eberhardt & Verbeken, Mycol. Res. 108: 1047 (2004). − Arcangeliella dolichocaulis Pegler, Kew Bull. 37: 267 (1982); Gastrolactarius dolichocaulis (Pegler) J.M. Vidal, Rev. Catalana Micol. 26: 77 “2004” 2005. – Type: Zambia, Luapala Prov., Nchelenge, Kawambwa scarp, at base of old termite mound in open forest, 22 Dec. 1977, A.C. Rose 7701 (K(M) 34217, holotype).
Selected descriptions and icons. Pegler, Kew Bull. 37: 267–269 (1982); Verbeken, Persoonia 17: 404–405 (2001).
Pileus 40–70 mm diam., applanate to depressed, pale ochraceous buff to light buff, smooth, glabrous, dry; margin entire, expanded, undulate. Hymenophore exposed, sinuate, glebulose, somewhat radiating to alveolate labyrinthoid, forming numerous, small irregular locules without any geotropic orientation, concolorous with pileus; tramal plates thin. Stipe well-developed, 40–65 × 8–13 mm, elongate, cylindric or tapering below, solid, concolorous with the pileus, smooth, glabrous. Context up to 10 mm thick at the disk, very pale grey, unchanging, firm. Latex copious, white, unchanging; taste of cedar-wood with a bitter aftertaste. Spore-deposit not observed. (According to Pegler 1982).
Basidiospores 7.8–8.5–9.7 × 6.7–7.7–9.0 mm, subglobose (Q = 1.05–1.10–1.21); ornamentation amyloid, consisting of an almost complete reticulum with ridges up to 1.5 mm high; plage distally amyloid. Basidia 4-spored, clavate to subfusiform, 35–45 × 9–12 mm. Cystidia and pseudocystidia not observed; hymenophoral trama consisting of thin‑walled hyaline hyphae which are more or less parallel and embedded in some slimy consistence; lactifers scarcely present. Pileipellis and stipitipellis a palisade with a well developed layer of isodiametric cells and a suprapellis of thin‑walled, 1‑ to 3‑septate cylindrical elements. Clamp-connections absent. – Fig. 93
Ecology. Miombo woodland.
Distribution. Only known from the type locality.
Revised specimens
Zambia. Luapala Prov., Nchelenge, Kawambwa scarp, Dec. 1977, A.C. Rose 7701 (K(M) 34217, holotype).
Notes. Pegler (1982) has already stated that the basidiomes of Arcangeliella dolichocaulis are completely agaricoid apart from the glebulose hymenophore and that the taxon fits better in Lactarius than in the genus Arcangeliella as defined by Pegler & Young (1979). Microscopically, this species has similar characters to the species of Lactarius subgenus Plinthogalus because of the reticulate and high spore ornamentation with distally amyloid plage and the structure of the pileipellis. Further observation on fresh collections are needed to investigate the presence of pseudocystidia.
71. Lactarius pulchrispermus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 205 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 1 April 1994, A. Verbeken 94-554 (BR 57696–78, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 216–217 & Pl. 238–241 (1996d); Verbeken & al., Syst. Geogr. Pl. 70: 201–203 (2000).
Pileus 12–55(–80) mm diam., thin, elastic to brittle, planoconvex to applanate and then slightly depressed, sometimes with a small papilla, often asymmetric; margin straight, slightly crenulate, to very irregularly and strongly striate, rarely irregularly deeply grooved; pellis not dehiscent, dry, smooth or felty, finely squamulose in the centre, sometimes a bit greasy, chamois-leather-like, pale orange to greyish orange, brownish to pale brown (5A4, 5B4, 5C4–5, 5D5, 6B3, 6C4 or 6D5–7), hygrophanous, with darker and paler patches, darker when bruised, unchanging, yellow or orange-red with KOH. Lamellae adnate to subdecurrent, rarely decurrent, unequal with lamellulae (3–5(–7) lamellulae between two lamellae, regular short-long-short pattern), dense (L+l = 3+9/cm to 5+15/cm) to moderately spaced but then with very abundant lamellulae of different lengths (in a regular short-long-short pattern, often 7 or more between two lamellae), moderately narrow (1–4 mm), thin, paper-like, rather tough, without venation, pale brown (5B4 to 5C4) to brownish orange (6C3–4); edge more or less concolorous, entire. Stipe eccentric or central, (10–)15–40(–100) × 2–7 mm, cylindrical, slender, tapering downwards, rarely remarkably bulbous and irregularly swollen at the base; pellis felty and almost pruinose to smooth, pale brown to greyish orange (4A3–4, 4C3 to 5D4), darker to dirty brown and dark brown when bruised, chambered to fistulous. Context thin (1–1.5 mm) in pileus, thin-fleshy, elastic and sometimes irregularly hollow in stipe, cream, buff to brownish, unchanging or changing salmon to greyish; taste mild, sometimes slightly acrid after a while, sometimes menthol-like, sometimes astringent and acrid; smell not particular or slightly spicy. Latex scarce to rather abudant, white, changing brown, sometimes unchanging; taste mild, spicy or slightly acrid. Spore-deposit dark buff to pale brown. – Pl. 44
Chemical reactions. Context unchanging with FeSO4 and HCl; soon dark grey, brown to black with KOH; unchanging (Verbeken 94-466), yellow (Verbeken 94-554) or orange-red (Verbeken 94-517) with NH4OH.
Basidiospores globose to subglobose, (7.1–)7.3–7.9–8.0–8.7 × 6.8–7.5–7.7–8.3 µm (Q = 1.00–1.03–1.05–1.12, n = 60); ornamentation amyloid, zebroid, composed of ridges and some warts, up to 2 mm high; plage distally amyloid. Basidia 45–55 × 10–13 mm, subclavate, 4-spored. Pleurocystidia very abundant, 60–85 × 10–12 mm, cylindrical to subfusiform, with rounded apex, thin-walled; content needle-like. Pleuropseudocystidia abundant, often very emergent, 2–3 mm diam., cylindrical to tortuous or corkscrew-like, often branched; content oleiferic to guttate. Lamellar edge sterile; marginal cells 35–70 × 5–9 mm, cylindrical to fusiform, with tapering, mucronate or rounded apex, sometimes septate; wall slightly thinckened; content often needle-like. Hymenophoral trama irregular; composed of interwoven hyaline hyphae and abundant lactifers. Pileipellis trichoderm-like; terminal elements (20–)40–70(–90) × 6–15 mm, cylindrical, clavate or fusiform, slightly thick-walled; pileocystidia abundant, 40–70(–90) × 6–15 µm, clavate, thick-walled or thin-walled, with needle-like content. Stipitipellis idem, terminal elements less frequently with thickened wall; caulocystidia abundant, 40–60(–70) × 6–12 µm, mostly thin-walled, with dense needle-like content. Clamp-connections absent. – Fig. 94
Ecology. Zambezian miombo woodland, mostly under Brachystegia spp.
Distribution. Widespread in the Zambezian region. Known from Burundi, Democratic Republic of Congo, Zambia and Zimbabwe.
Revised specimens
Burundi. Bururi Prov., Rumonge Forest Reserve, Nyamirambo, 11 March 1994, A. Verbeken 94-072 (BR 57582–61) & 94-073 (BR 57583–62); Ibid., 18 March 1994, A. Verbeken 94-221 (BR 57629–11); Ibid., 25 March 1994, A. Verbeken 94-396 (BR 27748–06); Ibid., 30 March 1994, A. Verbeken 94-466 (BR 57672–54); Ibid., 31 March 1994, A. Verbeken 94-517 (BR 57687–69); Ibid., 1 April 1994, A. Verbeken 94-554 (BR 57696–78, holotype; GENT, isotype); Nkayamba, near Rumonge, 4 Dec. 1992, B. Buyck 4653 (PC).
Democratic Republic of Congo. Shaba region, N of Kipopo, Jan. 1972, D. Thoen 5339 (BR 66314–63).
Malawi. Northern Prov., Mzimba distr., Ntanga Forest Reserve, 30 Jan. 2005, A. Verbeken 05-160 (GENT); Ibid., 31 Jan. 2005, A. Verbeken 05-175 (GENT); Ibid., Perekezi Forest Reserve, 14 Feb. 2005, F. Noe 05/263 (GENT); Southern Prov., Machinga distr., Liwonde Forest Reserve, 22 Jan. 2005, A. Verbeken 05-010 & 05-014 (all GENT).
Zambia. Northern Prov., Mpika distr., North Luangwa National Park, Mwaleshi Camp, 9 Feb. 1995, D. Shah‑Smith 202 (K(M) 35515); Mbereshi National Forest, 22 Feb. 1998, priv. herb. M. Groenendaal L5.
Zimbabwe. Manicaland Prov., along Mutare-Bvumba road, near Prince of Wales Viewpoint, 11 Feb. 1999, A. Verbeken 99-168 (GENT); 15.5 km along Mutare-Bvumba road, 11 Feb. 1999, A. Verbeken 99-178 (GENT).
Notes. The description is based on collections from Burundi supplemented with fieldnotes of Zimbabwe.
72. Lactarius miniatescens Verbeken & Van Rooij
Van Rooij & al., Nova Hedwigia 77: 233 (2003). − Type: Benin, Atacora Prov., Bassila, 21 June 2000, A. De Kesel 2841 (BR 126394–03, holotype; GENT, isotype).
Selected descriptions and icons. Van Rooij & al., Nova Hedwigia 77: 233–234, 243 (2003).
Pileus 35–40 mm diam., almost applanate to planoconvex and deeply depressed; pellis mat, tomentose, indehiscent, sometimes dark blond (5D4), usually dark brown (6E4 to 6F3) to blackish brown, after a while fading to brownish grey (6C2 to 6F2); margin striate to 10–13 mm. Stipe 40–45 × 6–8 mm, regularly cylindric, slender and straight, solid; pellis tomentose, pruinose, whitish to greyish brown (5A3), becoming more whitish at the base. Lamellae adnate, slightly decurrent, seldomly forked, distant (L+l = 10/cm), lamellulae 1/L with an irregular pattern (sometimes 1 l/l to 3 l/L, more or less in series), cream (4A3–4). Context very thin in pileus, turning red to orange red (8A6–8 to 8B6–7), especially under the stipiti-and pileipellis, remarkably intense in younger specimens, faint or lacking in older specimens, unchanging in the center; taste unremarkable, faint to strongly fungoid; smell unremarkable. Latex white, only observed in young specimens. Spore-deposit pale yellow to light yellow (4A3–4). – Pl. 45
Basidiospores globose, sometimes subglobose to broadly ellipsoid, 7.0–7.7–8.4–9.1 × 6.8–7.5–8.1–8.7 µm (n = 80, Q = 1.00–1.03–1.04–1.15); ornamentation amyloid, winged, composed of ridges, up to 2 µm high, almost completely zebroid, only few ridges branched; ridges often with a split aspect; isolated warts and short, lower ridges abundant between the higher wings; plage distinctly distally to almost totally amyloid. Basidia (55–)67–73 × 12–15 µm, clavate to subclavate, 4-spored (seldom 2-spored), slightly thick-walled (0.5–1 µm); sterigmata long, 6–9 × 2–4 µm; content mostly (strongly) guttate. Pleuromacrocystidia abundant, 50–70 × 9–13 µm, fusiform to irregularly cylindric, apex often moniliform or mucronate, content granular. Pleuropseudocystidia abundant, emergent, 2–6 µm diam., irregularly cylindric to tortuous, content granular, sometimes needle-like. Lamellar edge sterile; marginal cells 13–16 × 2–5 µm, cylindric to clavate, thin-walled to slightly thick-walled, hyaline. Hymenophoral trama cellular with abundant dark coloured lactifers. Pileipellis a hymeniderm to trichoderm; terminal elements in the suprapellis either short and clavate, 13–23 × 10–15 µm, or longer and subcylindric to subfusiform, 20–65 × 4–7(–10) µm, with brown intracellular pigmentation; subpellis thin, composed of short cylindrical to subglobose cells. Stipitipellis idem, terminal elements more frequently with slightly thickened walls. Clamp-connections absent. – Fig. 95
Ecology. Found in denser riverine forest dominated by Berlinia grandiflora, Uapaca somon, Lonchocarpus sericeus and Pterocarpus erinaceus.
Distribution. Known from Benin and Togo.
Revised specimens
Benin. Donga Prov., Bassila, 21 June 2000, A. De Kesel 2841 (BR 126394–03, holotype; GENT, isotype); Ibid., 10 Jun. 2002, A. De Kesel 3335 (BR 152237–44); Ibid., 10 Jun. 2002, A. De Kesel 3336 (BR 152236–43); Ibid., 17 Jun. 2004, A. De Kesel 3659 (BR 157048–05); Atacora Prov., Kota, 23 Sept. 2000, A. De Kesel 2918 (BR 129129–22).
Togo. Central Prov., Tchamba, 13 Jul. 2007, A. De Kesel 4338 (BR 163828–92).
Notes. This species resembles Lactarius nudus R. Heim, from which it differs macroscopically by the sharp contrast between its dark coloured cap and whitish stipe, as well as the more distant spacing of the lamellae. Microscopically, the ornamentation of the spores is rather zebroid and the hymeniderm is composed of rather rounded to clavate elements.
73. Lactarius sulcatulus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 210 (1996). − Type: Democratic Republic of Congo, Kivu, Irangi, April 1972, J. Rammeloo Z414 (GENT, holotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 179–180 & Pl. 152–153 (1996d); Verbeken, Persoonia 17: 387, 389–390 & Pl. 4 (2001).
Pileus 10–45 mm diam., very thin, planoconvex, applanate to broadly infundibuliform, with very distinct and acute papilla in the centre; margin thin, crenulate, undulate, slightly grooved, fimbriate to serrate in older specimens; pellis not dehiscent, dry, mat, wrinkled, with very fine radial venation superposed on the radial grooves which start in the centre, greyish yellow (4B4 to 4C4) to yellowish brown (5E5 to 5F5), light brown to yellowish brown (5D4 to 5E4) in the centre; papilla and extreme margin darker brown (5F6–8). Stipe 15–23 × 4–6 mm, cylindrical to very flattened and with a longitudinal fold in it, slightly tapering downwards, slender, mat, slightly felty, yellowish white to pale yellow (3A2–3 to 4A2–3), firm, solid. Lamellae broadly adnate to subdecurrent, unequal with abundant lamellulae of different lengths, distant (6/cm), rather broad (2–4(–5) mm), thin, rarely anastomosing, yellowish white (3A2) when young, cream when older (3A2 to 4A2); edge entire, concolorous. Context thin, whitish, usually regularly fistulose, slightly changing to reddish brown in the stipe, in older specimens reddish in the centre of the stipe; smell papaya‑like; taste agreeable, mild, a bit spicy. Latex present, scarce, watery, whitish to transparent, unchanging or changing slightly brownish; taste papaya‑like. Spore‑deposit pale coloured, not white. – Pl. 46
Chemical reactions. Context unchanging with NH4OH, NaOH, anilated H2O, phenol and FeSO4; reddish brown after a while with phenolaniline.
Basidiospores ellipsoid, 7.5–8.1–8.6 × 6.0–6.6–7.0(–7.2) mm (Q = 1.16–1.22–1.31, n = 20); ornamentation amyloid, zebroid, composed of ridges, mostly parallel, seldom branched; some isolated elongated warts present, ridges up to 1 mm high; plage inamyloid or with distal amyloid spot. Basidia 50–55(–60) × 9–11 mm, subcylindrical to subclavate, 4‑spored. Pleurocystidia absent. Pleuropseudocystidia abundant, mostly emergent, 10–13(–15) mm diam. in the upper part, cylindrical to clavate, with rounded apex, thin‑ walled or slightly thick‑walled; content needle‑like and oleiferic. Lamellar edge sterile; marginal cells 15–25 × 4–7 mm, cylindrical to irregularly fusiform, thin‑walled, hyaline. Hymenophoral trama irregular, composed of lactifers and hyaline, thin‑walled hyphae. Pileipellis hymeniderm‑like, two‑ layered, covered by a slime‑layer; terminal elements of suprapellis clavate to broadly clavate, 10–20 × 5–12 mm, thin‑walled; subpellis thin, composed of isodiametric cells of 10–15(–20) mm diam. Stipitipellis between a trichoderm and a hymeniderm; elements of the suprapellis 10–50 × 5–8 mm, cylindrical, with rounded apex, sometimes septate; subpellis rudimentary, composed of a few, small, isodiametric cells of 5–10(–20) mm diam., mixed with long and slender hyaline hyphae. Clamp‑connections absent. – Fig. 96
Ecology. Found in wetter Zambezian miombo woodland and Guineo‑Congolian rainforest.
Distribution. So far, only known from the Democratic Republic of Congo and Zimbabwe.
Revised specimens
Democratic Republic of Congo. Kivu Prov., Irangi, April 1972, J. Rammeloo Z414 (GENT, holotype).
Zimbabwe. Manicaland Prov., Chipinge, Busi farm, around Scott’s house, 12 Feb. 1999, A. Verbeken 99-187 (GENT); Penhalonga, hills behind Harris’ garden, 9 Feb. 1999, A. Verbeken 99-144 & 99-157 (all GENT).
74. Lactarius adhaerens R. Heim
Heim, Candollea 7: 375 (1938). – Type: Madagascar, South of Antanambé, 20 Dec. 1934, R. Heim J18 (PC, holotype).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 32–36 & Pl. 1b (1938); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 189 & Pl. 178–181 (1996d).
Pileus 40–45 mm diam., convex, then applanate and deeply depressed, infundibuliform; margin slightly incurved; pellis not dehiscent, slightly wrinkled to subtomentose, ochraceous orange near the margin, darker in the centre, with thick radial veins and anastomosing small veins; veins branched near the margin, becoming translucid and crenulate near the extreme margin. Lamellae decurrent with long tooth, unequal with lamellulae of three different lengths, distant, moderately broad (3–4.5 mm), thin, waxy, tough, brownish orange; edge entire, concolorous. Stipe 35 × 4–6 mm, 10 mm diam. at the base, long, slender, regularly cylindrical, subbulbous, smooth, brownish red, concolorous with the pileus, paler at the base, with a large collar of orange hairs around the base, solid or slightly fistulous in the middle. Context tough, compact, whitish in the pileus, cream in the stipe, unchanging. Latex very abundant, white, unchanging; taste astringent, mild. Spore-deposit probably whitish cream. (According to Heim 1938b). – Pl. 46
Chemical reactions. Context unchanging with gaiac and gaiacol. (According to Heim 1938b).
Basidiospores subglobose to ellipsoid, (7.1–)8.0–8.8–9.5(–10.7) × (6.5–)6.9–7.5–8.2(–8.8) mm (Q = 1.05–1.17–1.28, n = 30); ornamentation amyloid, composed of narrow ridges, up to 2 mm high, some isolate, some connected, forming a partial reticulum; wall rugose, rugged with lower ridges; plage amyloid in distal part. Basidia 40–50 × 10–15 mm, clavate to ventrose, 2-spored (sterigmata 10–15 × 2–3 mm) or 4-spored (sterigmata 5–10 × 1–2(–3) mm); content slightly granular. Pleurocystidia absent. Pleuropseudocystidia abundant, emergent, 3–5 mm diam., tortuous to corkscrew-like, often branched; wall locally slightly thickened; content oleiferic. Lamellar edge sterile; marginal cells (8–)10–20(–25) × 3–6 mm, cylindrical, conical, tapering towards the apex, thin-walled or slightly thick-walled, hyaline. Hymenophoral trama irregular, composed of narrow, hyaline hyphae and abundant lactifers; sphaerocytes absent. Pileipellis subcellular to a hymeniderm, composed of isodiametric to irregular cells which are (5–)7–15(–25) mm diam.; terminal elements slightly differentiated, rather subclavate or subfusiform. Stipitipellis similar; terminal elements rather cylindrical. Clamp-connections absent. – Fig. 97
Ecology. Found in Malagasy lowland rainforest.
Distribution. Only known from the type locality.
Revised specimens
Madagascar. Betsimisaraka Prov., South of Antanambé, 20 Dec. 1934, R. Heim J18 (PC, holotype).
Notes. The type-specimen consists of one basidiome in rather poor condition.
Heim (1938b) classified this species into the group of his Lactarius pandani. There might be a superficial resemblance, especially in the macroscopic characters, but the microscopic characters are totally different and similar to those of Lactarius acutus.
75. Lactarius desideratus Verbeken & Stubbe
Verbeken & al., Cryptog. Mycol. 29: 138 (2008). − Type: Cameroon, forêt de Dja, 30 Jan. 1993, A. Verbeken 07-47 (GENT, holotype).
Selected descriptions and icons. Verbeken & al., Cryptog. Mycol. 29: 138-140 (2008).
Pileus 9–20 mm diam., planoconvex, flattened or slightly depressed in the centre; surface striate-grooved, with soft pubescent reticulate veins in the centre and some veins also present near the margin, soft velvety-pubescent, somewhat opaque, pale yellow to pale yellowish brown or orange yellow (3A4 to 4B6), darker in young specimens (up to 5D8 but yellower). Lamellae broadly adnate with slightly decurrent tooth, distant (L+l = 7+21 halfway pileus), with 3 lamellulae between 2 lamellae, with a regular short-long-short-pattern, yellowish white (4A2), rather thick, 1–3 mm broad; edge entire, concolorous. Stipe 8–14 mm × 1–2 mm, subcylindrical, tapering downwards, greyish yellow (4B5 to 4C6) or somewhat paler yellow, very slightly felty. Context very thin-fleshed, hollow in stipe, dirty pale yellow, unchanging; taste agreeable; smell not remarkable. Latex watery white, not abundant, unchanging, unchanging when isolated on glass, but changing to dirty yellow when isolated on white paper. – Pl. 47
Chemical reactions. Context unchanging with FeSO4.
Basidiospores mostly broadly ellipsoid, sometimes subglobose, 6.5–7.2–7.8 × 5.7–6.3–6.4 µm (Q = 1.07–1.15–1.23, n = 20); ornamentation amyloid, strongly zebroid, composed of rather narrow ridges up to 1.5(2) µm high, arranged in a parallel pattern; abundant short ridges and warts present; plage distally amyloid. Basidia 35–55 × 10–11 µm, (2)4-spored, subclavate or somewhat irregular. Pleurocystidia absent. Pleuropseudocystidia rather abundant, slightly emergent, 4–6 µm diam., cylindrical to slightly tortuous, with slightly refringent walls. Lamellar edge sterile; marginal cells 28–52 × 4–6 µm, subcylindrical or subfusiform, often irregular or with some bulges, very thin-walled, hyaline to slightly granular. Lamellar trama mixed, with hyaline hyphae, small sphaerocytes and lactifers. Pileipellis a palisade; subpellis thin, 1 or 2 cell-layers thick; terminal elements of two types; first type subclavate or clavate, sometimes almost spherical, 15–25 × 5–15 µm, thin-walled or distinctly thick-walled; second type long cylindric and slender, 30–50 × 4–6 µm, always thin-walled. Stipitipellis similar, a thin palisade with two types of elements. – Fig. 98
Ecology. Guineo-Congolian rainforest. Found under Uapaca guineensis.
Distribution. Only known from the type locality.
Revised specimens
Cameroon. Eastern Prov., Réserve de faune du Dja, 10 April 2007, A. Verbeken 07-47 (GENT, holotype); Ibid., 11 April 2007, A. Verbeken 07-63 (GENT).
Notes. L. desideratus is most closely related to L. adhaerens R. Heim. Lactarius adhaerens differs macromorphologically by the darker and more ochraceous orange colours of the pileus and the large collar of orange hairs at the base of the stipe. The stipe is also brownish red, while our species in Cameroon has a pale yellow and smooth stipe. Microscopically, L. adhaerens differs by the larger spores (8.0–8.8–9.5(–10.5) × 6.9–7.5–8.2 mm), also ornamented with narrow and high ridges but in a less zebroid and rather subreticulate pattern.
There is a superficial resemblance with Lactarius uapacae, a species that was growing together with L. desideratus, and has comparable small and yellowish fruitbodies, but the microscopical features are completely different.
76. Lactarius kalospermus (Beeli) Verbeken & Walleyn
Verbeken, Edinburgh J. Bot. 53: 69 (1996). − Laccaria kalosperma Beeli, Bull. Soc. Roy. Bot. Belg. 66: 22 (1933). − Type: Democratic Republic of Congo, Binga, 20 Dec. 1929, M. Goossens-Fontana 859 (BR 7718–55, holotype).
Selected descriptions and icons. Verbeken, Edinburgh J. Bot. 53: 69-72 (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 178-179 & Pl. 149-151 (1996d).
Pileus 45–65 mm diam., firm, fleshy, convex, then applanate, with papilla when young, depressed in the centre when older; margin striate; pellis not dehiscent, smooth, radially plicate, with reticulate pattern between the grooves, fuliginous brown. Lamellae adnate to decurrent, unequal with lamellulae of different length, distant, yellowish orange to ferruginous brown; edge entire, concolorous. Stipe 50 × 5 mm, cylindrical, fuliginous brown, paler at the base. Context firm, solid, whitish, yellowish in stipe. Latex not observed. Spore-deposit not observed. – Pl. 48
Chemical reactions. Context immediately blue-green with gaiac; slowly brown with phenol; unchanging with FeSO4, NH3, Sulfovanilline, Pyrol and a-naphtol.
Basidiospores globose to subglobose, 7.7–8.3–9.0 × 7.4–7.9–8.5 mm (Q = 1.00–1.04–1.12; n = 20); ornamentation amyloid, composed of coarse and parallel ridges forming a zebroid pattern, sometimes even spiral-like; ridges < 2 mm high, narrow or broader and then often splitting; isolated fine lines also present; plage sometimes distally amyloid, sometimes verrucose. Basidia 50–65 × 10–14 mm, clavate, 4-spored; content needle-like or slightly granular. Pleurocystidia absent. Pleuropseudocystidia very abundant, cylindrical to irregularly tortuous towards the apex, often branched; content oleiferic, guttate. Lamellar edge sterile; marginal cells 10–20 × 3–5 mm, cylindrical, sometimes fusiform and mucronate; pseudocystidia also present. Hymenophoral trama subregular, composed of hyaline hyphae and abundant laticiferous hyphae. Pileipellis subcellular to hymeniderm-like, two-layered; terminal elements 14–40 × 5–9 mm, cylindrical to clavate or fusiform, often septate, content brown; with a rudimentary pseudoparenchymatic subpellis. Stipitipellis with longer, narrower and more regularly cylindrical elements than the pileipellis, 15–30 × 4–5 mm; without a pseudoparenchymatic layer. Clamp-connections absent. – Fig. 99
Ecology. This species has been found in drier Guineo‑Congolian rainforest with Gilbertiodendron dewevrei.
Distribution. Only known from Democratic Republic of Congo.
Revised specimens
Democratic Republic of Congo. Equateur Prov. Binga, Dec. 1929, M. Goossens-Fontana 859 (BR 7718–55, holotype); Ibid., Jan. 1936, M. Goossens-Fontana 1031 (BR 12316–94).
Notes. The type-material consists of a single basidiome and is, although crumbled, in rather good condition. Goossens-Fontana 1031 is in very poor condition but the striking resemblance of the water-colour and the spore characters with those of the type remove any doubt that they are conspecific.
The description is based on the type-material, completed with field notes and macrochemical observations on Goossens-Fontana 1031.
The species is best recognized by its zebroid spores. Similar characteristic very high and dense spore-ornamentation is rare in African representatives of the genus.
Heinemann (1946) was the first to indicate that Laccaria kalosperma is a Lactarius species belonging to the "Pterospori" (Lactarius sect. Plinthogali). It was, however, not included in Heim's studies (1955a, 1955b) of tropical African Lactarius species.
77. Lactarius congolensis Beeli
Beeli, Bull. Soc. Roy. Bot. Belg. 60: 164 (1928). – Type: Democratic Republic of Congo, Diobo Akuba, Dec. 1925, M. Goossens-Fontana 528 (BR 7714–51, holotype).
Lactarius craterelloides R. Heim & Gooss.‑Font. in Heim, Bull. Jard. Bot. Etat Bxl 25: 52 (1955). – Type: Democratic Republic of Congo, Binga, anno 1942, M. Goossens-Fontana 2081 (BR 7729–66, holotype).
Lactarius unicolor Gooss.‑Font. & R. Heim in Heim, Bull. Jard. Bot. Etat Bxl 25: 77 (1955). – Type: Democratic Republic of Congo, Binga, Oct. 1928, M. Goossens-Fontana 881 (BR 7719–56, holotype).
Excl.: Lactarius congolensis ss. Degreef (1990, = Lactarius kabansus); ss. Nzigidahera (1993, = Lactarius sp.).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: Pl. 3, fig. 4 & Pl. 5, fig. 4a–b (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 14, fig. 9 & Pl. 15, fig. 4 (1955); Verbeken, Edinburgh J. Bot. 53: 63–65 (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 185–186 & Pl. 166–167, 269 (1996d).
Pileus 40–120 mm diam., firm, convex, then deeply depressed; margin incurved when young, not striate; pellis not dehiscent, smooth to finely velutinate, fuliginous brown to dark brown with locally blackish tinge. Lamellae subdecurrent, unequal with irregular lamellulae, crowded (20/cm), narrow, thin, white to cream; edge brownish. Stipe 40–80 × 10–20 mm, cylindrical to subfusiform, smooth to finely velutinate, dark fuliginous ochre to brown. Context thick, white, changing reddish to brownish; taste mild. Latex abundant, opaque, white, unchanging; taste mild. Spore-deposit white. (According to Heim 1955a). – Pl. 48
Chemical reactions. Context slowly greyish blue with gaiac; brown-purple with phenol; unchanging with FeSO4, NH3, aniline and a-naphtol. (According to Heim 1955a).
Basidiospores subglobose to ellipsoid, 6.9–8.1–9.3 × 5.6–6.4–7.4 mm (Q = 1.10–1.26–1.51; n = 30); ornamentation amyloid, composed of a reticulum, without distinct isolated warts; ridges 1(–1.5) mm high; plage inamyloid. Basidia 35–45 × 10–11 mm, cylindrical to clavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia moderately abundant, 2–4(–5) mm diam., cylindrical to irregularly tortuous. Lamellar edge sterile; marginal cells 25–40 × 4–9 mm, irregularly tortuous, often capitate, sometimes obtuse; content brownish; pseudocystidia moderately abundant. Hymenophoral trama irregular, composed of hyaline hyphae and abundant lactifers, no sphaerocytes. Pileipellis a hymeniderm, one-layered; terminal elements 25–40 × 3–4(–7) mm, narrowly cylindrical to slightly tapering upwards, with intracellular brownish pigmentation. Stipitipellis with larger cells than the pileipellis, 15–35 × 6–8 mm. Clamp-connections absent. – Fig. 100
Ecology. Drier Guineo-Congolian rainforest with Gilbertiodendron dewevrei.
Distribution. So far, only known from Democratic Republic of Congo.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Diobo Akuba, Dec. 1925, M. Goossens-Fontana 528 (BR 7714–51, holotype); Binga, 31 Aug. 1929, M. Goossens-Fontana 837 (BR 12317–95); Ibid., Oct. 1928, M. Goossens-Fontana 881 (BR 7719–56); Ibid., 1942, M. Goossens-Fontana 2081 (BR 7729–66).
Notes. The synonymy of Lactarius craterelloides with L. congolensis was already suggested by Heim himself. The type-material of Lactarius congolensis consists of two specimens in rather poor condition. Goossens-Fontana 2081 consists of four rather poorly preserved specimens (mixed with two different Russula specimens) agreeing in all characters with the material of L. congolensis. The description is based on both collections.
According to Beeli (1928) this species is consumed by local populations.
78. Lactarius melanodermus R. Heim & Gooss.‑Font.
Heim, Bull. Jard. Bot. Etat Bxl 25: 50 (1955). – Type: Democratic Republic of Congo, Binga, Feb. 1947, M. Goossens-Fontana 4039 (BR 7734–71, holotype).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 50–52 & Pl. 5, fig. 3a–b (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 14, fig. 7 (1955) Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 162–165 & Pl. 184–185 (1996d).
Pileus 20–60 mm diam., convex, then depressed, irregular; margin thin, slightly involute, striate to grooved when older (up to midradius); pellis not dehiscent, finely tomentose, fuliginous brown, whitish pruinose when dry. Lamellae adnate to subdecurrent with decurrent tooth when older, unequal with lamellulae of same length, distant (3+2/cm), broad, thick, white, finally greyish; edge entire, (cream and pruinose on the exsiccatum). Stipe 40–60 × 5–10 mm, cylindrical, slightly subbulbous at the base, fuliginous brown, paler greyish when dry, firm. Context thin in the pileus, subtranslucid, white, then pale grey, changing pink when cut or bruised; taste plumbeous; smell not remarkable. Latex almost transparent then opaque and white; taste "acrid". Spore-deposit white. (According to Heim 1955a). – Pl. 49
Basidiospores subglobose to ellipsoid, (7.2–)7.8–8.5–9.2(–9.8) × (6.4–)6.8–7.3–7.8(–8.4) mm (Q = 1.01–1.16–1.24; n = 30); ornamentation amyloid, composed of ridges forming a complete reticulum; ridges narrow, up to 2 mm high, spore-wall between the ridges slightly verrucose but without distinct isolated warts; plage inamyloid. Basidia 49–65 × 8–17 mm, cylindrical to clavate, 4-spored. Pleurocystidia absent. Pleuropseudocystidia 4–6 mm diam., cylindrical to irregularly tortuous, obtuse, often slightly subterminally restricted; content densely oleiferic, sometimes granular or needle-like. Lamellar edge sterile, marginal cells cylindrical to flexuous, clavate or conical, 18–37 × 5–9 mm; content slightly granular; pseudocystidia present, often branched. Hymenophoral trama irregular, composed of hyaline hyphae and abundant lactifers, no sphaerocytes. Pileipellis hymeniderm-like, 50–70 mm thick, composed of isodiametric to irregular cells, 10–20 mm diam., forming erected chains, with a dark brown intracellular pigmentation; terminal fusiform or cylindrical cells sometimes present; extremities of laticiferous hyphae present. Stipitipellis as pileipellis; cells more elongate. Clamp-connections absent. – Fig. 101
Ecology. Found terrestrial or on dead wood in lowland rainforest, drier Guineo‑Congolian rainforest.
Distribution. Known from Cameroon, Democratic Republic of Congo and Gabon.
Revised specimens
Cameroon. South Western Prov., near Mundema, Korup National Park, 1984, I. Alexander 6 (E).
Democratic Republic of Congo. Equateur Prov., Binga, April 1928, M. Goossens-Fontana 703 (BR 12322–03); Ibid., Feb. 1947, M. Goossens-Fontana 4039 (BR 7734–71, holotype).
Gabon. Research Station of Ipassa-Makokou, 24 March 2005, M. Nguele 22 (BR 164548–36).
Notes. The microscopic description is based on Goossens-Fontana 703.
The species is macroscopically recognized by the colour-change of the context and the very dark tomentose pileus. It looks like Lactarius congolensis but differs from it by the grooved pileus-margin and the concolorous lamellar edge.
Alexander 6 shows spores which are slightly smaller and the ornamentation is up to 3.5 mm high. The spores measure 6.8–7.7–9.0 × 6.5–7.0–7.6 mm, Q = 1.03–1.07–1.15. The specimen is in better condition than Goossens-Fontana 703. A section through the pileipellis shows a regular palisade ("virescens"-like structure), with small cylindrical elements on the layer of isodiametric cells. The macroscopic characters are similar.
Heim, Bull. Jard. Bot. Etat Bxl 25: 53 (1955). – Type: Democratic Republic of Congo, Binga, Jan. 1935, M. Goossens-Fontana 1006 (BR 7725–62, holotype).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 53–56 & Pl. 5, fig. 5a–c (1955); Fl. Icon. Champ. Congo 4 & Pl. 14, fig. 8 (1955) Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 186–187 & Pl. 167–169 (1996d).
Pileus 15–50 mm diam., convex, then applanate, finally depressed; margin whitish, slightly recurved, striate with radiate veins; pellis not dehiscent, dark olivaceous, smooth, wrinkling in the centre, granular when dry. Lamellae adnate with decurrent tooth, unequal with lamellulae of two different lengths, dense, rather thin, 3.5 mm broad, white to cream, staining reddish; edge entire, concolorous. Stipe 30–45 × 5–8 mm, elongate, slender, irregularly cylindrical, tapering downwards, ochraceous to olivaceous in the lower part, paler in the upper part, smooth, irregularly fistulous. Context firm, white, changing red when bruised; taste "acrid". Latex abundant, white; taste very acrid. Spore-deposit white. (According to Heim 1955a). – Pl. 49
Chemical reactions. Context strongly reacting with gaiac (colour not indicated); brown with pyrogallol; wine-colour with phenol; unchanging with FeSO4, NH3 and aniline. (According to Heim 1955a).
Basidiospores globose to subglobose, sometimes broadly ellipsoid, (7.1–)7.6–8.2–8.8(–9.9) × (6.5–)7.0–7.6–8.1(–8.5) mm (Q = 1.03–1.09–1.14; n = 30); ornamentation amyloid, composed of ridges forming a partial reticulum; ridges up to 2 mm high, sometimes composed of narrower, parallel ridges, irregularly shaped and sized, warts present; spore-wall verrucose between the ornamentation; plage inamyloid. Basidia 45–56 × 12–14(–17) mm, clavate to ventricose, 4-spored; content slightly granular. Pleurocystidia 40–65 × 9–12 mm, irregularly clavate to fusiform, with obtuse apex, mucronate to irregular, sometimes forked; content granular. Pleuropseudocystidia scarce, cylindrical, 2–4(–5) mm. Lamellar edge sterile, marginal cells cylindrical to clavate, septate, 15–35 × 4–8 mm; content slightly granular. Hymenophoral trama irregular, composed of hyaline hyphae and abundant laticiferous hyphae. Pileipellis a trichoderm, one-layered, 55–75 mm thick; terminal elements cylindrical and obtuse to conical, tapering upwards, sometimes with a slightly granular content, 23–40 × (3–)4–7(–8) mm, thin-walled. Stipitipellis similar; terminal elements 30–45 × 3–8 mm. Clamp-connections absent. – Fig. 102
Ecology. Found in drier Guineo-Congolian rainforest with Gilbertiodendron dewevrei.
Distribution. Only known from the type locality.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Binga, Jan. 1935, M. Goossens-Fontana 1006 (BR 7725–62, holotype).
Notes. The type consists of one full-grown and two young basidiomes, in rather poor condition.
Heim (1955a) mentioned that he observed two-spored basidia. We have not observed them, which is probably due to the poor condition of the material. The elements of the hymenium needed a long KOH-treatment before swelling.
Heim described (1955a) the cystidia as being directly connected with laticiferous hyphae. We observed cystidia with the same size and shape as those Heim described, but they are not connected with lactifers though the content is granular and they often arise deep in the hymenium. They are true pleurocystidia.
80. Lactarius rumongensis Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 208 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 7 March 1994, A. Verbeken 94-006 (BR 57558–37, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 180–181 & Pl. 154–157 (1996d); Verbeken, Persoonia 17: 385–386 (2001).
Pileus 25–75 mm diam., thin, planoconvex to applanate; margin first regular, then irregular and strongly crenulate, sometimes grooved and striate; pellis not dehiscent, smooth in the centre, concentrically wrinkling towards the margin, with remarkable veins forming a reticulum at the margin, yellowish white, greyish yellow to brownish grey (4A2, 4B3, 5C2–3 to 5D2–3), paler towards the margin, hygrophanous. Lamellae adnate, unequal with lamellulae (1 to 3 between 2 lamellae, regular pattern), very distant (L+l = 3+5 to 3+6/cm), 4–10 mm broad, elastic, paper‑like, with remarkable venation when older, white to creamy; edge entire, concolorous. Stipe 15–50 × 5–10 mm, cylindrical, subbulbous at the base, sometimes tapering downwards, long and slender, dry, smooth, slightly darker than pileus, brownish grey, paler towards the base, solid, firm. Context thin, firm, whitish; taste mild, smell pleasant. Latex not abundant to quite abundant, white to water‑like; taste mild. Spore‑deposit dark cream to light brown. – Pl. 49
Chemical reactions. Context unchanging with FeSO4.
Basidiospores subglobose, 7.4–8.4–8.8–9.8(–10.0) × 6.8–7.8–8.0–8.9 mm (Q = 1.01–1.08–1.09–1.17; n = 60); ornamentation amyloid, forming a complete reticulum, ridges 0.5–1 mm high; plage sometimes distally amyloid. Basidia 40–60 × 10–12 mm, cylindrical to clavate, 4‑spored, exceptionally 2‑spored. Pleurocystidia absent. Pleuropseudocystidia abundant, 4–6 mm diam., cylindrical to tortuous, with rounded apex. Lamellar edge sterile; marginal cells 25–40 × 4–5(–6) mm, cylindrical to fusiform, septate, sometimes branched. Hymenophoral trama subregular to irregular, with hyaline hyphae and laticiferous hyphae. Pileipellis a hymeniderm; elements 10–20 × 3–5 mm, fusiform, sometimes clavate, sometimes thick‑walled; terminal elements of laticiferous hyphae present, not abundant. Stipitipellis a regular hymeniderm; elements 13–25 × 3–6 mm, cylindrical to clavate, sometimes slightly thick‑walled. Clamp‑connections absent. – Fig. 103
Ecology. Miombo woodland, (?) drier Guineo‑Congolian rainforest with Gilbertiodendron dewevrei.
Distribution. Known from Burundi, Democratic Republic of Congo and Malawi.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, May 1993, B. Buyck 5103 & 5135 (all PC); Ibid., March 1994, A. Verbeken 94-006 (BR 57558–37, holotype; GENT, isotype), 94-018 (BR 57568–47), 94-065 (BR 57576–55), 94-067 (BR 57578–57), 94-068 (BR 57579–58), 94-074 (BR 57584–63), 94-077 (BR 57587–66), 94-106 (BR 57596–75), 94-107 (BR 57597–76), 94-216 (BR 57624–06), 94-294 (BR 57640–22), 94-437 (BR 57663–45), 94-519 (BR 57689–71) & 94-469 (BR 57675–57); Ibid., 1 April 1994, A. Verbeken 94-572 (BR 57704–86); Nkayamba, just N of Rumonge, 1 March 1993, B. Buyck 4987 (PC).
Democratic Republic of Congo. Equateur Prov., Binga, 10 Nov. 1928, M. Goossens-Fontana 808 (BR 4951–04).
Malawi. Southern Prov., Machinga distr., Liwonde Forest Reserve, 22 Jan. 2005, A. Verbeken 05-013 (GENT).
81. Lactarius sulcatus Verbeken & Walleyn
Verbeken, Persoonia 17: 390 (2001). − Type: Zimbabwe, road from Mutare to Bvumba at peg 15.5 km, 11 Feb. 1999, A. Verbeken 99-176 (GENT, holotype).
Selected descriptions and icons. Verbeken, Persoonia 17: 390–392 & Pl. 5 (2001).
Pileus 45–55 mm diam., slightly planoconcave; margin directed slightly upwards in mature specimens, strongly crenulate, deeply and densily grooved (up to 1 cm long); surface velvety, chamois-leather-like, yellowish brown to brown (5E5 to 6E4), slightly darker in the centre and in the grooves. Stipe 20–40 × 7–11 mm, almost regularly cylindric, tapering at the base, smooth, soft, concolorous but paler, paler to whitish at the top and at the base. Lamellae broadly adnate, distant (4–5/cm halfway radius), with abundant lamellulae (often in a regular pattern), whitish (4A2), papery thin, not very brittle; edge even, concolorous. Context thin-fleshed near margin, moderately thick in pileus centre, solid in stipe, whitish, slightly greyish-brown in older specimens; taste mild; smell not particular. Latex not abundant, white, unchanging; taste mild. Spore-deposit not observed. – Pl. 49
Chemical reactions. Context unchanging with gaiac and FeSO4.
Basidiospores globose to subglobose, 7.2–8.1–8.9 × 7.0–7.4–8.0 µm (Q = 1.01–1.07–1.12, n = 20); ornamentation composed of rounded ridges up to 1.5 µm high, forming a subcomplete and rather dense reticulum; plage distally amyloid. Basidia 55–65 × 10–12(–15) µm, cylindric to subclavate, 4-spored; sterigmata 6–10 × 1–2 µm. Pleurocystidia absent. Pleuropseudocystidia scarce, never emergent, 4–6(–9) µm diam., irregularly cylidric, with a very dense granular content. Lamellar edge sterile, composed of marginal cells; marginal cells irregularly cylindric to subfusiform, 15–40 × 4–8 µm, hyaline, thin-walled. Hymenophoral trama mainly composed of hyphae, with some swollen compartimens present, but no distinct nests of sphaerocytes. Pileipellis 60–80 µm thick, a palisade embedded in a slime layer; terminal elements subcylindric to irregular, 20–40 × 3–6 µm, thin-walled (walls not gelatinizing), hyaline; cellular layer 30–40 µm thick, composed of rather small (10–20 µm diam.) globose cells. Pileipellis 80–120 µm thick, a trichopalisade embedded in a slime layer composed of chains of elements; terminal elements 15–40 × 3–6 µm, irregularly cylindrical to subtortous; lower elements globose up to 20 µm diam. or irregular, with pale brown intracellular pigmentation. Stipitipellis a trichoderm. Clamp‑connections absent. – Fig. 104
Ecology. Zambezian miombo woodland. Found under Brachystegia spiciformis.
Distribution. Only known from the type locality.
Revised specimens
Zimbabwe. Manicaland Prov., road from Mutare to Bvumba at peg 15,5 km, 11 Feb. 1999, A. Verbeken 99-176 (GENT, holotype).
82. Lactarius saponaceus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 209 (1996). – Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, 25 March 1994, A. Verbeken 94-353 (BR 57650–32, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 181–182 & Pl. 157–159 (1996d); Verbeken, Persoonia 17: 387–388 & Pl. 3 (2001); De Kesel & al., Guide des champignons comestibles du Bénin: 172 (2002).
Pileus 50–65 mm diam., applanate and slightly depressed in the centre; margin regular, straight, faintly striate, paler; pellis not dehiscent, dry, felty, greyish yellow to orange grey (4A3, 4B3 to 5B2), brown in the centre, pale yellow at the margin (4A3). Stipe 15–40 × 5–13 mm, eccentric to central, cylindrical, tapering downwards, pale brown to greyish brown, white at the base, dry, felty. Lamellae adnate to decurrent, quite dense (L+l = 8+7 to 8+24/cm), unequal with 1–3 lamellulae between 2 lamellae, paper‑like, elastic, thin, pale yellow (4A3); edge entire, dark grey to dark brown (very conspicous in young specimens). Context firm, thick in pileus (4–5 mm), solid in stipe, white, unchanging; taste astringent in Verbeken 94-353, strongly soap‑like and disgusting in Verbeken 94-302. Latex not abundant, although very abundant when pileus and stipe are separated, white, unchanging; taste soap‑like and disgusting in Verbeken 94-302. Spore-deposit pale brown to dark cream. – Pl. 50
Chemical reactions. Context unchanging with FeSO4, NH4OH, HCl.
Basidiospores globose to subglobose, sometimes ellipsoid, 7.0–7.7–8.1–9.3 × 6.8–7.0–7.5–8.2 mm (Q = 1.00–1.08–1.11–1.22; n = 80); ornamentation amyloid, composed of ridges up to 1 mm high, forming an incomplete reticulum with isolated warts and ridges; plage inamyloid. Basidia 50–65 × 9–11 mm, cylindrical to slightly fusiform, 4‑spored. Pleurocystidia absent. Pleuropseudocystidia very abundant, 4–6 mm diam., emergent, irregularly cylindrical, mucronate to tortuous; content oleiferic, sometimes guttulate. Lamellar edge sterile; marginal cells 15–35 × 4–6 mm, cylindrical to fusiform, with pale brown content. Hymenophoral trama irregular, composed of hyaline hyphae; lactifers abundant. Pileipellis a hymeniderm, 40–50 mm thick; terminal elements of suprapellis cylindrical to clavate, 15– 25(–35) × 5–12 mm, without brown pigment; subpellis composed of short to cylindrical cells. Stipitipellis a hymeniderm; terminal elements longer and narrower, 10–30 × 5–7 mm. Clamp‑connections absent. – Fig. 105
Ecology. Zambezian miombo woodland, Sudanian woodland with Isoberlinia,Burkea africana or gallery forests with Berlinia grandiflora.
Distribution. Known from Benin, Burundi, Malawi, Togo, Zambia and Zimbabwe.
Revised specimens
Benin. Atacora Prov., Kota, 7 Jun. 2002, A. De Kesel 3309 (BR 152200–07); Borgou Prov., Wari Maro, 21 Aug. 1997, A. De Kesel 1959 (BR 74950–66); Ibid., 20 Sept. 1998, A. De Kesel 2281 (BR 112774–60); Borgou Prov., Wari Maro, 15 June 2000, A. De Kesel 2803 (BR 126356–62); Ibid., 18 Sept. 2000, A. De Kesel 2897 (BR 129108–01); Ibid., 13 Sept. 2001, A. De Kesel 3161 (BR 149828–60); Zou Prov., Ouèssè, Forêt Classée de Gbadji, 13 Jun. 2004, A. De Kesel 3595 (BR 157208–68).
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, 25 March 1994, A. Verbeken 94-302 (BR 57643–25), 94-353 (BR 57650–32, holotype; GENT, isotype).
Malawi. Southern Prov., Machinga distr., near Machinga, Malosa Forest Reserve, 25 Jan. 2005, F. Noe 05/05 (GENT); Northern Prov., Mzimba distr., Perekezi Forest Reserve, 28 Jan. 2005, A. Verbeken 05-084 (GENT); Ibid., Ntanga Forest Reserve, 30 Jan. 2005, A. Verbeken 05-158 & 05-159 (all GENT); Ibid., 31 Jan. 2005, A. Verbeken 05-176 (GENT); Mtanga tanga forest, near Chicangana village, 19 Feb. 2005, F. Noe 05/395 (GENT).
Togo. Central Prov., Forêt Classée d’Alédjo, 11 Jul 2007, A. De Kesel 4297 (BR 158418–17)
Zimbabwe. Manicaland Prov., along Mutare-Bvumba road, near Prince of Wales Viewpoint, 11 Feb. 1999, A. Verbeken 99-171 (GENT); Masvingo Prov., south of Lake Mutirikwe, around Norma Jean’s lodge, 16 Feb. 1999, A. Verbeken 99-224 (GENT); Mashonaland East Prov., Bromley, Liemba farm, 3 Feb. 1999, A. Verbeken 99-106 (GENT).
83. Lactarius pusillisporus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 206 (1996). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, A. Verbeken 94-285 (BR 57638–20, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 182–183 & Pl. 160–161 (1996d); Verbeken, Persoonia 17: 383–385 (2001).
Pileus 30–75 mm diam., planoconvex when young, applanate to planoconcave, sometimes subumbilicate or subumbonate, mostly regular; margin regular, incurved then straight, faintly striate, paler; pellis not dehiscent, dry, velvet‑like, felty in the centre, smoother and wrinkling towards the margin, brownish orange, reddish blond to light brown and yellowish brown (5C3–4 to 5D5 to 5E5); centre dark brown to brownish buff (6F5 to 6F3); margin pale yellow (4A3). Lamellae adnexed, narrowly adnate to adnate, unequal with lamellulae of different lengths, moderately spaced (L+l = 3+6 to 8+11/cm), paper‑like, thin, white to cream, pale brown when older; edge entire, dark grey to dark brown. Stipe 35–60 × 5–10 mm, slightly eccentric to central, cylindrical, long and slender, slightly tapering downwards, concolorous with pileus, paler towards the apex and the base, dry, felty. Context firm, elastic, white to cream, sometimes changing pale greyish; taste not remarkable (very sweet in Verbeken 94-427); smell sweet, agreeable. Latex quite abundant to very abundant, white, unchanging; taste mild to a little astringent. Spore‑deposit pale brown. – Pl. 51
Chemical reactions. Context unchanging with FeSO4 (but pale greyish green in Verbeken 94-427), unchanging with NH4OH, KOH and HCl.
Basidiospores subglobose to ellipsoid, 5.3–5.8–6.5–7.1 × 4.5–5.1–5.9–6.5 mm (Q = 1.01–1.10–1.16–1.25; n = 100); ornamentation amyloid, composed of ridges up to 0.75 mm high, forming an almost complete reticulum with some isolated warts; plage inamyloid. Basidia 35–45 × 7–11 mm, cylindrical to slightly clavate, 4‑spored. Pleurocystidia absent. Pleuropseudocystidia not abundant, 3–6 mm diam., emergent, cylindrical, with rounded apex; content oleiferic. Lamellar edge sterile; marginal cells 15–35 × 3–5 mm, irregularly cylindrical to fusiform, sometimes clavate, content pale brown. Hymenophoral trama irregular, composed of hyaline hyphae; lactifers not abundant, sphaerocytes absent. Pileipellis a trichoderm to trichopalisade, 70–100 mm thick; terminal elements cylindrical, long, slender, 20–60 × 3–5 mm, arising from broadly cylindrical to almost spherical basal elements. Stipitipellis a trichoderm; terminal elements 25–45 × 3–5 mm, septate. Clamp‑connections absent. – Fig. 106
Ecology. Miombo woodland.
Distribution. So far, only known from Burundi.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, March 1994, A. Verbeken 94-285 (BR 57638–20, holotype; GENT, isotype); Ibid, A. Verbeken 94-011 (BR 57562–41), 94-022 (BR 57570–49), 94-044 (BR 57573–52) & 94-427 (BR 57659–41).
84. Lactarius kabansus Pegler & Piearce
Pegler & Piearce, Kew Bull. 35: 487 (1980). – Type: Zambia, Kitwe, purchased from Chimwemwe market, 21 Dec. 1982, G. Piearce FP600/3 (K(M) 159481, holotype).
Misappl.: Lactarius hispidulus ss. Degreef (1990), L. craterelloides ss. Degreef (1990).
Selected descriptions and icons. Buyck, Champignons comestibles de l'Ouest du Burundi: 106, fig. 80 (1994); Härkonen & al., Karstenia 35: Suppl.: fig. 1 (1995); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 190–192 & Pl. 182–187 (1996d); Karhula & al., Karstenia 38: fig. 22 (1998); Ryvarden & al., An introduction to the larger fungi of South Central Africa: 100 (1994); Verbeken, Persoonia 17: Pl. 7 (2001); Härkonen & al., Norrlinia 10: 87 (2003).
Pileus 20–70 mm diam., firm, elastic, thin, convex, slightly depressed to infundibuliform; margin strongly incurved, regular and involute when young, striate and irregularly lobed when older, always deflexed downwards; pellis almost totally dehiscent, smooth, dry, finely felty when young, glabrous and even somewhat shiny when older, grey to brownish grey (5D4 to 5E4) to brown and dark brown (6E4–5 to 6F4–5). Lamellae decurrent to strongly decurrent, unequal with lamellulae of 3 different lengths (regular pattern), crowded (L+l = 6+16 – 8+20/cm), thin, narrow, 1.5–3 mm broad, thick, paper-like, pale yellow to light orange (4A3–4 to 5A5); edge entire, concolorous, darker brown towards the margin of the pileus. Stipe 20–40 × 4–15 mm, cylindrical, on the whole slightly expanded downwards, tough, concolorous with pileus, dry, smooth and glabrous, sometimes with whitish tomentum at the base. Context thin, firm, chambered to fistulous in the stipe, white to cream, light orange then greyish red when bruised; taste mild and pleasant; smell not particular or fungoid. Latex only abundant when young and fresh, watery, white, unchanging; taste mild. Spore-deposit white. – Pl. 51
Chemical reactions. Context unchanging with FeSO4, NH4OH, HCl and KOH.
Basidiospores ellipsoid to elongate, 7.5–8.5–9.0–10.0 × 5.2–6.2–6.7–7.2 mm (Q = 1.21–1.32–1.39–1.53; n = 100); ornamentation amyloid, composed of ridges up to 1 mm high and a few isolated warts, forming a dense reticulum; plage distally amyloid or rather rugose. Basidia 35–45 × 9–11(–12) mm, narrowly clavate to narrowly utriform, 4-spored. Pleurocystidia absent. Pleuropseudocystidia rather abundant, emergent and distinct, 7–9(–10) mm diam. in the upper part, sometimes tapering and then 2–3 mm in the lower part, cylindrical to flexuous; content oleiferic with guttules. Lamellar edge sterile; marginal cells 15–35 × 6–10(–13) mm, short, narrowly clavate and inflated, thin-walled, often septate, even branched, with an intracellular brown pigmentation. Hymenophoral trama irregular, composed of hyaline hyphae, 2–7 mm diam., intermixed with numerous laticiferous hyphae, (2–)9–12 mm diam.; sphaerocytes lacking. Pileipellis a well-developed, 20–40 mm thick hymeniderm; terminal elements 10–35 × 8–15 mm, clavate to obovoid and broadly clavate, with brown intracellular pigmentation. Stipitipellis as pileipellis; terminal elements smaller, 8–20 × 5–10 mm. Clamp-connections absent. – Fig. 107
Distribution. Known from Burundi, Democratic Republic of Congo, Kenya, Malawi, Tanzania, Zambia and Zimbabwe.
Ecology. Growing in open Zambezian miombo woodland, growing under various trees (Brachystegia, Isoberlinia, Uapaca, etc.). Common, especially on sandy soils. Especially abundantly fruiting at the beginning of the rain season.
Revised specimens
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, March 1994, A. Verbeken 94-218 (BR 57626–08), 94-301 (BR 57651–33), 94-344 (BR 57645–27) & 94-425 (BR 57658–40); Nkayamba, just N of Rumonge, March 1993, B. Buyck 5001 (PC); Ibid., Nov. 1993, B. Buyck 5227 (PC); Ibid., Dec. 1993, B. Buyck 5282 & 5342 (all PC); Mugara, 18 Nov. 1978, J. Rammeloo 5787 (BR 8769–39); Cankuzo Prov., Murango, 3 March 1994, B. Nzigidahera 62 (PC).
Democratic Republic of Congo. Shaba region., Luiswishi, 15 Feb.1990, J. Degreef 90-020 (BR 5046–02); Biano, Lubudi, 18 March 1990, J. Degreef 90-190 (BR 5114–70); Kipopo, 2 Dec. 1971, D. Thoen 5075 (BR 66271–20); Ibid., 10 Dec. 1959, M.C. Schmitz-Levecq 174 (BR 12299–77); Ibid., 24 Dec. 1960, M.C. Schmitz-Levecq 262 (BR 12297–75); Shilatembo, 24 Jan. 1972, D. Thoen 5545 (BR 66338–87).
Kenya. Coast Prov., Kwale distr., Shimba hills, Giriama Point, Nature Reserve, June 1994, D. Ivory IV 867 (BR 49368–92).
Malawi. Central Prov., Dedza distr., Chongoni, 11 Feb. 1972, J. Williamson 603 (K(M) 38477).
Tanzania. Western Prov., Tabora distr., purchased from Urambo market, 13 Dec. 1991, T. Saarimäki & al. 1089 (H); Southern Highlands, Iringa distr., Mafinga, Sao Hill Misitu, 12 Feb. 1993, T. Saarimäki & al. 1590 (H).
Zambia. Copperbelt Prov., Chati‑forest, near Kitwe, Dec. 1990, B. Buyck 3133, 3134, 3169, 3180 & 3205 (all PC); Kitwe, purchased from Kwacha market, Dec. 1976, G. Piearce FP462 (K(M) 38491; E, paratype); Ibid., purchased from Chimwemwe market, Dec. 1976, G. Piearce FP600/3 (K(M) 159481, holotype); Luapala Prov., Samfya, 29 Dec. 1990, B. Buyck 3277 (PC); Mansa, Kabunda mission, 3 Jan. 1993, B. Buyck 3362 (PC); Western Prov., Mwinilunga, 29 Jan. 1991, B. Buyck 3512 (PC); Mongu, May 1979, G. Piearce FP622/2 (K(M) 38496); Southern Prov., Mazabuka, Nat. Irr. Res., 14 Jan. 1974, Rappaport s.n. (K(M) 38500); Choma distr., May 1979, G. Piearce FP615/2 (K(M) 38495); farm Gibsons, 16 Jan. 1996, B. Buyck & G. Eyssartier 96-059 (PC); Lusaka, near airport, 9 Feb. 1996, B. Buyck & G. Eyssartier 96-332.
Zimbabwe. Midlands Prov., Mvuma, Central Estates, 29 Jan. 1995, C. Sharp 353/95 (BR 38491–79); Ibid., Gadzanga section, 23 Jan. 1994, C. Sharp 290/94 (BR 38490–78); Ibid., Baru section, 27 Jan. 1999, A. Verbeken 99-033 (GENT); Ibid., paddock 77, 5 Feb. 1999, A. Verbeken 99-126 (GENT); Midlands Prov., Mvuma, Mtao forest, 12 Feb. 1999, Gansemans in A. De Kesel 2743 (BR 115758–37); Manicaland Prov., along Mutare-Bvumba road, near Prince of Wales Viewpoint, 11 Feb. 1999, A. Verbeken 99-179 (GENT); Chipinge, Busi farm, forest around Scott’s house, 12 Feb. 1999, A. Verbeken 99-205 (GENT); Mguzu, 7 Feb. 1999, A. De Kesel 2430 (BR 112600–80).
Notes. The macroscopic description is based on Pegler & Piearce (1980) and fieldnotes of Buyck and Verbeken. The microscopic description is based on the type-material and B. Buyck 3169 & 3362.
Livingstone observed in 1867 that in Zambia's Northern Prov. people lived on mushrooms and leaves during famine periodes. One of the consumed species he mentioned, called "Nakabansa", most probably refers to Lactarius kabansus (Piearce 1985). Lactarius kabansus is still a very popular edible mushroom throughout the Zambezian miombo woodland area (e.g. Buyck 1994, Härkonen & al. 1995, Pegler & Piearce 1980). It can be eaten also in the raw state, without being parboiled.
85. Lactarius sciaphilus Verbeken & C. Sharp
Verbeken & C. Sharp, Cryptog. Mycol. 24: 128 (2003). – Type: Zimbabwe, Mvuma, Beacon Hill Section Central Estates, 30 Dec. 1996, C. Sharp 482/96 (GENT, holotype).
Selected descriptions and icons. Verbeken & C. Sharp, Cryptog. Mycol. 24: 126–129 (2003).
Pileus 70–120 mm diameter, centrally depressed to infundibuliform; margin entire, slightly in-rolled, sometimes sulcate and wavy; pellis smooth, mat, hazel, umber or with shades of sienna at centre. Stipe 37–42 × 12–25 mm, shortly cylindrical to tapering at base, smooth, mat, firm, hazel to umber with sepia base. Lamellae adnate to subdecurrent, prolonging in a definite line at the stipe-apex; lamellulae present and especially numerous at margin; sparse to moderately dense (L+l = 5–8/cm mid-radius), thick, waxy or papery, very brittle, 8(19)mm broad, pale orange when young, saffron, orange, fulvous, becoming sienna in older specimens. Context white in pileus; solid, pithy, white in stipe, to orange at base; smell not distinctive. Latex white, abundant. Spore-deposit white to pale salmon.
Chemical reactions. Context in pileus becoming red with guaiacol, no reaction with FeSO4 and KOH.
Basidiospores broadly ellipsoid to ellipsoid, 7.9–8.7–9.3–10.3(–10.6) × 6.7–7.5–7.6–8.4 µm (Q=1.06–1.15–1.23–1.33; n=60); ornamentation amyloid, composed of ridges up to 1µm high and a few isolated warts forming a dense reticulum; plage distally amyloid. Basidia 45–55(–65) × 10–13 µm, narrowly clavate, 4-spored; sterigmata long (up to 8.5 µm), rather slender and curved. Pleurocystidia absent. Pleuropseudocystidia rather scarce, cylindric, 4–8 µm broad. Lamellar edge sterile; marginal cells 20–30 × 5–8 µm, subcylindric to fusoid, with an intracellular brown pigmentation. Hymenophoral trama irregular, mixed; some sphaerocytes present; lactifers present. Pileipellis a well developed trichoderm up to 70 µm thick; some of the underlying cells slightly swollen (up to 20 µm diam.); terminal cells subcylindric, subfusiform, never rounded nor clavate, 30–65 × 8–10(–12) µm, with brown intracellular pigmentation. – Fig. 108
Ecology. Drier Zambezian miombo woodland.
Distribution. Only known from Zimbabwe.
Revised specimens
Zimbabwe. Midlands Prov., Mvuma, Beacon Hill Section, Central Estates, south of Beacon Hill Range, 20 Jan. 1995, C. Sharp 331/95 (GENT); Ibid., Beacon Hill Homestead, 30 Dec. 1996, C. Sharp 482/96 (GENT, holotype); Ibid., south of Beacon Hill Range, 1 Jan. 1997, C. Sharp 505/97 (GENT); Ibid., below gate of Beacon Hill Homestead, 3 Jan. 1997, C. Sharp 521/97 (GENT); Ibid., Beacon Hill Homestead, leg. G. Sharp 18 Jan. 1997, C. Sharp 571/97 (GENT).
Notes. Macroscopically Lactarius sciaphilus differs from L. kabansus in overall size and habitat preference. Lactarius sciaphilus reaches 85 mm in height (50 mm in L. kabansus) with a pileus diameter up to 120 mm (up to 70 mm in L. kabansus). Both species occur in miombo woodland but L. sciaphilus prefers deep shade and thicker leaf litter while L. kabansus is more common in the sparse litter of open patches, often with cryptogamic crusts in degraded woodland (C. Sharp, pers. observation). Microscopically, Lactarius sciaphilus is characterised by the broader spores and larger basidia. The spores in L. kabansus are 7.5–8.5–9.0–10.0 × 5.2–6.2–6.7–7.2 µm and subsequently the Q-value is larger (Q = 1.21–1.32–1.39–1.53, n=100); basidia in L. kabansus measure 35–45 × 9–11(12) µm. Furthermore, the marginal cells and terminal elements in the pileipellis are fusiform rather than clavate and rounded. Sphaerocytes are present in the hymenophoral trama and lactifers are less abundant than in L. kabansus.
Lactarius tenellus Verbeken is another close relative of L. kabansus and differs by the slender habit, usually smaller size, a more greyish-brown cap with crenulate margin, white lamellae and lack of orange tinges in the stipe base (Verbeken 2001). All these species are edible and mostly not distinguished by local populations (Verbeken & Sharp 2003).
86. Lactarius tenellus Verbeken & Walleyn
Verbeken, Persoonia 17: 392 (2001). − Lactarius kabansus var. pallidus Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 202 (1996), non Lactarius pallidus Pers.: Fr. (1797). − Type: Burundi, Rumonge Forest Reserve, Nyamirambo hill, A. Verbeken 94-157 (BR 57611–90, holotype; GENT, isotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 192–193 & Pl. 188 (1996d); Verbeken, Persoonia 17: 392–393 & Pl. 8 (2001), De Kesel & al., Guide des champignons comestibles du Bénin: 164 (2002).
Pileus 13–35 mm diam., planoconvex to applanate and slightly depressed to infundibuliform, sometimes with distinct acute papilla; margin incurved, regular and involute when young, then regularly grooved, crenulate; pellis almost totally dehiscent, smooth, dry, finely felty when young, very minutely cracking, grey to brownish grey or buff (5C3–4 to 5D3–4 to 6D3 to 6E3). Lamellae decurrent to strongly decurrent with tooth, unequal with lamellulae of 3 different lengths (regular pattern), very dense, thin, narrow, 1.5–3 mm broad, thick, rounded to slightly slightly running off, paper-like, whitish to pale yellow (4A3–4 to 5A2); edge entire, concolorous. Stipe 15–30 × 4–10 mm, cylindrical to irregular, concolorous with pileus or darker (5D3 to 5E3), white at the extreme basis, dry, smooth and glabrous or slightly velvety and soft, sometimes with whitish tomentum at the base. Context thin, firm, solid or fistulous in the stipe, white to cream, unchanging; taste mild and pleasant; smell not particular or a bit chemical, like paint. Latex only abundant when young and fresh, watery, white, unchanging; taste mild. Spore-deposit white. – Pl. 52
Chemical reactions. Context unchanging with FeSO4, NH4OH, HCl and KOH. Latex unchanging with KOH.
Basidiospores ellipsoid to elongate, 7.5–8.5–9.0–10.0 × 5.2–6.2–6.7–7.2 mm (Q = 1.21–1.32–1.39–1.53; n = 100); ornamentation amyloid, composed of ridges up to 1 mm high and a few isolated warts, forming a dense reticulum; plage distally amyloid or rather rugose. Basidia 35–45 × 9–11(–12) mm, narrowly clavate to narrowly utriform, 4-spored. Pleurocystidia absent. Pleuropseudocystidia rather abundant, emergent and distinct, 7–9(–10) mm diam. in the upper part, sometimes tapering and then 2–3 mm in the lower part, cylindrical to flexuous; content oleiferic with guttules. Lamellar edge sterile; marginal cells 15–35 × 6–10(–13) mm, short, narrowly clavate and inflated, thin-walled, often septate, even branched, with an intracellular brown pigmentation. Hymenophoral trama irregular, composed of hyaline hyphae, 2–7 mm diam., intermixed with numerous laticiferous hyphae, (2–)9–12 mm diam.; sphaerocytes lacking. Pileipellis a well-developed, 20–40 mm thick hymeniderm, terminal elements 10–35 × 8–15 mm, clavate to obovoid and broadly clavate, with brown intracellular pigmentation. Stipitipellis as pileipellis; terminal elements smaller, 8–20 × 5–10 mm. Clamp-connections absent. – Fig. 109
Ecology. Zambezian miombo woodland, Sudanian savannah woodland. Found terrestrial under ectomycorrhizal trees such as Brachystegia, Julbernardia, Isoberlinia, Uapaca.
Distribution. Widespread in tropical Africa. Known from Benin, Burundi, Democratic Republic of Congo, Kenya, Malawi, Tanzania, Togo, Zambia and Zimbabwe.
Revised specimens
Benin. Borgou Prov., Adjimon, 11 Sept. 2001, A. De Kesel 3145 (BR 149790–22); Wari Maro, 21 Aug. 1997, A. De Kesel 1951 (BR 129213–09); Ibid., 19 June 1998, A. De Kesel 2154 (BR 129213–09); Ibid., 28 June 1998, A. De Kesel 2215 (BR 112724–10); Ibid., 9 Sept. 2001, A. De Kesel 3109 (BR 149852–84) ; Ibid., 16 June 2002, A. De Kesel 3415 (BR 152136–40) Atacora Prov., Kota, 29 Sept. 2000, A. De Kesel 2967 (BR 129213–09); Ibid., Koussokouango, 21 June 2004, A. De Kesel 3716 (BR 157190–50); Zou Prov., Savalou, 13 June 2004, A. De Kesel 3598 (BR 157243–06).
Burundi. Bururi Prov., Nyamirambo, Rumonge Forest Reserve, March 1994, A. Verbeken 94-470 (BR 57676–58); Nkayamba, just N of Rumonge, March 1994, A. Verbeken 94-157 (BR 57611–90, holotype; GENT, isotype); Ibid., Dec. 1991, B. Buyck 4024 (PC); Cankuzo Prov., Murango, 11 March 1994, B. Nzigidahera 64 (PC); Ibid., Dec. 1994, B. Buyck 5671 (PC).
Democratic Republic of Congo. Shaba, Kipopo, 6 March 1959, M.C. Schmitz-Levecq 72 (BR 12301–79); Ibid., 9 Jan. 1961, M.C. Schmitz-Levecq 286 (BR 12295–73).
Kenya. Coastal Prov., Shimba Hills (Kayatelezia plots), 4 Dec. 2004, A. De Kesel 3975 (BR 162769–03)
Malawi. Southern Prov., Zomba distr., near R.C. Cathedral, Jan. 1974, J. Williamson 762 (K(M) 38482) & 769 (K(M) 38485); Ibid., Blantyre, 13 Jan. 1972, J. Williamson 563 (K(M) 38474); Machinga distr., Liwonde Forest Reserve, 22 Jan. 2005, A. Verbeken 05-015 (GENT); Northern Prov., Mzimba distr., Ntanga Forest Reserve, 30 Jan. 2005, A. Verbeken 05-171 (GENT).
Tanzania. Morogoro Prov., Morogoro distr., Dec. 1992, E. Munyanziza 2-5 & 2-6 (all WAG); Lake Prov., Mwanza distr., Geita Forest Reserve, 5 Jan. 1971, D.L. Ebbels F5 (K(M) 38487).
Togo. Centrale Prov., Parc National Fazao, 20 July 2007, A. De Kesel 4463 (BR 163798–62)
Zambia. Copperbelt Prov., Chati‑forest, near Kitwe, Dec. 1990, B. Buyck 3176 & 3222 (all PC); Luapala Prov., Mansa, Kwanfumwe, 5 Jan. 1991, B. Buyck 3410 (PC); Northern Prov., North Luangwa National Park, 25 Jan. 1995, D. Shah‑Smith 164 (K(M) 35506); road from Mpongwe to Lusaka, Mikati, 18 Jan. 1996, B. Buyck & G. Eyssartier 96-094 & 96-095 (all PC); Lusaka, near airport, 8 Feb. 1996, B. Buyck & G. Eyssartier 96-330 & 96-331 (all PC); Ibid., 9 Feb. 1996, B. Buyck & G. Eyssartier 96-333 (PC).
Zimbabwe. Manicaland Prov., Chipinge, Busi farm, around Scott’s house, 12 Feb. 1999, A. Verbeken 99-204 (GENT); Penhalonga, hills behind Harris’ garden, 9 Feb. 1999, A. Verbeken 99-147 (GENT); Midlands Prov., Mvuma, Central Estates, Beacon Hill section, ridge at the ostrich incubator, 26 Feb. 1999, A. Verbeken 99-008 & 99-009 (all GENT); hills north of the ostrich incubator, 15 Feb. 1999, A. Verbeken 99-213 (GENT); Baru section, 27 Jan. 1999, A. Verbeken 99-031 (GENT); Masvingo Prov., south of Lake Mutirikwe, around Norma Jean’s lodge, 16 Feb. 1999, A. Verbeken 99-225 (GENT).
Notes. The species differs from Lactarius kabansus Pegler & Piearce as well microscopically (terminal elements of the pileipellis obovoid to broadly clavate, 10–20 × 8–15 µm in L. kabansus; terminal elements of the pileipellis cylindrical, long and slender, 10–40 × 6–10 µm in L. tenellus), as macroscopically: Lactarius tenellus has a rather dark brown cap when very young, which together with its contrasting white lamellae, makes it reminiscent of L. lignyotus. The cap colour typically fades to grey, almost without brownish components, while L. kabansus has brown to dark brown colours, eventually also very pale when specimens are exposed to the sun. L. kabansus is a usually more robust species, with an irregular, deflexed margin, while L. tenellus is a slender species with a crenulate margin. The initially pure white lamellae of L. tenellus, become more yellowish, but not as soon and as ochraceous orange as in L. kabansus. In case of doubt, the cutting of the stipe base seems an inambiguous test: the context in the base of the stipe is white and remains white in L. tenellus, while in all encountered specimens of L. kabansus, even in old and insect-infected ones, the stipe base shows an orange colour when cut. As to the spores, no conspicuous differences have been observed.
A popular edible species (e.g. De Kesel & al. 2002, Yorou & De Kesel 2002), often mixed with L. kabansus.
Heim, Bull. Jard. Bot. Etat Bxl 25: 73 (1955). − Type: Guinea, near Macenta, April 1939, R. Heim D74 (PC, holotype).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 73–75 (1955); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 187–188 & Pl. 170–177 (1996d).
Pileus 10–35 mm diam., thin, deeply depressed to infundibuliform, sometimes with papilla in the centre, irregular; margin crenate, incurved when young, long and regularly striate when older; pellis not dehiscent, finely tomentose, velvety cracked, smooth and greasy, wrinkling in the centre, brown (6D6), dark brown, snuff brown, prolonging on the edge of the lamellae, sometimes with locally darker spots (6E5). Lamellae deeply decurrent, prolonging in ridges on the stipe, unequal with lamellulae of two different lengths, moderately distant, narrow (< 3 mm), rather thick, elastic, honey-coloured, yellowish orange, orange-brown (5A6 to 5B6); edge entire, concolorous, but with same colour as the pileus near the edge of the pileus. Stipe 30–35 × 5–7 mm, irregularly cylindrical, subfusiform, tapering downwards and curved at the base, solid, very tough, brownish, reddish brown, dark brown in the upper half, paler at the apex, mixed with orange in the under half, sometimes with greyish yellow felty layer at the base. Context white, then cream to ochraceous in pileus; white, then ochraceous, finally orange in the stipe; brownish orange under the pellis; taste mild; smell fish-like. Latex abundant, fluid, white, changing sometimes red after more than half an hour and then also staining context reddish (in Schreurs 1473); taste mild, a bit fetid. Spore-deposit not observed. – Pl. 53
Chemical reactions. Context unchanging with FeSO4.
Basidiospores subglobose to ellipsoid, 6.1–6.9–7.9–8.3(–9.0) × 5.5–6.0–6.6–7.0(–7.5) mm (Q = 1.04–1.13–1.20–1.28, n = 70); ornamentation amyloid, composed of irregular, spiny warts or ridges, up to 1.5 mm high, isolated or connected by low and fine connective lines; plage distally amyloid; wall locally rugose. Basidia 35–45(–50) × 8–12(–14) mm, cylindrical to clavate, 4-spored, sometimes 2-spored; content sometimes granular. Pleurocystidia scarce, 30–40 × 8–10 mm, cylindrical to subfusiform, with acute apex, hyaline, thin-walled; content slightly granular. Pleuropseudocystidia very abundant, emergent, 6–10 mm diam., cylindrical to subclavate, with rounded apex; content oleiferic, guttate. Lamellar edge sterile; marginal cells 15–40 × 5–8 mm, short and rounded or longer and fusiform, tapering upwards, sometimes septate, hyaline, thin-walled; content granular. Hymenophoral trama regular, composed of hyaline hyphae, 2–7 mm diam.; lactifers abundant. Pileipellis hymeniderm-like; terminal elements 15–35 × 8–15 mm, cylindrical, clavate to obovoid and broadly clavate, with brown intracellular pigmentation; subpellis interwoven. Stipitipellis as pileipellis; terminal elements cylindrical, slender, hair-shaped, 25–40 × 4–6(–12) mm; subpellis regular, composed of parallel hyphae. Clamp-connections absent. – Fig. 110
Ecology. Found terrestrial or on rotten wood in drier Guineo-Congolian rainforest with Gilbertiodendron dewevrei, lowland rainforest, dry evergreen forest (Muhulu).
Distribution. Known from Cameroon, Democratic Republic of Congo, Gabon and Guinea.
Revised specimens
Cameroon. South Western Prov., near Mundema, Korup National Park, transect P18, March 1991, R. Watling 24191 (E); Ibid., transect P17-18, March 1991, R. Watling 25222 (E); Ibid., March 1991, R. Watling 25406 (E).
Democratic Republic of Congo. Equateur Prov., Binga, Dec. 1928, M. Goossens-Fontana 882 (BR 12311–89); Shaba region, Luiswishi, 13 Feb. 1986, J. Schreurs 1091 (BR 7928–71); Ibid., 22 March 1986, J. Schreurs 1473 (BR 8244–96).
Gabon. Ogooué-Ivindo Prov., Zoula, near Mekambo, 19 March 2005, J. Degreef 291 (BR 166507–55) & 298 (BR 166506–54); Mekambo, 4 Oct. 2006, J. Degreef 458 (BR 164535–23).
Guinea. Upper Guinea, road to Kérouané, around Macenta, April 1939, R. Heim D74 (PC, holotype).
Notes. The type consists of one basidiome in very bad condition. It has been preserved in alcohol, which has evaporated. The spores and basidia of the type-specimen are slightly larger than those of the three specimens of Cameroon. The other characters are similar.
The macroscopic description is based on the notes from Heim (1955a), completed with fieldnotes from Watling en Schreurs. Watling 25222 & 25406 showed no latex. The taste of latex and context is very slightly acrid in Schreurs 1091. The microscopic description is especially based on Heim D74 and Watling 24191. The spores in Schreurs 1473 are somewhat deviating from the other specimens.
The species is easily recognized by the small, dark and papillate pileus combined with the orange lamellae and the reddish tinge in the stipe. It differs from the closely related Lactarius adhaerens by the presence of true cystidia.
In Gabon this species is consumed by the pygmee population and sometimes sold on market (Degreef, pers. comm.).
88. Lactarius pseudolignyotus R. Heim
Heim, Bull. Jard. Bot. Etat Bxl 25: 56 (1955). − Type: Ivory Coast, Tiapleu Forest, March 1939, R. Heim C19 (PC, holotype).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 56–58 & Pl. 4, fig. 6 (1955); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 190 & Pl. 181 (1996d).
Pileus 25–50 mm diam., strongly depressed, without papilla, irregular; margin deeply deflexed downwards, grooved, crenulate; pellis cream to chamois-colour, with fine, dark, ochraceous brown squamules, darker near the margin. Lamellae adnate to decurrent, prolonging in longitudinal grooves on the stipe, distant (L+l = 24/total), moderately large (4 mm), rather thick, orange; edge broadly sinuose, concolorous. Stipe 20–25 × 5–11 mm, cylindrical, broader towards the apex, tapering downwards, squamulose, concolorous to the pileus, somewhat paler, orange at the base, fistulous. Context white in the pileus, orange in the stipe, creamish orange in the lamellae; taste mild; smell fish-like. Latex abundant, white, opalescent; taste mild. Spore-deposit cream. (According to Heim 1955a).
Chemical reactions. Context unchanging with gaiac and gaiacol. (According to Heim 1955a).
Basidiospores ellipsoid, 6.9–8.1–9.3 × 5.6–6.4–7.1 mm (Q = 1.14–1.28–1.36, n = 30); ornamentation amyloid, composed of ridges, up to 2 mm high, forming an almost complete reticulum; some very low and short, isolated ridges present; plage inamyloid. – Fig. 111
Ecology. Guineo‑Congolian rainforest.
Distribution. Known only from Ivory Coast.
Revised specimens
Ivory Coast. Near Danané, Tiapleu forest, March 1939, R. Heim C19 (PC, holotype).
Notes. The type-specimen consists of one basidiome in very bad condition. It was first preserved in alcohol, but the alcohol has evaporated and the material is as hard as stone. Only the spores could be observed. A surface view of the pileipellis is given. Heim mentions that the pileipellis is filamentous with cylindrical elements of 3.6–5 mm diam., terminal elements are incrusted, claviform or piriform, 12–28 × 7.5–12.5 mm.
We do not exclude that this species is conspecific with the previous one. More collections and good fieldnotes are needed to find out. One of the questions is how stable the macroscopical features of the pileipellis are (colour, presence of scales).
Sect. Pseudofuliginosi Verbeken
Lactarius sect. Pseudofuliginosi Verbeken, Persoonia 17: 392 (2001).
− Type species: Lactarius atro-olivinus Verbeken & Walleyn.
Key to tropical African species
1. Pleuromacrocystidia abundant; pileus with blackish olivaceous tinges; spores ornamented with narrow, fine ridges ............................................................................................ 89. L. atro-olivinus
Pleuromacrocystidia absent; pileus rather brownish; spores more heavily ornamented ............ 2
2. Ascending elements in the pileipellis 5–9 mm broad, subclavate or subfusiform; pileus mat and very finely felty ....................................................................... 90.1. L. undulatus var. undulatus
Ascending elements in the pileipellis narrow (2–4 µm), cylindric, more poorly represented; pileus smooth and shiny, slightly viscid ................................................ 90.2. L. undulatus var. rasilis
Species descriptionS
Notes. The description is based on the type specimen. This taxon differs from the type variety in two depending characters (microscopical structure of the pileipellis and macroscopic aspect of the cap), and is therefore considered a variety and not a separate species. More material is needed to understand the variation of characters in these similar taxa.
89. Lactarius atro-olivinus Verbeken & Walleyn
Verbeken, Persoonia 17: 394 (2001). – Type: Zimbabwe, Manicaland Prov., hill north of Penhalonga, 9 Feb. 1999, A. Verbeken 99-143 (GENT, holotype).
Selected descriptions and icons. Verbeken, Persoonia 17: 394–396 & Pl. 6 (2001).
Pileus 30–100 mm diam., regularly infundibuliform, with a typical Clitocybe-shape, with a distinct, sometimes suddenly deep and narrow umbilicus; margin incurved and staying slightly incurved, broadly wavy, crenulate in older specimens; surface smooth, shiny, with very small, almost indistinct radially wrinkles, very slightly sticky, very dark brown, almost blackish at margin, greyish brown (7F3–4) but more intense towards the centre (6F4 to 6E3); overall tinges dark brown to blackish brown with an olivaceous shade. Stipe 30–60 × 10–16 mm, subcylindric, slightly tapering at the base, smooth, dryer than the pileus, with a somewhat pale, almost pruinose aspect which makes it look like plaster, greyish brown (6E3), not unicolorous, locally grey (6C1) in young ones, with plastery greyish tinge, with a very dark narrow zone just beneath the lamellae, and right underneath this darker zone a bit paler than in the rest of the stipe. Lamellae broadly adnate, moderately distant to distant (L+l = 4 to 7/cm halfway radius), papery thin, not very brittle, with very abundant lamellulae of many different lengths (no regular pattern), dark blonde (5D4), staining dark in old specimens. Context rather thick and firm, brownish-grey (same tinge as pileus, but paler); smell a bit fatty, faintly as L. quietus; taste first mild, but soon very acrid, burning. Latex moderately abundant, white, slightly cream, drying dark olivaceous brown (5E4) after more than 30 minutes, unchanging with KOH, a bit cream on paper. Spore-deposit not observed. – Pl. 53
Chemical reactions. Context immediately blue with gaiac. Latex unchanging with KOH.
Basidiospores globose to subglobose, 8.3–9.2–9.3–10.0 × 7.7–8.3–8.5–9.2 µm (Q = 1.00–1.08–1.10–1.17, n = 40); ornamentation amyloid, composed of narrow and acute ridges, up to 2.5 µm high, forming a very incomplete reticulum; plage distally amyloid. Basidia 50–65 × 12–15 µm, 4-spored, subclavate, with guttulate or densily granular content; sterigmata 9–11 × 1–3 µm. Pleuromacrocystidia abundant, sometimes slightly emergent, 50–70 × 6–8 µm, narrowly fusiform, with tapering or mucronate apex; content needle-like. Pleuropseudocystidia rather abundant, never emergent, 3–4 µm diam., narrow, cylindric, with rounded apex. Lamellar edge sterile; composed of marginal cells which are narrowly cylindric, 30–45 × 4–6 µm, hyaline, thin-walled. Hymenophoral trama mixed; filamentose with some sphaerocytes present; lactifers abundant. Pileipellis an ixotrichoderm, 100–120 µm thick, composed of very loosely interwoven repent and ascending hyphae; hyphae 2–4 µm diam., thin-walled, some with a brown intracellular pigmentation; some incrustations also present; embedded in a slime layer but never with gelatinizing walls. Stipitipellis a trichoderm, locally a cutis, not embedded in a slime-layer, but composed of hyphae of 4–6 µm diam. and with a rather rigid wall; no pigmentation present. Clamp-connections absent. – Fig. 112
Ecology. Zambezian miombo woodland, in Zimbabwe growing under Brachystegia spiciformis.
Distribution. Known from Malawi, Zambia and Zimbabwe.
Revised specimens
Benin. Donga Prov., Forêt Classée de Bassila, 17 Jun. 2004, A. De Kesel 3661 (BR 157046–03).
Malawi. Northern Prov., Mzimba distr., Mtanga tanga forest, near Chicangana village, 19 Feb. 2005, F. Noe 05/399 (GENT); Ibid., Perekezi Forest Reserve, 14 Feb. 2005, F. Noe 05/255 (GENT).
Zambia. Kitwe, 7 Dec. 1978, G. Piearce FP593/2 (K(M) 38492).
Zimbabwe. Manicaland Prov., 15.5 km peg along Mutare-Bvumba road, 11 Feb. 1999, A. Verbeken 99-175 (GENT); Penhalonga, hill behind Harris’ garden, 9 Feb. 1999, A. Verbeken 99-143 (GENT, holotype).
Togo. Central Prov., Forêt Classée d’Alédjo, 11 Jul. 2007, A. De Kesel 4280 (BR 163674–35); Plateau Prov., Ile, route de l'ancien campement, 23 May 2008, A. De Kesel 4538 (BR 163490–45).
90. Lactarius undulatus Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 210 (1996).
90.1 Lactarius undulatus var. undulatus
– Type: Cameroon, Korup National Park, 30 Jan. 1989, R. Watling 21468 (E, holotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 219 & Pl. 243–245 (1996d); Verbeken, Persoonia 17: 396–398 (2001).
Pileus 75 mm diam., broadly infundibuliform with an umbilicate centre; margin straight, strongly undulate, strongly striate to grooved; pellis not dehiscent, mat, very finely felty, sepia-colour to horn-colour, ochraceous towards the centre. Stipe 42 × 11 mm, long, slender, cylindrical, mat, very finely felty, clay-buff to hazel-buff, cream at the apex. Lamellae adnate to slightly decurrent, unequal with lamellulae of different lenghts, very distant (L+l = 5/cm), cream to pale milky coffee-colour. Context with very acrid taste; smell unpleasant to mealy. Latex very abundant, white, becoming grey and then a bit brown; taste very acrid. Spore-deposit not observed.
Basidiospores ellipsoid, 8.2–10.0–11.8(–12.4) × (6.1–)6.7–7.9–9.2 mm (Q = 1.16–1.26–1.36, n = 20); ornamentation amyloid, composed of ridges up to 2(–2.5) mm high, forming a reticulum with strongly winged aspect; some low isolated warts present; wall sometimes rugose; plage strongly distally amyloid. Basidia 45–55 × 11–14 µm, subfusiform to fusiform, 4-spored; content guttulate. Pleurocystidia absent. Pleuropseudocystidia rather scarce, narrowly cylindrical, with rounded apex, 3–4 mm diam.; content oleiferic. Lamellar edge sterile; marginal cells not abundant because of the presence of narrow, parallel hyphae which border the lamella; some marginal cells narrowly cylindrical, with rounded apex, 10–30 × 3–5 mm, hyaline, thin-walled. Hymenophoral trama subregular, composed of hyaline, narrow hyphae; lactifers abundant. Pileipellis ixotrichoderm-like, two-layered; suprapellis composed of ascending elements, 20–60 × 5–9 mm, subclavate or subfusiform, sometimes septate, thin-walled, often with brownish content; subpellis composed of narrow, hyaline hyphae, locally with brownish content. Stipitipellis two-layered; suprapellis composed of ascending elements, 20–50 × 5–8 mm, cylindrical, subclavate or subfusiform, sometimes septate, thin-walled, often with brownish content; subpellis ixocutis-like, composed of narrow, hyaline hyphae, locally with brownish content. Clamp-connections absent. – Fig. 113
Ecology. Found in lowland rainforest.
Distribution. Only known from the type locality.
Revised specimens
Cameroon. South Western Prov., Korup National Park, near Mundema, transect P’-P24, 30 Jan. 1989, R. Watling 21468 (E, holotype).
90.2 Lactarius undulatus var. rasilis Verbeken
Verbeken, Bull. Jard. Bot. Nat. Belg. 65: 211 (1996). – Type: Democratic Republic of Congo, Kivu, Irangi, April 1972, J. Rammeloo Z415 (GENT, holotype).
Selected descriptions and icons. Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 220–221 & Pl. 246–248 (1996d); Verbeken, Persoonia 17: 398–400 (2001).
Pileus 70 mm diam., thin, elastic, broadly infundibuliform, with small, broad papilla in the centre; margin slightly deflexed, strongly undulate and crenulate, strongly striate to grooved; pellis not dehiscent, smooth, shiny, humid and slightly sticky, dark brown (5E6 to 5F6), paler in the centre, ochraceous brown in the centre. Stipe 55 × 8 mm, long, slender, cylindrical, slightly curved, smooth, mat, slightly striate in the upper part, brown (5E4), pale greyish in the upper part, fragile, fistulous, with ochraceous rhizomorphs at the base. Lamellae adnate to slightly decurrent, unequal with abundant lamellulae of different lenghts, very distant (L+l = 5/cm), rather broad (5–7 mm), moderately thick, elastic, not fragile, pale yellow to greyish yellow (4AB3); edge entire, concolorous. Context rather firm, thin, fragile in stipe, dirty white (6B2), changing pale greyish-brown in pileus, changing brownish orange (6C3) in stipe; taste very acrid; smell not remarkable. Latex very abundant, watery, white, changing to yellowish when drying, later greyish; taste very acrid. Spore-deposit pale cream. – Pl. 54
Chemical reactions. Context unchanging with anilated H2O, NH4OH, phenolaniline; brownish with NaOH; pale pinkish with phenol.
Basidiospores subglobose to ellipsoid, 8.2–9.2–10.3 × 7.3–8.1–8.8 mm (Q = 1.05–1.14–1.24; n = 20); ornamentation amyloid, composed of ridges up to 2(–2.5) mm high, forming a reticulum with strongly winged aspect; some low isolated warts present; wall sometimes rugose; plage strongly distally amyloid. Basidia 50–65 × 9–11 µm, cylindrical to subfusiform, 4-spored. Pleurocystidia absent. Pleuropseudocystidia scarce, narrowly cylindrical, with rounded apex, 3–4 mm diam.; content oleiferic. Lamellar edge sterile; marginal cells not abundant because of the presence of narrow, parallel hyphae which border the lamella; some marginal cells narrowly cylindrical, with rounded apex, 13–25 × 3–5 mm, hyaline, thin-walled. Hymenophoral trama subregular, composed of hyaline, narrow hyphae; lactifers abundant. Pileipellis one-layered, ixotrichoderm-like, composed of recumbent, narrow hyphae, more or less parallel; some terminal, very narrow (2–4 mm), ascending elements present, embedded in a slime-layer; locally brownish intracellular pigmentation. Stipitipellis ixocutis-like, composed of narrow, reumbent, parallel hyphae; locally brownish intracellular pigmentation. Clamp-connections absent. – Fig. 114
Ecology. Guineo‑Congolian rainforest.
Distribution. Only known from Democratic Republic of Congo.
Revised specimens
Democratic Republic of Congo. Kivu Prov., Irangi, April 1972, J. Rammeloo Z415 (GENT, holotype); Ibid., station IRSAC, 3 Jan. 1972, P. Van der Veken 8962 (GENT).
91. Lactarius badius Verbeken
Verbeken, Edinburgh J. Bot. 53: 54 (1996).
Clitocybe castanea Beeli, Bull. Soc. Roy. Bot. Belg. 60: 79 (1927, non Lactarius castaneus W.F. Chiu 1945). − Type: Democratic Republic of Congo, Motina valley, Bangala, March 1924, M. Goossens-Fontana 404 (BR 4909–59, holotype).
Selected descriptions and icons. Verbeken, Edinburgh J. Bot. 53: 54–56 (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 218–219 & Pl. 242–243 (1996d).
Pileus 80 mm diam., firm, planoconvex, slightly depressed in the centre; margin slightly incurved; pellis smooth, glabrous, chestnut-brown. Lamellae adnate, unequal with lamellulae, moderately narrow (4 mm), whitish. Stipe 100 × 14–20 mm, clavate, glabrous, smooth, whitish, with reddish brown shade, firm, solid. Context firm, rather thin in pileus (2 mm), white. Latex not observed. Spore-deposit not observed. (According to Beeli 1927). – Pl. 54
Basidiospores ellipsoid to elongate, 5.9–6.5–7.0(–8.0) × 4.0–4.7–5.6 mm (Q = 1.23–1.40–1.60; n = 20); ornamentation amyloid, composed of irregularly shaped and sized warts; warts sometimes aligned or forming ridges, never forming a complete reticulum, < 0.3 mm high; plage inamyloid. Basidia 32–40 × 6–8 mm, cylindrical to subclavate, 4-spored. Pleurocystidia scarce, 40–55 × 6–9 mm, clavate or fusiform, with rounded or tapering apex, arising deep in the subhymenium; content granular to needle-like. Pleuropseudocystidia rather abundant, 6–9 mm diam., often emergent, clavate or fusiform, with rounded or tapering apex; content granular or needle-like. Lamellar edge fertile. Hymenophoral trama composed of sphaerocytes and lactifers. Pileipellis a trichoderm; ascending terminal elements 10–30 × 4–6 mm, cylindrical to fusiform, tapering upwards; content yellowish brown. Stipitipellis composed of interwoven, long and slender hyphae which are 3–6 mm diam., thin-walled, sometimes branched; content locally yellowish brown. Clamp-connections absent. – Fig. 115
Ecology. Collected in a swamp forest.
Distribution. Only known from the type locality.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Bangala, valley of the Motima, March 1924, M. Goossens-Fontana 404 (BR 4909–59, holotype).
Notes. The macroscopic description is based on Beeli (1927b) and on the water-colour of Goossens-Fontana. The microscopic description is based on our own observations on the type material which consists of one basidiome in good condition.
This species is particularly characterized by the very small and elongate spores. It is also remarkable that true pleurocystidia and pseudocystidia have the same size and shape. True pleurocystidia arise rather deep in the subhymenium, but there is a septum and they are not connected with a lactifer, unlike the pseudocystidia.
Beeli (1933) stated that Clitocybe isolata Beeli seemed identical with his C. castanea although the remains of the type-specimen of that species do not belong to the Russulaceae.
The pileipellis structure argues for a relationship to section Edules or Amari, but other characters are not reconciliable with these sections. Furthermore, the lack of macroscopic data makes it impossible to classify this species with certitude.
According to Beeli (1927b) this species is consumed by local populations.
92. Lactarius caperatus R. Heim & Gooss.‑Font.
Heim, Bull. Jard. Bot. Etat Bxl 25: 36 (1955). – Type: Democratic Republic of Congo, Binga, Aug. 1947, M. Goossens-Fontana 4078 (BR 7735–72, lectotype designated here).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 36–38 & Pl. 3, fig. 1a–c (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 14, fig. 5a–b (1955).
Pileus 40–70 mm diam., fleshy, strongly curved, then applanate, orbicular-sinuose, depressed, sometimes infundibuliform, with or without papilla; margin straight, sometimes slightly incurved; pellis not dehiscent, rugose-venose, concentrically wrinkled, with gelified veins; veins more abundant near the margin, but more prominent in the centre; dry, smooth, vivid ochraceous orange, dull ochre near the margin and darker brownish orange near the centre in dried condition. Lamellae decurrent, unequal, dense, narrow, thin, brittle, whitish to cream, then pale ochraceous. Stipe 40–70 × 10 mm, first curved, then straight, long and regularly cylindrical, almost smooth, especially in the upper part longitudinally striate, concolorous with the pileus or slightly paler, whitish tomentose at the base, solid, longitudinally deeply grooved in dry condition. Context firm and thick in the pileus, caseous, cream; taste very acrid. Latex abundant, whitish translucid, unchanging; taste very acrid. Spore-deposit white. (According to Heim 1955a). – Pl. 54
Basidiospores globose to subglobose, sometimes broadly ellipsoid, 7.0–7.6–8.3(–8.4) × 6.1–6.9–7.7 µm (Q = 1.01–1.11–1.22, n = 20); ornamentation amyloid, composed of rounded to irregular warts, up to 0.3(–0.5) µm high, mostly isolated, sometimes aligned; plage not amyloid. Basidia 50–60 × 10–12 µm, subclavate to subcylindric, 4-spored. Pleuromacrocystidia abundant, slightly emergent, 55–65 × 8–10 µm, thin-walled, cylindric, slightly clavate, usually rounded on top, with dense needle-like content. Pleuropseudocystidia scarce, not emergent, 5–7 µm diam., cylindric, slightly tortuous. Hymenophoral trama cellular. Lamellar edge sterile; marginal cells irregularly shaped, shortly cylindrical to subfusiform, thin-walled, hyaline, 6–30 × 5–10 µm; cheilomacrocystidia subcylindric to subclavate, 30–40 × 7–9 µm, with needle-like content. Pileipellis a trichopalisade, 70–100 µm thick; terminal elements cylindric or slightly irregular, sometimes subclavate, with rounded apex, 25–30 × 4–6 µm, thin-walled, but walls somewhat slightly refringent; some elements dermatocystidia-like because of the needle-like content. Clamp-connections absent. – Fig. 116
Ecology. Found in drier Guineo‑Congolian rainforest with Gilbertiodendron dewevrei.
Distribution. Only known from the type locality.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Binga, Jan. 1947, M. Goossens-Fontana 4038 (BR 7733–70); Ibid., Aug. 1947, M. Goossens-Fontana 4078 (BR 7735–72, lectotype designated here).
Notes. In the original description two collections are cited, without indicating a type. Therefore, a lectotype is selected here.
93. Lactarius hispidulus R. Heim
Heim, Bull. Jard. Bot. Etat Bxl 25: 75 (1955). – Type: Democratic Republic of Congo, Binga, 9 Aug. 1929, M. Goossens-Fontana 725 (BR 7717–54, holotype).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 75-76 & Pl. 6, fig. 5a–b (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 15, fig. 3 (1955).
Pileus 22–34 mm diam., curved, convex, depressed; margin involute; pellis completely squamose, tomentose, rough, ochraceous reddish. Lamellae emarginate to subadnate, unequal, rather distant (L+l = 30/total), rather broad halfway radius, rather thick, cream. Stipe 25–30 × 3–5 mm, cylindrical, tapering downwards, smooth, unicolorous, ochraceous reddish, darker than the pileus, fistulous. Context white; taste acrid; smell pepper-like, acrid. Latex very abundant, white; taste acrid. Spore-deposit white. (According to Heim 1955a). – Pl. 54
Basidiospores subglobose to broadly ellipsoid, 7.4–8.2–8.4–9.3 × 6.2–7.2–7.3–8.3 mm (n = 40, Q = 1.05–1.14–1.17–1.24); ornamentation amyloid, up to 1.5(2) µm high, composed of a rather spiny and acute short ridges, forming a broken and irregular reticulum; some isolated spines also present; plage not amyloid or slightly distally amyloid. Basidia 35–45 × 10–12 mm, cylindrical to subclavate, 4-spored, with guttate content. Pleurocystidia absent. Pleuropseudocystidia rather abundant, sometimes slightly emergent, but usually not emergent, 3–4 µm diam., cylindrical. Lamellar edge sterile; marginal cells occupying the whole margin and continuing up to the sides of the lamellae, (20–)30–40(–45) × 5–10 µm, variously shaped, some cylindric to subclavate, some fusiform, with slightly refringent walls. Hymenophoral trama composed of hyphae and abundant laticiferous hyphae. Pileipellis a trichoderm, composed of broad (up to 8 µm) and rigid hyphae with refringent and slightly thickened walls, multi-septate, forming long, erecting tufts, up to 250 µm high; within the tufts hyphae sometimes branching and sometimes with irregular bulges or pseudoclamps. – Fig. 117
Ecology. Guineo-congolian rainforest.
Distribution. Only known from Democratic Republic of Congo and Guinea.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Binga, 9 Aug. 1929, M. Goossens-Fontana 725 (BR 7717–54, holotype).
Guinea. Forêt humide de Ziama, piste d’Aningueria, 27 July 2000, A. Bâ 151 & 152 (all GENT).
Notes. The microscopical description is based on the type material and on Bâ 151.
Heim described the species as a small Lactarius helvus with larger and more heavily ornamented and clearly reticulate spores and acrid latex. He proposed to classify the species in Lactarius section Floccosi Kühner (invalid).
94. Lactarius russuliformis (Beeli) Verbeken
Verbeken, Edinburgh J. Bot. 53: 60 (1996). – Omphalia russuliformis Beeli, Bull. Soc. Roy. Bot. Belg. 60: 83 (1927, as "russulaeformis"). – Type: Democratic Republic of Congo, Eala, June 923, M. Goossens-Fontana 136 (BR 5014–67, holotype).
Lactarius pellicularis R. Heim, Bull. Jard. Bot. Etat Bxl 25: 85 (1955).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 85–87 & Pl. 6, fig. 1a–d (1955); Heim, Fl. Icon. Champ. Congo 4: Pl. 15, fig. 9 (1955); Verbeken, Edinburgh J. Bot. 53: 60–63: (1996); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 221–222 & Pl. 249–251 (1996d).
Pileus (18–)25–35 mm diam., very thin, planoconvex to depressed and infundibuliform; centre with papilla; margin striate to grooved; pellis not dehiscent, pellicular, smooth, cupreous red with a lilac tinge. Lamellae decurrent (sometimes adnate-decurrent), unequal with lamellulae, distant (L+l = 5+4/cm), medium broad (< 6 mm), rather thick, brittle, yellowish white, with a pink tinge. Stipe 20–30 × 4–7 mm, long, slender, cylindrical to subfusiform, sometimes curved, smooth, cupreous red with a lilac tinge, paler at the apex, fragile, often fistulous. Context granulate, brittle, ochraceous pink, more intense pink near the base; taste bitter and very acrid; smell bitter. Latex white to yellowish; taste acrid. Spore-deposit white. – Pl. 54
Chemical reactions. Pileus greenish with FeSO4, brown with gaiac.
Basidiospores ellipsoid, 7.9–9.2–9.8–10.9 × 7.3–7.8–8.4–9.0 mm (Q = 1.07–1.16–1.19–1.26; n = 60); ornamentation amyloid, composed of slender spines, (1–)1.5–2.0 mm high, fairly regularly connected by fine connective lines; wall with slightly amyloid spots; plage inamyloid. Basidia 36–55 × 10–14 mm, cylindrical to clavate, 4-spored; content slightly granular; sterigmata 6–10 × 1–2.5 mm. Pleurocystidia absent. Pleuropseudocystidia fairly abundant, 6–7(–9) mm diam., cylindrical to moniliform; content densely oleiferic, sometimes slightly granular. Lamellar edge sterile; marginal cells 10–15(–20) × 4–6 mm, shortly cylindrical to clavate, often septate. Hymenophoral trama irregular, composed of hyaline, thin-walled hyphae, 3–4 mm diam.; oleiferic and laticiferous hyphae abundant, < 9 mm diam. Pileipellis a palisade, hymeniderm-like, 60–100 mm thick; elements of the suprapellis 15–35 × 7–10 mm, cylindrical to fusiform and tapering upwards, slightly thick-walled (< 0.5 mm), sometimes branched and septate. Stipitipellis a trichoderm; hyphae long and slender, often septate; terminal elements 10–20 × 5–8 mm, isodiametric to cylindrical or fusiform, thin-walled. Clamp-connections absent. – Fig. 118
Ecology. Found in swamp forest and Guineo-Congolian rainforests, also on dead wood.
Distribution. Only known from Democratic Republic of Congo.
Revised specimens
Democratic Republic of Congo. Equateur Prov., Binga, on the ground, 25 July 1931, M. Goossens-Fontana 926 (BR 7720–57); Eala, June 1923, M. Goossens-Fontana 136 (BR 5014–67, holotype); Tshopo Prov., 5 km NNE of Batiabongena, 27 April 1984, B. Buyck 1589 (BR 12748–41).
Notes. The type-specimen is in a bad condition but both spore ornamentation and size, as well as the water-colour, agree with the type of Lactarius pellicularis (except that on the spores of Goossens-Fontana 136 some shorter and more rounded warts are also observed).
The description is based on Goossens-Fontana 926 and Buyck 1589 (but spore measurements are based on all three available collections). The colour of the context in Buyck 1589 is brownish, the taste is acrid; its pileus measures 1.8 cm diam. and is brown with a reddish tinge (6–7E8) in the centre and light orange to greyish orange (5A4–5B5) near the margin, some lamellae are forked.
Heim (1955) put "Lactarius pellicularis" and L. chamaeleontinus in Lactarius section Pelliculares R. Heim (invalid, descr. gall.). Both species share indeed the pellicular pileus with deeply striate to grooved margin and vivid colour, the fistulous stipe, the distant lamellae and the dirty white, acrid latex but are considered here as not closely related given the totally different pileipellis-structure and the different spore-ornamentation. Lactarius roseolus is macroscopically somewhat similar to L. russuliformis but the spores are very different. Also the structure of the pilei- and stipitipellis of this species are very similar to those of Lactarius russuliformis.
1. Lactarius fulgens R. Heim
Heim, Candollea 7: 377 (1938). – Type: Madagascar, North of the Isle of St.‑Marie, 14 Dec. 1934, R. Heim I40 (PC, holotype).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 44–45 & Pl. 1d (1938); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 193 & Pl. 189 (1996d).
Pileus 30–40 mm diam., irregularly convex, then depressed; margin strongly recurved, crenulate; pellis finely tomentose, thick, brownish yellow to orange. Lamellae arcuate-decurrent, crowded, fairly thick, narrow (1–1.2 mm), orange; edge entire, concolorous. Stipe 30 × 5–10 mm, tapering downwards, finely tomentose, cream, becoming orange when bruised or with age, white near apex; solid. Context white; taste mild; smell of linseed oil. Latex abundant, white, unchanging; taste mild. Spore-deposit not observed. (According to Heim 1938b).
Chemical reactions. Context weakly greenish grey, then greenish with gaiac; first negatif, then dark lilac with gaiacol; first negative, then lilac with pyramidon; unchanging with FeSO4. (According to Heim 1938b).
Basidiospores subglobose to ellipsoid, 4.6–5.3–5.8 × 3.8–4.5–5.0 mm (Q = 1.06–1.19–1.31 n = 30); ornamentation amyloid, composed of irregular ridges and warts, forming an almost complete reticulum; ridges up to 0.8 mm high; plage inamyloid. Basidia 35–45 × 7–9 mm, cylindrical, 4-spored. True pleurocystidia abundant, 35–50 × 4–8 mm, cylindrical, tapering upwards, often very irregularly branched at the apex, or with irregular bulges; content granular. Pleuropseudocystidia fairly abundant; slightly emergent, 2–5 mm diam., cylindrical to tortuous, with rounded apex; content oleiferic. – Fig. 119
Ecology. Malagasy lowland rainforest.
Distribution. Only known from the type locality.
Revised specimens
Madagascar. Betsimisaraka Prov., North of the Isle of St. Marie, 14 Dec. 1934, R. Heim I40 (PC, holotype).
Notes. The type-material consist of one basidiome which had been preserved in alcohol. As the alcohol has evaporated in the course of time, the specimen is in very bad condition. The spore ornamentation can hardly be observed in Melzer's reagents; neither could the structure of the pilei- and stipitipellis be identified. The former consists of small and short cylindrical elements, 10–30 × 3–6 mm, thin-walled and with granular content. Those were only observed in surface view.
The macroscopic description is completely based on the notes of Heim (1938b), the microscopic description on own observations of the type-specimen.
Apparently, the species shares some macroscopic characters with Lactarius acutus, but here the pileipellis is brownish yellow to orange against dark brown in L. acutus. The pileus is not papillate in Lactarius fulgens, the spores in L. fulgens are smaller with a lower ornamention.
2. Lactarius fulgens var. africanus R. Heim
Heim, Bull. Jard. Bot. Etat Bxl 25: 70 (1955).− Type: Ivory Coast, Toulépleu forest, near Danané, 22 March 1939, R. Heim B21 (PC, holotype).
Selected descriptions and icons. Heim, Bull. Jard. Bot. Etat Bxl 25: 70–72 (1955); Verbeken, Biodiversity of the genus Lactarius Pers. in tropical Africa: 194 & Pl. 189 (1996d).
Pileus 40–50 mm diam., irregularly convex, then depressed; margin first recurved, then straight; pellis not dehiscent, mat, finely wrinkled, thin, smooth, finely tomentose, dull reddish brown. Lamellae sinuose-subdecurrent with decurrent tooth, crowded, fairly thick, rather narrow (< 4.5 mm), pale yellow; edge entire, concolorous. Stipe 30 × 5–10 mm, cylindrical, tapering downwards, smooth, finely tomentose, cream, solid, firm. Context rather thick (7–8 mm) in pileus, granulose in stipe, cream in stipe, pinkish under stipitipellis, dull reddish in pileus; taste mild; smell not particular. Latex scarce, "not coloured". Spore-deposit not observed. (According to Heim 1955a).
Chemical reactions. Context in pileus reacting strongly with gaiac, in stipe more weakly with gaiac; unchanging with FeSO4, NH3 and phenol. (According to Heim 1955a).
Basidiospores globose to subglobose, sometimes ellipsoid, 5.2–6.0–6.8 × 4.7–5.4–6.1 mm (Q = 1.01–1.11–1.17; n = 30); ornamentation amyloid, composed of irregular ridges and warts, forming an almost complete reticulum; ridges up to 0.8 mm high; plage inamyloid. Basidia 35–45 × 8–10 mm, cylindrical to subclavate, 4-spored. Pleurocystidia not observed. Pleuropseudocystidia fairly abundant, 2–5 mm diam., cylindrical to tortuous, with rounded apex; content oleiferic. Hymenophoral trama irregular, composed of narrow, hyaline hyphae and abundant lactifers. – Fig. 120
Ecology. Guineo‑Congolian rainforest.
Distribution. Only known from the type locality.
Revised specimens
Ivory Coast. Toulépleu forest, near Danané, 22 March 1939, R. Heim B21 (PC, holotype).
Notes. The type-material consists of a small piece of a basidiome, in very bad condition. The spore-ornamentation is hardly visible in Melzer's reagens; most probably the specimen has been preserved in alcohol before drying completely. Given the bad condition of the specimen, it was impossible to identify the structure of stipiti- and pileipellis. A surface view of the pileipellis could be observed. Probably it consists of pseudoparenchymatous subpellis with a suprapellis composed of slender cylindrical elements, 20–35 × 4–8 mm.
Lactarius fulgens var. africanus most probably represents a good taxon but more collections and observations are needed to clarify its taxonomic level.
Heim, Candollea 7: 381 (1938). – Type: Madagascar, Antanambé, 18 Dec. 1934, R. Heim I75 (PC, holotype).
Selected descriptions and icons. Heim, Prodr. Fl. Mycol. Madagascar 1: 64–66 (1938).
Pileus 50–70 mm diam., irregular, strongly depressed in the centre; margin thin, deeply striate; pellis dehiscent, smooth, dry, creamish orange, bright ochraceous in the centre. Lamellae subadnate, distant, moderately narrow, rather thick, cream to pinkish. Stipe 30 × 14–20 mm, short cylindrical, tapering downwards, felty, white, then cream, solid, then fistulous; with abundant, white mycelium at the base. Context soft and cotton-like, brittle, white, unchanging; smell crustacean-like; taste slightly acrid. Latex rather scarce, hyaline, unchanging; taste slightly acrid. Spore-deposit white. (According to Heim 1938b).
Chemical reactions. Context unchanging with pyramidon and FeSO4; slowly blue-green with gaiac; slowly pink, then purple with gaiacol. (According to Heim 1938b).
Basidiospores broadly obovoid, 7.8–9 × 7.2–7.8 mm; ornamentation composed of distant, unequal, low warts, isolated or connected by fine connective lines; plage inamyloid. Basidia 40–45 × 9 mm, fusiform, 4-spored; sterigmata 8 mm, spiniform.
Ecology. Found in Malagasy lowland rainforest.
Distribution. Only known from the type locality.
Revised specimens
Madagascar. Betsimisaraka Prov., Antanambé, Dec. 1934, R. Heim I75 (PC, holotype).
Notes. The type-specimen consists of one half basidiome in very bad condition. It is extremely brittle and does not swell in potassiumhydroxide. The macroscopic, as well as the microscopic notes, are taken from Heim. The microscopic description is very incomplete; Heim mentioned that the other microscopic characters could not be specified because of the poor condition of the preserved material.
The species could not be classified due to the significant amount of missing data. Heim (1938b) mentions that this is clearly a species of the group “Pruinosi” Quél., with scarce, hyaline and hardly acrid latex, with a undistinct smell, between that of his group “Subdulces” R. Heim (invalid) and the “Olentes Bataille”. Lactarius striatus is easily characterized by its dehiscent and deeply striate pileipellis, its agregated mycelium and its hyaline latex. Heim proposed to put the species in a a group “Striatini” R. Heim (invalid.), opposed to the “Pruinosi” with smooth pileipellis, which would then be named “Laevini” R. Heim (invalid)
1. Lactarius lignyotus ss. Dade (1940)
Notes. Based on the collection Bustar & Codron s.n. (K!). It does not belong to Lactarius.
2. Lactarius pellucidus Gooss.‑Font. & R. Heim
Heim, Bull. Jard. Bot. Etat Bxl 25: 61 (1955). Russula pellucida (Gooss.‑Font. & R. Heim) Buyck, Bull. Jard. Bot. Nat. Belg. 59: 241 (1989). – Type: Democratic Republic of Congo, Binga, Feb. 1942, M. Goossens-Fontana 2079 (BR 6139–28, holotype).
Notes. This is a Russula species (Buyck 1993).
3. Lactarius piperatus ss. Pegler & Piearce (1980) & Piearce (1981)
Notes. Records of this species are based on Piearce FP600/1a (K(M) 34213) which represents Lactarius densifolius.
4. Lactarius trivialis ss. To & Ob (2005)
Notes. Reported from Akoko land, Nigeria. Material not obtained, but this identification is considered as presumably wrong.
5. Lactarius uvidus ss. Bresadola. & Saccardo (1899) & De Wildeman & Durand (1901)
Notes. Records of this species are based on the collection Dewèvre 111 (BR 12289–67) which belongs to Russula sp.
6. Lactarius vellereus ss. Morris (1984, 1987, 1990) & Williamson (1975)
Notes. Records of this species are based on collection Williamson 629 (K(M) 34216), which represents Lactarius medusae.
De Wildeman, Bull. Jard. Bot. Etat Bxl. 4: 25 (1914, non Lactarius velutinus Bertillon 1865). – Russula velutina (Bres.) Buyck, Bull. Jard. Bot. Nat. Belg. 58: 476 (1988). – Type: Democratic Republic of Congo, Longo, region of Bambatas, 1910, leg Allard in Vanderyst 162 (BR 12287–65, holotype).
Notes. This is a Russula species (Buyck 1993).
8. Lentinus clitocyboides Henn.
Bot. Jahrb. Syst. 30: 45 (1902). – Russula clitocyboides (Henn.) Verbeken & Buyck, in Verbeken & al., Cryptog. Mycol. 17: 7 (1996). – Type: Cameroon, Bipindi, Sept. 1898, Zenker 1899 (type not traced; K!, PC, isotypes).
Russula subfistulosa var. apsila Buyck, Bull. Jard. Bot. Nat. Belg. 60: 193 (1990). – Type: Democratic Republic of Congo, Binga, Sept. 1938, M. Goossens-Fontana 2099c (BR 4927–77, holotype).
Notes. This species does not belong to Lactarius as suggested by Pegler (1983: 229) but to Russula (Verbeken & al. 1996).
Translation of the keys into French
Clé pratique des sections du genre Lactarius en Afrique tropicale
1. Carpophores gastéroïdes ou secotioïdes........................................... sect. Plinthogali s.l.
Carpophores avec un stipe et des lamelles normalement développés......................................... 2
2. Espèces associées à des Pinus exotiques; chapeau brun rouge........ sect. Russularia
Espèces associées à des arbres indigènes (principalement Caesalpinioideae, Uapaca, Papilionoideae, Dipterocarpaceae) .. 3
3. Stipe avec un anneau distinct, plus ou moins fugace ....................... sect. Lactariopsis
Stipe sans anneau; restes de voile présents seulement à la marge du chapeau ou absents.. 4
4. Marge du chapeau avec des restes de voile .................................. sect. Chamaeleontini
Restes de voile absents ........................................................................................................................ 5
5. Contexte noircissant .............................................................................................................................. 6
Contexte non noircissant (brunissant, immuable etc.) .................................................................... 7
6. Hyménium avec des lamprocystides; spores ornementées, à petites verrues ou crêtes ....... sect. Rubroviolascentini
Hyménium sans lamprocystides; spores à ornementation ailée, d’au moins 1 µm de haut .... sect. Nigrescentes
7. Sporée et lamelles brun chocolat à maturité; chapeau visqueux ............. sect. Chromospermi
Sporée blanche à jaunâtre ou rose pâle, au plus brun clair; chapeau sec ou visqueux ......... 8
8. Chapeau visqueux à marge fortement poilue OU stipe scrobiculé .. sect. Piperites s.l.
Chapeau visqueux ou sec, à marge non poilue, stipe jamais scrobiculé .................................... 9
9. Chapeau pruineux, orange et zoné concentriquement à l’état humide; arête des lamelles fimbriée, appendiculée, avec des gouttelettes transparentes à l’état frais; stipe rétréci vers la base et clairement creux ...... sect. Aurantiifolii
Pas cette combinaison de caractères ............................................................................................... 10
10. Hyménium avec des lamprocystides bien visibles ................ sect. Pseudogymnocarpi
Hyménium sans lamprocystides .................................................................. 11
11. Pileipellis constitué d’une palissade avec une couche supérieure d’éléments anticlinaux, à paroi épaisse (lampropalissade); contexte (lamelles) brunissant souvent au froissement .......12
Pileipellis à structure différente, dans le cas d’une palissade sans éléments à paroi épaisse ....... 13
12. Ornementation sporale typiquement composée de verrues arrondies distinctes et presque isolées, séparées par quelques connectifs très fins .................... sect. Phlebonemi
Ornementation sporale plus ou moins réticulée ....................... sect. Polysphaerophori
13. Ornementation sporale zébroïde ......................................... sect. Plinthogali
Ornementation sporale différente ............................................................ 14
14. Chapeau mat (brun, grisâtre, café au lait, chamois), sans teintes orange ou rougeâtres; spores souvent ailées ou réticulées, parfois (sub)globuleuses ........................................................ 15
Chapeau coloré différemment, souvent plus vif, avec des teintes orange ou rougeâtres ..... 17
15. Spores petites, 6–7 × 4–5.5 µm, ellipsoïdes à allongées; pileipellis en trichoderme ......... L. badius
Spores plus grandes ou subglobuleuses; pileipellis en palissade ou trichopalissade .......... 16
16. Pileipellis en palissade ou hyméniderme, sec, souvent duveteux ou finement tomenteux à légèrement squamuleux; contexte virant parfois au rosâtre au froissement ................. sect. Plinthogali
Pileipellis en trichoderme, lisse, parfois brillant ou subvisqueux; contexte ne virant pas au rosâtre au froissement ........................... sect. Pseudofuliginosi
17. Carpophores assez petits; chapeau ocracé, zoné de manière concentrique à l’état frais; stipe fistuleux; pileipellis en cutis ou trichoderme, composé d’hyphes longs entrelacés ..... sect. Amari
Pas cette combinaison de caractères ............................................................................................... 18
18. Pileipellis en palissade, à couche supérieure composée d’éléments anticlinaux; basides longues et minces; spores ellipsoïdes à allongées, verruqueuses à plus ou moins réticulées, plage souvent avec une tache amyloïde au centre; chapeau sec, orange, jaunâtre, rougeâtre, rouge brun (sub)globuleux ......................... sect. Rugati
Pileipellis à structure différente ......................................................................................................... 19
19. Pileipellis en ixocutis (à localement ixotrichoderme) de couleur crème. sect. Piperites
Pileipellis sec ou visqueux mais pas en ixocutis, chapeau orange si en ixotrichoderme ...... 20
20. Pileipellis le plus souvent avec des poils épars à paroi épaisse; chapeau brillant et lisse, sec à visqueux, marge souvent striée ou sulquée; pleuropseudocystides émergentes à très émergentes, surtout larges et fusiformes ..... sect. Chamaeleontini
Pileipellis sec, sans poils à paroi épaisse ....................................................................................... 21
21. Chapeau < 35 mm diam., rouge ocre, rouge brunâtre, tomenteux à finement écailleux ............. L. hispidulus
Pas cette combinaison de caractères ............................................................................................... 22
22. Basidiomes rappelant une petite Russula rougeâtre ou rosâtre; ornementation sporale composée d’épines minces de 1.5–2 µm de haut, reliées régulièrement par de fins connectifs..................................... L. russuliformis
Pas cette combinaison de caractères ............................................................................................... 23
23. Chapeau orange ocracé vif; lamelles denses; spores à verrues arrondies généralement isolées; pleuromacrocystides abondantes, légèrement émergentes et à contenu dense constitué de cristaux à l’aspect d’aiguilles ......... L. caperatus
Pas cette combinaison de caractères ............................................................................................... 24
24. Pileipellis sec, en trichoderme ou composé de chaînes de cellules subcylindriques à subsphériques (chapeau souvent craquelé); piléomacrocystides absentes ....... sect. Edules
Chapeau sec à légèrement visqueux; pileipellis en cutis avec des macrocystides distinctes; chapeau souvent rougeâtre; pseudocystides souvent ramifiées et diverticulées ........ sect. Russulopsidei
Sect. Lactariopsis (Henn.) Singer
1. Eléments en forme de poil à paroi épaisse rares dans le pileipellis, pleurocystides présentes ................................................. sect. Chamaeleontini, 6. L. indusiatus
Pileipellis entièrement constitué d’éléments en forme de poil à paroi épaisse, pleurocystides absentes ........ 2
2. Ornementation sporale bien développée, jusque 1 mm de haut ..... 1. L. annulatoangustifolius
Ornementation sporale à peine visible, généralement jusque 0.2 mm de haut ............................. 3
3. Spores ellipsoïdes à allongées, Q = (1.15)1.3–1.5(1.65), spores jusque 11.5 mm de long; ornementation sporale formant presque un réseau complet ................................... 4. L. heimii
Spores ellipsoïdes, parfois subglobuleuses, Q = (1.09)1.19–1.32(1.45), spores généralement inférieures à 9(10) mm de long; ornementation sporale formant un réseau incomplet, toujours avec des verrues distinctes .... 4
4. Pileipellis remarquablement épais, feutré, blanc jaunâtre, persistant, se rompant en étoile; spores 8.4–9.2 mm de long; connu uniquement du miombo en région zambézienne ......... 5. L. velutissimus
Pileipellis plus fin, submembraneux .................................................. 5
5. Basidiomes fragiles, brun jaunâtre à ocracé pâle; spores petites, 7.1–7.8 ´ 5.5–6.3 mm, à ornementation très faible; cellules marginales tortueuses et ramifiées .............. 2. L. zenkeri
Basidiomes plus fermes, ocre pâle à jaunâtre pâle; spores plus grandes, 8.0–8.6 ´ 6.4–7.0 mm, ornementation 0.2 mm de haut; cellules marginales étroitement utriformes à côniques .......... 3. L. pelliculatus
Sect. Chamaeleontini Verbeken
1. Leptocystides hyméniales assez abondantes, 65–80 × 8–10 µm, cylindriques à subclavées, moniliformes ou terminées par un rostre. Voile présent chez les jeunes basidiomes se présentant comme un indusium arachnoïde entre le stipe et la marge du chapeau, parfois persistant sous forme d’un anneau très petit chez les basidiomes plus âgés; éléments en forme de poil à paroi épaisse (très) rares dans le pileipellis; contexte virant au rosé ou au rougeâtre, puis au vert grisâtre ................................ 6. L. indusiatus
Pleurocystides absentes; éléments en forme de poil à paroi épaisse absents à abondants dans le pileipellis; restes de voile ne formant jamais d’anneau ou absents .................................... 2
2. Pileipellis sans éléments à paroi épaisse ................................................................ 10. L. laevigatus
Pileipellis avec des éléments en forme de poil à paroi épaisse plus ou moins abondants, parfois difficiles à trouver ...... 3
3. Carpophore ferme; chapeau blanc sale à crème jaunâtre, souvent couvert de terre ......... 13. L. emergens
Pas cette combinaison de caractères ....................................................... 4
4. Jeune chapeau et stipe entièrement blanchâtres pruineux, nombreux éléments terminaux à paroi épaisse dans le pileipellis ....... 11. L. pruinatus
Chapeau et stipe pas entièrement pruineux, éléments terminaux à paroi épaisse rares ou nombreux dans le pileipellis ..... 5
5. Espèce de petite taille, chapeau < 2 cm diam.; tous les éléments terminaux du pileipellis à paroi épaisse, subfusiformes à subcylindriques, 15–35 × 6–9 µm, spores en moyenne 10 µm de long ...... 12. L. uapacae
Chapeau le plus souvent > 2 cm diam.; éléments du pileipellis à paroi épaisse épars, plus longs et plus fins, spores en moyenne plus courtes ............................... 6
6. Basidiocarpes plus ou moins entièrement orange jaunâtre brillant, souvent collants; pseudocystides effilées ou délicatement capitées. Espèce associée à Brachystegia spp. ........ 14. L. brachystegiae
Basidiocarpes colorés différemment; pseudocystides pas délicatement capitées ....................... 7
7. Ornementation sporale peu visible, pas plus de 0.2 µm de haut ........................ 9. L. sesemotani
Ornementation sporale jusque 0.5 µm de haut ................................................................................... 7
8. Chapeau jusque 5 cm diam., fin et fragile, orange pâle, ocracé ou brun rougeâtre; marge regulièrement striée à cannelée ..................... 7. L. chamaeleontinus
Chapeau jusque 8 cm diam., plus ferme, jaune pâle à jaune paille; marge irrégulière ........... 8. L. madagascariensis
Sect. Russulopsidei Verbeken
1. Trame de l’hyménophore composée principalement de sphérocystes .......................................... 2
Trame de l’hyménophore irrégulière à régulière, composée principalement d’hyphes hyalins, plus ou moins parallèles, occasionnellement avec quelques sphérocystes .................................. 6
2. Ornementation sporale composée d’épines côniques, pointues à légèrement arrondies, jusque 1.5 mm de haut, reliées par de fins connectifs formant un réseau presque complet ....... 22. L. corbula
Ornementation sporale composée de verrues irrégulières, arrondies et obtuses, isolées ou reliées par de fins connectifs ....... 3
3. Marge des lamelles brun rougeâtre ....................................... 19. L. rufomarginatus
Marge des lamelles concolore .................................................................... 4
4. Goût doux; éléments du pileipellis sans protubérances remarquables ......... 20. L. pseudotorminosus
Goût très âcre; éléments du pileipellis avec des protubérances diverticulées remarquables ..... 5
5. Stipe courtement cylindrique; lamelles assez distantes (3+7 to 4+8/cm); chapeau virant au brun foncé grisâtre à partir de la marge en vieillissant et en pourrissant; latex immuable; pleurocystides généralement présentes et abondantes ............................ 15. L. ruvubuensis
Stipe long et fin; lamelles très serrées (22–32/cm); chapeau devenant plus pâle en vieillissant; latex parfois grisâtre à noirâtre en séchant; pleurocystides absentes ............ 16. L. longipes
6. Latex bleu verdâtre en séchant; spores très faiblement ornementées; chapeau unicolore ......... 17. L. cyanovirescens
Latex immuable en séchant; ornementation sporale 0.2–0.5 mm de haut; chapeau distinctement plus foncé au centre et plus pâle vers la marge ........................................ 7
7. Pileipellis avec des dermatocystides clairement moniliformes; lamelles serrées et étroites (2–2.5 mm); latex légèrement âcre .............. 21. L. claricolor
Certains éléments du pileipellis lactifères et rappelant des dermatocystides; lamelles assez distantes et plus larges (3–13 mm); latex âcre brûlant ............................................ 18. L. urens
Sect. Edules Verbeken
1. Pileipellis entièrement composé d’hyphes longs et étroits entrelacés ........................................... 2
Pileipellis composé de chaînes d’éléments subcylindriques à subsphériques ........................... 3
2. Goût acre brûlant; lamelles distantes, jaune pâle à jaune d’or; pleurocystides absentes ........ 26. L. aureifolius
Goût doux; lamelles très serrées, de couleur crème; pleurocystides assez abondantes ........ 27. L. densifolius
3. Chapeau < 3 cm diam., rose à brun rougeâtre, basidiocarpe souvent fragile et délicat ......... 30. L. roseolus
Chapeau plus grand, basidiocarpe généralement ferme ................................................................. 4
4. Lamelles peu à très distantes; stipe court et épais ............................................................................. 5
Lamelles (très) serrées; stipe long et fin ...................................................................... 28. L. inversus
5. Chapeau à teinte quelque peu olivacée; lamelles ramifiées dichotomiquement vers la marge; marge du chapeau striée et translucide .................................................... 29. L. phlebophyllus
Chapeau jamais à teinte olivacée; lamelles non ramifiées dichotomiquement; marge parfois striée chez les vieux spécimens, non translucide .......................................... 6
6. Chapeau, spécialement à l’état jeune, brun rougeâtre et brun cannelle; lamelles orange, orange jaunâtre à orange brunâtre clair; éléments du pileipellis contenant souvent de petits granules foncés intracellulaires ..... 25. L. latifolius
Chapeau principalement jaunâtre à couleur crème, à teinte rougeâtre seulement sur les exsiccata; lamelles de couleur crème; éléments du pileipellis sans granules intracellulaires ........ 7
7. Ornementation sporale jusque 0.3 µm de haut, constituée d’éléments verruqueux plus ou moins reliés; macrocystides absentes ........... 23. L. edulis
Ornementation sporale moins prononcée, jamais plus de 0.2 µm de haut; macrocystides très abondantes et portant des protubérances, comme les pseudocystides ......... 24. L. nodosicystidiosus
Sect. Rubroviolascentini (Singer) Verbeken
1. Chapeau ocre brunâtre à orange rougeâtre; contexte virant au noirâtre, puis au rougeâtre; spores très faiblement ornementées à presque lisses; plage non amyloïde ........ 32. L. rubroviolascens
Chapeau chamois clair; contexte virant au rouge puis au lilas et noircissant finalement fortement par le latex; spores avec un réseau distinct incomplet, plage amyloïde au centre ........ 33. L. denigricans
Sect. Pseudogymnocarpi Verbeken
1. Pleurocystides abondantes et émergentes .................................. 2
Pleurocystides présentes mais peu visibles, non émergentes; stipe et lamelles remarquablement jaunes à jaune verdâtre ................................... 40. L. luteopus
2. Spores en moyenne 12 µm de long, 9 µm de large; chapeau < 30 mm diam., d’un rouge carmin saisissant ............................... 39. L. carmineus
Spores plus petites, chapeau coloré différemment ............................................................................ 3
3. Chapeau devenant blanchâtre pruineux en vieillissant; arête des lamelles orange ou brunâtre; éléments du pileipellis très allongés, jusque 300 mm de long......................... 38. L. medusae
Chapeau non blanchâtre pruineux; arête des lamelles concolore; éléments du pileipellis jusque 100 mm de long .................. 4
4. Stipe blanc, contrastant avec le chapeau orange .............................................................................. 5
Stipe orange pâle à jaune pâle, plus concolore avec le chapeau ................................................... 6
5. Spores allongées, en moyenne Q = 1.33–1.60; chapeau jusque 7 cm diam., trame de l’hyménophore avec des sphérocystes ......................................................... 36. L. longisporus
Spores ellipsoïdes, en moyenne Q = 1.20–1.27; chapeau < 3 cm diam., trame de l’hyménophore sans sphérocystes ................................. 37. L. pumilus
6. Marge des lamelles fertile; ornementation sporale constituée d’un réseau incomplet sans verrues distinctes ..... ....................... 34. L. gymnocarpoides
Marge des lamelles stérile; ornementation sporale constituée d’un réseau partiel et de quelques verrues isolées ....................................... 35. L. pseudogymnocarpus
Sect. Rugati Verbeken
1. Pileipellis avec des nuances orange, plutôt tomenteux, non veiné ou ridé, au plus légèrement ridé; stipe court comparé au diamètre du chapeau ..................................................................... 2
Pileipellis avec des nuances brun rougeâtre, finement mais distinctement veiné et ridé ou craquelé; stipe plus long (3–8 cm) ................................................................................................. 3
2. Lamelles blanc pur, contrastant avec l’orange ou l’orange jaunâtre du stipe et du chapeau ......... 41. L. volemoides
Lamelles orange ............................................................................................... 42. L. pseudovolemus
3. Lamelles denses, chapeau souvent craquelé .................................................. 45. L. xerampelinus
Lamelles (modérément) espacées, non craquelées .......................................................................... 4
4. Chapeau rouge orange à brun; spores 9 × 6.5–7 mm; verrues petites, surtout allongées jusque 0.1 mm de haut; éléments du stipitipellis (20–)50–100 ´ 4–6 mm, à paroi épaisse, extrêmement longs et tortueux à la base du stipe ................................................................ 43. L. rubiginosus
Chapeau brun foncé à l’état jeune, puis plus clair et plus rougeâtre, parfois avec des taches ocracées; spores plus petites, 7.9–8.4 ´ 5.9–6.3 mm, verrues plus arrondies, jusque 0.2 mm de haut; éléments du stipitipellis 10–40 ´ 2–4 mm, à paroi fine, à paroi épaisse uniquement à la base ......................... 44. L. kivuensis
Sect. Polysphaerophori Singer
1. Chapeau orange jaunâtre, orange, orange brunâtre, parfois plus ocracé en vieillissant ........... 3
Chapeau ni orange ni rouge, blanc jaunâtre, chamois, brunâtre ................................................... 2
2. Chapeau pâle, couleur crème, blanc jaunâtre, tomenteux; lamelles modérément distantes; ornementation sporale jusque 1 µm de haut, subréticulée à réticulée ............. 51. L. foetens
Chapeau chamois à brunâtre, fortement pruineux à granuleux; ornementation sporale jusque 0.2 µm de haut, consistant en verrues souvent alignées, mais ne formant jamais de réseau ........... 52. L. brunnescens
3. Eléments terminaux du pileipellis à paroi très épaisse (jusque 3 µm), souvent 10–15 mm de large; lamelles distantes, décurrentes par une dent ........................................... 46. L. gymnocarpus
Eléments terminaux du pileipellis à paroi moins épaisse, généralement <10 µm de large; lamelles le plus souvent serrées à modérément distantes (si distantes, éléments terminaux du pileipellis < 8µm de large) ...... 4
4. Stipe blanc ................................................................................................................... 49. L. tanzanicus
Stipe jaune à orange ou orange brunâtre, généralement brillant ................................................... 5
5. Ornementation sporale peu visible, au plus 0.2 µm de haut, consistant en verrues allongées non reliées; plage non amyloïde; stipe jaune à orange pâle ou orange brunâtre, à zone blanc pur distincte au sommet .... 48. L. albocinctus
Ornementation sporale jusque 0.5 µm de haut, subréticulée avec une tache amyloïde distincte au centre; stipe parfois aussi à zone blanchâtre au sommet .......................................................... 6
6. Spores ellipsoïdes, 7.3–8.4 ´ 5.5–6.0 mm; en moyenne Q = 1.20–1.50 ............... 47. L. flammans
Spores allongées, 8.2–10.3 ´ 5.2–6.3 mm; en moyenne Q = 1.40–1.75 ........ 50. L. goossensiae
Sect. Phlebonemi R. Heim ex Verbeken
1. Eléments du pileipellis capités, sphériques au sommet ........................................ 57. L. nonpiscis
Eléments du pileipellis non capités ...................................................................................................... 2
2. Chapeau zoné, ocracé brunâtre, devenant plus pâle de couleur crème ....... 56. L. pisciodorus
Chapeau jamais zoné, ne devenant jamais très pâle ........................................................................ 3
3. Pleurocystides présentes et abondantes ...................................................................... 55. L. arsenei
Pleurocystides absentes ......................................................................................................................... 4
4. Chapeau à veines sinueuses gélifiées remarquables, très fortement ridé, jaune ocre à centre orange à ocre brunâtre ou brun foncé; latex virant d’ocre à brun clair, puis brun foncé; connectifs dans l’ornementation sporale rares, quoique présents sur à peu près toutes les spores; éléments terminaux du pileipellis à paroi très épaisse (paroi jusque 1 µm d’épaisseur) ............. 53. L. phlebonemus
Chapeau sans veines, presque lisse, feutré, orange ocracé vif; latex blanc, immuable, contexte virant au brun; connectifs dans l’ornementation sporale très rares, présents seulement sur quelques spores; éléments terminaux du pileipellis à paroi légèrement épaissie (paroi jusque 0.5 µm d’épaisseur) ............ 54. L. angustus
Sect. Piperites (Fr. ex J. Kickx f.) Burl. s.l.
1. Chapeau à marge fortement poilue ............................................................................ 59. L. barbatus
Chapeau à marge lisse .................................................................. 2
2. Stipe plus clair que le chapeau, scrobiculé; goût doux à amer; ornementation sporale composée de verrues principalement isolées ....................................................... 60. L. afroscrobiculatus
Stipe concolore avec le chapeau, non scrobiculé; goût âcre brûlant; ornementation sporale composée de verrues principalement alignées .............................................. 61. L. acrissimus
Sect. Amari Verbeken
1. Spores en moyenne 9.5–10 × 8.5 µm; goût doux et variable (farineux, raphanoïde); lamelles libres à adnées, plutôt espacées .............................. 62. L. subamarus
Spores en moyenne 7.0 × 5.7 µm; goût âcre et amer; lamelles fortement décurrentes, serrées....
................................. 63. L. amarus
Sect. Nigrescentes Verbeken
1. Spores ellipsoïdes, Q = 1.25–1.7 ....................................................................... 2
Spores globuleuses à subglobuleuses, rarement ellipsoïdes, Q = 1–1.2 ..................................... 3
2. Spores entièrement ailées; chapeau et stipe jaune grisâtre à brun jaunâtre; chapeau sans papille, non veiné ... 67. L. orientalis
Spores partiellement ailées, à verrues côniques et crêtes plus basses; chapeau et stipe brun foncé; chapeau avec une papille distincte .................................................... 68. L. griseogalus
3. Ornementation sporale 1–2 mm de haut; spores 7.3–7.5 ´ 6.5–6.7 mm; latex finalement noir bleuté ............ 65. L. melanogalus
Ornementation sporale jusque 1(1.3) mm de haut; spores 7.9–8.3 ´ 7.3–7.6 mm; latex finalement chamois à crème ........................ 66. L. baliophaeus
Sect. Plinthogali (Burl.) Singer s.l.
1. Carpophore gastéroïde ou secotioïde ................................................................. 2
Carpophore avec un stipe et des lamelles normalement développés ............................................ 3
2. Carpophore gastéroïde, jaune ocracé, hypogé ou semi-hypogé; spores subglobuleuses, 8–11 µm de long; ornementation à ailes prononcées, jusque 3(3.5) µm, formant un réseau presque complet .......... 69. L. angiocarpus
Carpophore épigé, à stipe normalement développé; hyménophore sinueux, rappelant une gleba ou alvéolé labyrinthoïde ..................... 70. L. dolichocaulis
3. Ornementation sporale zébroïde .............................................................. 4
Ornementation sporale en réseau ou épineuse......................................................... 9
4. Pleurocystides abondantes ...................................................................... 5
Pleurocystides absentes ......................................................... 6
5. Jeune chapeau brun clair à orange grisâtre; contexte ne virant pas au rougeâtre ni au rosâtre; pleurocystides émergentes, à contenu à l’aspect d’aiguilles; pileocystides abondantes ....... 71. L. pulchrispermus
Jeune chapeau brun foncé; contexte virant au rouge, rouge orange au froissement; pileocystides absentes ................................. 72. L. miniatescens
6. Spores ellipsoïdes ornementées, à crêtes arrondies régulières, jusque 1 µm de haut; chapeau brun grisâtre à brun jaunâtre, 10–45 mm diam.; stipe creux et fragile ........ 73. L. sulcatulus
Spores ellipsoïdes à globuleuses, à ornementation jusque 2 µm de haut .................................... 7
7. Chapeau > 4 cm diam., brun fuligineux; ornementation sporale consistant en crêtes grossières et parallèles .................................. 76. L. kalospermus
Chapeau petit, < 2 cm diam., à teintes jaunâtres à brun clair; spores à aspect épineux ............. 8
8. Chapeau à teintes brunes; stipe à base distincte hirsute; spores 8–9.5 × 7–8.5 µm, partiellement réticulée .... 74. L. adhaerens
Chapeau jaune pâle à jaune vif; base du stipe non hirsute; spores 7–8 × 5.5–6.5 µm, non réticulées ........ 75. L. desideratus
9. Contexte virant au rosâtre ou au rougeâtre au froissement; ornementation sporale ailée ....... 10
Contexte ne virant pas au rose ou au rougeâtre; ornementation sporale ailée ou épineuse... 12
10. Pleurocystides présentes; chapeau brun olivacé ........................................................ 79. L. nudus
Pleurocystides absentes; chapeau fuligineux brun à brun foncé ............................................... 11
11. Arête des lamelles brunâtre; ornementation sporale 1(1.5) µm ..................... 77. L. congolensis
Arête des lamelles concolore; spores très fortement ornementées, à ailes réticulées jusque 2 µm de haut, distinctement verruqueuses entre les crêtes............................. 78. L. melanodermus
12. Lamelles distantes ....................................................................... 13
Lamelles (modérément) serrées ................................................. 14
13. Chapeau grisâtre clair à gris brunâtre, marge non sulquée ......................... 80. L. rumongensis
Chapeau brun jaunâtre plus foncé à brun foncé, marge sulquée ........................ 81. L. sulcatus
14. Arête des lamelles colorée (brun) ...................................................... 15
Arête concolore avec les lamelles ................................................ 16
15. Spores 7.7–8.1 ´ 7.0–7.5 mm; pileipellis en hyméniderme, éléments terminaux cylindriques à clavés, jusque 12 mm de large ...................... 82. L. saponaceus
Spores 5.8–6.5 ´ 5.1–5.9 mm; pileipellis en trichoderme ou trichopalissade, éléments terminaux long et minces, jusque 5 mm de large .................... 83. L. pusillisporus
16. Chapeau couvert de veines radiales épaisses; stipe avec un large collier de poils orange autour de la base; ornementation sporale épineuse(-zébroïde); basides en partie 2-spores; pleuropseudocystides ramifiées et tortueuses ................................................ 74. L. adhaerens
Pas cette combinaison de caractères; ornementation sporale moins haute, épineuse ou réticulée; partie terminale des pleuropseudocystides cylindrique et large; contexte souvent orange à la base du stipe ...... (Pseudolignyoti ad int.) 17
17. Ornementation sporale jusque 1 µm de haut, réticulée; espèces des forêts claires ................ 18
Ornementation sporale jusque 1.5 mm de haut, épineuse; petits carpophores; espèces des forêts ombrophiles
............................................................................................................................................................ 20
18. Chapeau jusque 4 cm diam., brun foncé à l’état jeune, vite grisâtre plus pâle, marge plus ou moins crénelée; jeunes lamelles blanches contrastant avec le chapeau et le stipe (rappelant L. lignyotus); contexte de la base du stipe jamais orange; éléments terminaux du pileipellis cylindriques, longs et minces, 10–40 ´ 6–10 mm ........... 86. L. tenellus
Chapeau plus grand, restant brun à brun foncé; marge non crénelée; contexte de la base du stipe orange; lamelles rapidement orange ocracé .................................................................... 19
19. Basidiocarpes jusque 5 cm de haut; chapeau jusque 7 cm diam.; éléments terminaux du pileipellis obovoïdes à largement clavés, 10–20 ´ 8–15 mm; espèce commune du miombo, poussant dans la litière peu épaisse des clairières ......................................... 84. L. kabansus
Basidiocarpes jusque 9 cm de haut; chapeau jusque 12 cm diam.; éléments terminaux du pileipellis subcylindriques, subfusiformes, jamais arrondis ni clavés, 30–65 × 8–10(12) µm; espèce du miombo préférant l’ombre et la litière épaisse .............................. 85. L. sciaphilus
20. Chapeau de couleur crème à squamules brun ocracé foncé; chapeau ne portant jamais de papille ... 88. L. pseudolignyotus
Chapeau brun foncé, finement tomenteux; chapeau portant parfois une papille ......... 87. L. acutus
Sect. Pseudofuliginosi Verbeken
1. Pleuromacrocystides abondantes; chapeau à teintes olivacées noirâtres; spores ornementées, à fines crêtes étroites .................... 89. L. atro-olivinus
Pleuromacrocystides absentes; chapeau plutôt brunâtre; spores plus lourdement ornementées ......... 2
2. Eléments ascendants dans le pileipellis 5–9 mm de large, subclavés ou subfusiformes; chapeau mat et très finement feutré .................................................... 90.1. L. undulatus var. undulatus
Eléments ascendants dans le pileipellis étroits (2–4 µm), cylindriques, plus rares; chapeau lisse et brillant, légèrement visqueux .................................................. 90.2. L. undulatus var. rasilis
Basso M. T. (1999) Lactarius Pers. Fungi Europei 8. Mycoflora, Alassio.
Beeli M. (1927a) Contribution à l'étude de la flore mycologique du Congo. Fungi Goossensiani II. Bull. Soc. Roy. Bot. Belg. 59: 101‑112, Pl. 1‑2.
Beeli M. (1927b) Contribution à l'étude de la flore mycologique du Congo. Fungi Goossensiani IV. Bull. Soc. Roy. Bot. Belg. 60: 75‑87, Pl. 1‑2.
Beeli M. (1928) Contribution à l'étude de la flore mycologique de Congo. Fungi Goossensiani V. Bull. Soc. Roy. Bot. Belg. 60: 153‑174.
Beeli M. (1933) Contribution à l'étude de la flore mycologique de Congo. Fungi Goossensiani X. Bull. Soc. Roy. Bot. Belg. 66: 14‑31.
Beeli M. (1936) Contribution à l'étude de la flore mycologique du Congo. Fungi Goossensiani XI. Bull. Jard. Bot. Etat Bxl 14: 83‑91.
Berthet P. & Boidin J. (1966) Observations sur quelques hyménomycètes récoltés en République Camerounaise. Cah. Maboké 4: 27‑54, Pl. 4.
Bresadola J. & Saccardo P.A. (1899) Fungi congoensis. In Durand Th. & De Wildeman E., Matériaux pour la flore du Congo. Bull. Soc. Roy. Bot. Belg. 38: 152‑168.
Buyck B. (1988) Russules nouvelles d'Afrique centrale. Bull. Jard. Bot. Nat. Belg. 58: 467‑476.
Buyck B. (1989a) New taxa of Central African Russulaceae. Bull. Jard. Bot. Nat. Belg. 59: 241‑253.
Buyck B. (1989b) Révision du genre Russula en Afrique centrale. Ph.D. dissertation, Rijksuniversiteit Gent, part I: 318 pp., 98 fig.; part II: 590 pp., 582 fig.; part III: 444 photos SEM (unpubl.).
Buyck B. (1991) The study of microscopic features in Russula. 1. Spores and basidia. Russulales Newsl. 1: 8-26.
Buyck B. (1993) Russula I. Fl. Illustrée Champignons Afrique Centrale 15: 335-408, Pl. 55-68.
Buyck B. (1994) Ubwoba: les champignons comestibles de l'Ouest du Burundi. Bruxelles, Admin. Gén. Coop. Dévelopm., Publ. Agric. 34: 123 p.
Buyck B. & Nzigidahera B. "1995" (1996) Ethnomycological notes from Western Burundi. Belg. J. Bot. 128: 131‑138.
Buyck B. & Verbeken A. (1995) Studies in tropical African Lactarius species. 2. Lactarius chromospermus Pegler. Mycotaxon 56: 427‑442.
Buyck B., Verbeken A. & Eberhardt U. (2007) The genus Lactarius in Madagascar. Mycol. Res. 111: 787-798.
Buyck B., Hofstetter V., Eberhardt U., Verbeken A. and Kauff, F. (2008) Walking the thin line between Russula and Lactarius: the dilemma of Russula subsect. Ochricompactae. Fungal Diversity 28: 15-40.
Chipompha N.W.S. (1994) Some mushrooms of Malawi: agarics of Central and Northern Malawi. In Seyani J.H. & Chikuni A.C. (eds.) Proceedings XIIIth Plenary Meeting AETFAT congress, Malawi, 2‑11 April 1991, vol. 1. Zomba (Malawi), p. 649‑658.
Clémençon H. (1972) Zwei verbesserte Präparierlösungen für die microskopische Untersuchung von Pilze. Z. Pilzk. 38: 49-53.
Dade H.A. (1940) A revised list of Gold Coast fungi and plant diseases. Bull. Misc. Inform. Kew 1940: 205‑247.
Degreef J. (1990) Inventaire, valeur alimentaire et culture de champignons comestibles du Shaba. Thesis (unpubl.): 103 p. Univ. Gembloux (Belgium), Fac. Agric. Sci.
De Kesel A., Codjia J.T.C. & Yorou N.S. (2002) Guide des champignons comestibles du Bénin. Jardin botanique national de Belgique, Meise (Belgium) & Cotonou (Benin), CECODI, 274 p.
De Wildeman E. (1914) Additions à la flore du Congo. Bull. Jard. Bot. Etat Bxl. 4: 1‑241.
De Wildeman E. & Durand T. (1901) Reliquiae Dewevreanae. Enumération des plantes récoltées par A. Dewèvre en 1895‑1896. Ann. Mus. Congo, Sér. Bot. 3, 1(1): 269‑287.
Donk M.A. (1949) New and revised nomina generica conservanda proposed for Basidiomycetes (Fungi). Bull. Bot. Jard. Buitenzorg, Ser. III, 18: 83-168.
Earle F.S. (1909) The genera of North American gill fungi. Bull. New York Bot. Gard. 5: 373-451.
Eberhardt E. & Verbeken A. (2004) Sequestrate Lactarius species from tropical Africa: L. angiocarpus sp. nov. and L. dolichocaulis comb. nov. Mycol. Res. 108: 1042-1052.
Eichelbaum F. (1906) Beiträge zur Kenntnis der Pilzflora des Ostusambaragebirges. Verh. Naturwiss. Vereins Hamburg 14: 1‑92.
Härkönen M., Saarimäki T. & Mwasumbi L. (1995) Edible mushrooms of Tanzania. Karstenia 35, Suppl.: 1‑92.
Härkönen M., Niemelä T. & Mwasumbi L. (2003) Tanzanian mushrooms. Edible, harmful and other fungi. Norrlinia 10: 1-200.
Heilmann-Clausen J., Verbeken A. & Vesterholt J. (1998) The genus Lactarius. Fungi of Northern Europe 2. Svampetryk, Danish Mycological Society, Denmark.
Heim R. (1937) Observations sur la flore mycologique malgache V. Les Lactario-Russulés à anneau: ontogénie et phylogénie. Rev. Mycol. (Paris) 2: 4-17, 61-75, 109-117.
Heim R. (1938a) Diagnoses latines d'espèces et variétés nouvelles de Lactario‑russulés du domaine oriental de Madagascar. Candollea 7: 374‑393.
Heim R. 1937 (1938b) Les lactario‑russulés du domaine oriental de Madagascar. Prodr. Fl. Mycol. Madagascar Dépendances 1: 1‑196, Pl. 1‑8.
Heim R. (1943) Remarques sur les formes primitives ou dégradées de Lactario‑Russulés tropicaux. Boissiera 7: 266‑280.
Heim R. (1955a) Les lactaires d'Afrique intertropicale (Congo belge et Afrique noire française). Bull. Jard. Bot. Etat Bxl 25: 1‑91, Pl. 1‑6.
Heim R. (1955b) Lactarius. Fl. Icon. Champ. Congo 4: 81‑97, Pl. 13‑15.
Heim R. (1966) Sur un lactaire hygroporoide tropical à lait grisonnant Lactarius griseogalus Heim sp. nov. Israel J. Bot. 15: 158‑162.
Heim R. (1967) Breves diagnoses latinae novitatum genericarum specificarumque nuper descriptarum. Rev. Mycol. (Paris) 32: 201‑210.
Hennings P. “1901” (1902) Fungi camerunenses novi III. Bot. Jahrb. Syst. 30: 39‑57.
Hesler L.R. & Smith A.H. (1979) North American species of Lactarius. Michigan Press, Ann Arbor, 841 pp., 154 pl.
Holmgren P.K. & Holmgren N.H. (1998). Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden's Virtual Herbarium. [Continuously updated at http://sweetgum.nybg.org/ih/]
Homola R.L. & Kimball J. (1975) Scanning Electron Microscopy of spore ornamentation in the genus Lactarius. Michigan Bot. 14(4): 179-189.
Horak E. (1968) Synopsis generum agaricalium (Die Gattungstypen der Agaricales). Beitr. Kryptogamenfl. Schweiz. 13: 1‑741.
Josserand M. (1941) Etude sur l'ornamentation sporique des Lactaires et de quelques autres espèces à spores amyloides. Bull. Trimestriel Soc. Mycol. France 56: 7-38.
Karhula P. (1997) Tansanian rouskut ja niiden käyttö pro gradu – Tutkielma. Master thesis, Helsinki University.
Karhula P., Härkönen M., Saarimäki T., Verbeken A. & Mwasumbi L. (1998) Tanzanian mushrooms and their uses 6. Lactarius. Karstenia 38: 49-68.
Kirk P.M. & Ansell A.E. (1992) Authors of fungal names. International Mycological Institute, Kew. [Continously updated at: http://www.indexfungorum.org/Names/AuthorsOfFungalNames.asp]
Kornerup A. & Wanscher J.H. (1978) Methuen Handbook of Colour. Methuen, London.
Lalli G. & Pacioni G. (1992) Lactarius sect. Lactifluus and allied species. Mycotaxon 44: 155‑195.
Lebel T. & Tonkin J. (2007) Australasian species of Macowanites are sequestrate species of Russula (Russulaceae, Basidiomycota). Austr. Syst. Bot. 20: 355-381.
Maghembe J.A. & Redhead J.F. (1980) A survey of ectomycorrhizal fungi in pine plantations in Tanzania. E. African Agric. For. J. 45: 203‑206.
Malençon G. (1931) Considérations sur les spores des Russules et des Lactaires. Bull. Trimestriel Soc. Mycol. France 47: 72-86.
Montoya L., Bandala V.M. & Mata M. (2007) Studies on Lactarius: two new records from Costa Rica and additional information from Mexico. Mycotaxon 99: 279-290.
Morris B. (1984) Macrofungi of Malawi. Some ethnobotanical notes. Bull. Brit. Mycol. Soc. 18: 48‑57.
Morris B. (1987) Common mushrooms of Malawi: 108 p., 8 pl. Oslo, Fungiflora.
Morris B. (1990) An annotated check‑list of the macrofungi of Malawi. Kirkia 13: 323‑364.
Munyanziza E. (1994) Miombo trees and mycorrhizae. Ecological strategies, a bases for afforestation: 192 p. PhD Thesis., Univ. Wageningen (The Netherlands).
Nuytinck J. Verbeken A., Delarue S. & Walleyn R. (2003) Systematics of European sequestrate lactarioid Russulaceae with spiny spore ornamentation. Belg. J. Bot. 136: 145-153.
Nzigidahera B. (1993) Contribution à l'étude systématique et écologique des champignons ectomycorrhiziques de la forêt claire de Nkayamba. Thesis (unpubl.), Univ. Burundi, Fac. Sci., 146 p.
Onguene N.A. (2000) Diversity and dynamics of mycorrhizal associations in tropical rain forests with different disturbance regimes in South Cameroon: 167 p. PhD Thesis., Univ. Wageningen (Netherlands).
Pegler D.N. (1969) Studies on African Agaricales. II. Kew Bull. 23: 219‑249.
Pegler D.N. (1977) A preliminary agaric flora of East Africa. Kew Bull., Addit. Ser. 6: 1‑615.
Pegler D.N. (1982) Agaricoid and boletoid fungi (Basidiomycota) from Malawi and Zambia. Kew Bull. 37: 254‑271.
Pegler D.N. (1983) The genus Lentinus, a world monograph. Kew Bull., Addit. Ser. 10: 1‑281.
Pegler D.N. & Piearce G.D. (1980) The edible mushrooms of Zambia. Kew Bull. 35: 475‑491.
Pegler D.N. & Young T.W.K. (1982) Russulaceae Roze. World Pollen Spore Fl. 10: 1-35.
Piearce G.D. (1981) An introduction to Zambia's wild mushrooms, and how to use them: 28 p. Zambia, Forest Department.
Redeuilh G., Verbeken A. & Walleyn R. (2001) Étude nomenclaturale des taxons infragénériques dans le genre Lactarius (Basidiomycota, Russulaceae). Mycotaxon 77: 127-143. [Continously updated at: http://www.mtsn.tn.it/russulales-news/]
Reijnders A.F.M. (1963) Les problèmes du développement des carpophores des Agaricales et de quelques groupes voisins. Dr. W. Junk, Den Haag, 412 p., 55 pl.
Romagnesi H. (1967) Les Russules d’Europe et d’Afrique du Nord. Paris, Bordas, 998 p. + 1 pl.
Ryvarden L., Piearce G.D. & Masuka A.J. (1994) An introduction to the larger fungi of South Central Africa: 200 pp. Harare (Zimbabwe), Baobab books.
Sanon K.B., Bâ A.M. & Dexheimer J. (1997) Mycorrhizal status of some fungi fruiting beneath indigenous trees in Burkina Faso. Forest Ecol. Management 98: 61-69.
Singer R. (1942) Das System der Agaricales. II. Ann. Mycol. 40: 1‑132.
Singer R. (1946) New and interesting species of basidiomycetes. II. Pap. Michigan Acad. Sci. 32: 103‑150, Pl. 1.
Singer R., Araujo I. & Ivory M.H. (1983) The ectotrophically mycorrhizal fungi of the neotropical lowlands, especially central Amazonia. Beih. Nov. Hedw. 77: 1‑352.
Thoen D. & Ba A.M. (1989) Ectomycorrhizas and putative ectomycorrhizal fungi of Afzelia africana Sm. and Uapaca guineensis Müll. Arg. in southern Senegal. New Phytol. 113: 549‑559, 2 pl.
Thoen D. & Ducousso M. (1989) Champignons et ectomycorrhizes du Fouta Djalon. Bois Forêts Trop. 221: 45‑63.
To A. & Ob A. (2005) Proximate and mineral composition of four edible mushroom species from South Western Nigeria. African J. Biotechnology 4(10): 1084-1088.
Van der Westhuizen G.C.A. & Eicker A. (1994) Field guide to the mushrooms of southern Africa: 207 p. Cape town, Struik Publishers.
Van Rooij P., De Kesel A. & Verbeken A. (2003) Studies in tropical African Lactairus species (Russulales, Basidiomycota) 11. Records from Benin. Nova Hedwigia 77: 221-251.
Verbeken A. (1995a) Further notes on Lactarius edulis Verbeken & Buyck. Russulales News 3: 17‑22.
Verbeken A. (1995b) Studies in tropical African Lactarius species. 1. Lactarius gymnocarpus Singer ex R. Heim and allied species. Mycotaxon 55: 515‑542.
Verbeken A. (1996a) Studies in tropical African Lactarius species. 4. Species described by P. Hennings and M. Beeli. Edinburgh J. Bot. 52: 49‑79.
Verbeken A. (1996b) Studies in tropical African Lactarius species. 3. Lactarius melanogalus and related species. Persoonia 16: 209‑223.
Verbeken A. (1996c) New taxa of Lactarius (Russulaceae) in tropical Africa. Bull. Jard. Bot. Nat. Belg. 65: 197‑213.
Verbeken, A. (1996d) Biodiversity of the genus Lactarius Pers. in tropical Africa. PhD thesis, Ghent University (Belgium).
Verbeken A. (1998a) Studies in tropical African Lactarius species. 5. A synopsis of the subgenus Lactifluus (Burl.) Hesler & A.H. Sm. emend. Mycotaxon 66: 363-386.
Verbeken A. (1998b) Studies in tropical African Lactarius species. 6. A synopsis of the subgenus Lactariopsis (Henn.) R. Heim emend. Mycotaxon 66: 387-418.
Verbeken A. “2000” (2001a) Studies in tropical African Lactarius species. 8. A synopsis of the subgenus Plinthogali. Persoonia 17: 377, Pl. 1-8.
Verbeken A. (2001b) Worldwide systematics of Lactarius: a state of the art. Micol. Vegetat. Mediterranea 16: 71-88.
Verbeken A. & Buyck B. (2001) Diversity and ecology of tropical ectomycorrhizal fungi in Africa. In Watling R., Frankland J.C., Ainsworth A.M., Isaac S. & Robinson C. (eds.) Tropical Mycology, Vol. 1. Wallingford, CABI Publishing, p. 11-24.
Verbeken A., Buyck B. & Walleyn R. (1996) Lentinus clitocyboides Hennings is a Russula. Cryptog. Mycol. 17: 7‑10.
Verbeken A. & Sharp C. (2003) A new Lactarius species from Zimbabwe. Cryptog. Mycol. 24: 125-129.
Verbeken A., Stubbe D. & Nuytinck J. (2008) Two new Lactarius species from Cameroon. Cryptog. Mycol. 29: 137-143.
Verbeken A. & Walleyn R. “1999” (2000) Studies in tropical African Lactarius species. 7. A synopsis of the section Edules and a review on the edible species. Belg. J. Bot. 66: 175-184.
Verbeken A., Walleyn R., Sharp C. & Buyck B. (2000) Studies in tropical African Lactarius species. 9. Records from Zimbabwe. Syst. Geogr. Pl. 70: 181-215.
Walker A. (1931) Champignons comestibles de la Basse‑Ngounié (Gabon). Rev. Int. Bot. Appl. Agric. Trop. 11: 240‑247.
Williamson J. (1975) Useful plants of Malawi (revised and extended edition). Fungi: 312‑336. Zomba, University of Malawi.
Williamson J. (1976) A list of some fungi collected in Malawi between 1971 and 1974. Soc. Malawi J. 29: 46‑53.
Yorou S.N., De Kesel A., Sinsin B. & Codjia J.T.C. (2002). Diversité et productivité des champignons comestibles de la forêt classée de Wari Maro (Bénin). In E. Robbrecht, J. Degreef & I. Friis (eds) Plant Systematics and Phytogeography for the understanding of African Biodiversity. Proceedings of the XVIth AETFAT Congress (2000, Meise, National Botanic Garden of Belgium). Syst. Geogr. Pl. 71:613-625.
Yorou S.N. & De Kesel A. (2002). Connaissances ethnomycologiques des peuples Nagot du centre du Bénin (Afrique de l'Ouest). In E. Robbrecht, J. Degreef & I. Friis (eds) Plant Systematics and Phytogeography for the understanding of African Biodiversity. Proceedings of the XVIth AETFAT Congress (2000, Meise, National Botanic Garden of Belgium). Syst. Geogr. Pl. 71:627-637.
Zeller H. (1982) Les champignons principaux de Madagascar (Malgaches): 34 p. Antananarivo, Institut Pasteur de Madagascar.
Accepted names are bold, synonyms are in normal font.
Afrobarbati (sect.)
Amari (sect.)
Armillaria pelliculata
Aurantiifolii (sect.)
Chamaeleontini (sect.)
Chromospermi (sect.)
Clitocybe castanea
Edules (sect.)
Laccaria kalosperma
Lactariopsis (sect.)
Lactariopsis zenkeri
Lactarius acrissimus
Lactarius acutus
Lactarius adhaerens
Lactarius afroscrobiculatus
Lactarius albocinctus
Lactarius amarus
Lactarius angiocarpus
Lactarius angustus
Lactarius annulatoangustifolius
Lactarius arsenei
Lactarius atro-olivinus
Lactarius aurantiifolius
Lactarius aureifolius
Lactarius badius
Lactarius baliophaeus
Lactarius baliophaeus var. orientalis
Lactarius barbatus
Lactarius brachystegiae
Lactarius brunnescens
Lactarius caperatus
Lactarius carmineus
Lactarius chamaeleontinus
Lactarius chromospermus
Lactarius cinnamomeus
Lactarius claricolor
Lactarius congolensis
Lactarius corbula
Lactarius craterelloides
Lactarius cyanovirescens
Lactarius denigricans
Lactarius densifolius
Lactarius desideratus
Lactarius dolichocaulis
Lactarius edulis
Lactarius emergens
Lactarius flammans
Lactarius foetens
Lactarius fulgens
Lactarius fulgens var. africanus
Lactarius goossensiae
Lactarius griseogalus
Lactarius gymnocarpoides
Lactarius gymnocarpus
Lactarius heimii
Lactarius hepaticus
Lactarius hispidulus
Lactarius indusiatus
Lactarius inversus
Lactarius kabansus
Lactarius kabansus var. pallidus
Lactarius kalospermus
Lactarius kivuensis
Lactarius laevigatus
Lactarius latifolius
Lactarius lignyotus
Lactarius longipes
Lactarius longisporus
Lactarius luteopus
Lactarius madagascariensis
Lactarius medusae
Lactarius melanodermus
Lactarius melanogalus
Lactarius miniatescens
Lactarius nodosicystidiosus
Lactarius nonpiscis
Lactarius nudus
Lactarius orientalis
Lactarius pandani
Lactarius pandani f. aurantiacus
Lactarius pandani f. pallidus
Lactarius pandani f. intermedius
Lactarius pelliculatus
Lactarius pellucidus
Lactarius phlebonemus
Lactarius phlebophyllus
Lactarius piperatus
Lactarius pisciodorus
Lactarius pruinatus
Lactarius pseudogymnocarpus
Lactarius pseudolignyotus
Lactarius pseudotorminosus
Lactarius pseudovolemus
Lactarius pulchrispermus
Lactarius pumilus
Lactarius pusillisporus
Lactarius roseolus
Lactarius rubiginosus
Lactarius rubroviolascens
Lactarius rufomarginatus
Lactarius rumongensis
Lactarius russuliformis
Lactarius ruvubuensis
Lactarius saponaceus
Lactarius sciaphilus
Lactarius sesemotani
Lactarius striatus
Lactarius subamarus
Lactarius sulcatulus
Lactarius sulcatus
Lactarius tanzanicus
Lactarius tenellus
Lactarius trivialis
Lactarius uapacae
Lactarius undulatus
Lactarius undulatus var. rasilis
Lactarius undulatus var. undulatus
Lactarius unicolor
Lactarius urens
Lactarius uvidus
Lactarius vellereus
Lactarius velutinus
Lactarius velutissimus
Lactarius volemoides
Lactarius xerampelinus
Lactarius zenkeri
Lentinus clitocyboides
Omphalia russuliformis
Phlebonemi (sect.)
Piperites (sect.)
Plinthogali (sect.)
Polysphaerophori (sect.)
Pseudofuliginosi (sect.)
Pseudogymnocarpi (sect.)
Rubroviolascentini (sect.)
Rugati (sect.)
Russula annulatoangustifolia
Russula clitocyboides
Russula sesemotani
Russula subfistulosa var. apsila
Russularia (sect.)
Russulopsidei (sect.)