ARTICLE 25

et et like and et al et al et al et al. RPB. et al./// Cantharellus , has recently Key words: Africa F&(V Cantharellus "&J LSU Molecular phylogeny & Botryobasidium et al677W( et al. .//G ?# Craterellus et al .//V [ Cantharellus cibarius-

is et al.///P ! # ! IMA FUNGUS · VOLUME 3 · NO 1: 25–38 NO 1: 3 · · VOLUME FUNGUS IMA Cantharellus et al .//G ( et al.677WP Afrocantharellus Cantharellaceae and the closely related et al. .//V " with different datasets, whereas LSU still seems to with different ( +JM = et al6777N Craterellus consistently were monophyletic and sister- 1 branch attraction in some types of analyses, particularly in = ?# primarily had a background in incongruent placement of Tulasnella # Tulasnella chanterelles has been demonstrated, suggesting a species < ! al 6772 N & # 9 P .//. ?# et al .//V genes such as RPB.# = 6772"&J# et al.//GF#"&J 0F 6777+ al.//7" N '+N0F+JM "&J'# of or in Cantharellales 6772! been investigated using molecular data, and reviewed by been investigated using molecular data, Moncalvo genes, particularly relationships as reconstructed from different with respect to the placement of Tulasnella and mtSSU, and groups in analyses based on LSU, SSU, Cantharellus

as from et al

J< FKK# Cantharellus Cantharellus . ( species from the Afrocantharellus , was described, and species known from and C. symoensii +& . et al./66 Z et al..///0 species are widely collected Afrocantharellus , Cantharellus C. splendens, J†J#4 Cantharellus, as presently delineated, C. platyphyllus 6.­

Cantharellus Cantharellus based on LSU and a concatenated data set are provided, along with descriptions of and a based on LSU and a concatenated data set et al.//7J from Africa, including detailed investigations of from %9.//6 et al.//Q9P./66 A new genus in the Cantharellaceae new genus in the A

Afrocantharellus NJ"[MM#0#6QNWR.GVMJ N?M#NJ([<GR6W7NJ&( ./66% Cantharellus %9 includes several well-known and highly esteemed edible includes several well-known and highly "F X=1 Cantharellus ### 6772 others are mentioned in a list of Cantharellus 90677R! 677QF F # F % 0 & E # P 67RQ F#F Abstract: N0F+JM+JM3RQ="&J.3F&(V"# # ' ' # F In contrast to most of and key to the four species and one form of Afrocantharellus Article info: J1.R!./6.XF16V?./6.X(1.6>./6. © 2012 International Mycological Association © 2012 International Mycological conditions: distribute and transmit the work, under the following are free to share - to copy, You Attribution: Molecular studies of the ‘cantharelloid clade’ & Q R Katanga, describing &# Cantharellaceae INTRODUCTION .G0F#J P 67VV . 1 NN& gen. stat. nov. is part of a rich diversity of African African of a rich diversity is part of stat. nov. gen. Afrocantharellus Cantharellaceae Non-commercial:No derivative works:Any of the above conditions can be waived if you get For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Nothing in this license impairs or restricts the author’s moral rights. permission from the copyright holder. "# notes and descriptions of new subgenera and sections in Cantharellus which in most cases have a vase-shaped or funnel-shaped which in most cases have a vase-shaped wrinkled, veined basidiome and a spore-bearing smooth, T 2V F 67 F % .//G VOLUME 3 · NO. 1 16/RR7Q3./6./G/6/2 "#$%4"*%

.//VF < Cantharellus cibarius, considered to &# Q#R?#et family level, but are manageable and cause much less data al.//VCantharellus and this complicates ?#et al.//V the circumscription of Cantharellus s. str. Additionally, Buyck Although LSU- and mtSSU-based analyses previously 9P./66

ARTICLE # # < C. relationships in Cantharellaceae ?# et al. .//V cibarius F&(V P# F&(=V # phylogenetic inference in Cantharellaceae K 9 species are hidden under traditional phenotype-based species 6777# Eet al.//QP# in Boletales = ; of recent studies of basidiomycetes based on molecular data < # " for the protein coding gene tef-1, leading to the recognition < 0 F Cantharellus et @et al.//V!=#et al ./66 al.//W0et al./6.& in the Cantharellus cibarius < on molecular data should be adopted when a morphological MJF 9 P ./66 9 P .//Q species concept is inapplicable in the sense that it is not presented preliminary results of a four gene phylogeny for 0 Cantharellus, employing mtSSU, LSU, and two protein- =QR coding loci, tef-1 and RPB.ca.2R P.//V# continents were sampled suggesting the recognition of at least between molecular phylogenies and morphological features < P# evaluated a posteriori< stated that more studies on a larger data set were needed & <F# understanding and reassessment of the phylogeny of ##Q Cantharellus based on the inclusion of molecular data derived RPet al.677W.///(et al.6777P9 from the rich diversity of African Cantharellus species based N.//69P.//.+et al..//2 +JMRQ="&J.F&(V' Binder et al.//R?.//R?#et al..//V Cantharellus ! et al. 677W N et al. ./// Dunham et al..//G&9P.//2Pet al. MATERIALS AND METHODS .//R†'F #E# Taxon and sequence sampling of the genus in Africa, this might well have hampered our All Cantharellus understanding of the phylogeny of Cantharellus and the & QR !6FM>JMN & # #.//2M.//WJ the molecular analyses so far published of phylogenetic # relationships of Cantharellus s. lat.< E&F##V/ ' & # E? <#0P <&et al.//Q& rich diversity of Cantharellus& ## 2/ diversity of CantharellusF< #E&F and the inclusion of data on African Cantharellus may thus #6/]' <# E& in comprehensiveness and phylogenetic relationships in 1=JNMAVM <=JN<‹5# the measured <5#AV5 JN #X ‹ 3 Current species recognition in Cantharellus %et al..//6&et al.//QJ In Cantharellus, as currently circumscribed, the distinction 67V7 < ! RQJM"&J. F&(V given the few and variable morphological characters ' .6 ./ Cantharellus #9P./66 respectively, +JM GV !<C. cibariusQCcibarius’ Craterellus " WW ' refers to any yellowish chanterelle, and C. cibarius is no @ # '&6 When the status of nominal species and morphological '@& variability within the species was not clear, sometimes these the phylogenetic position of African Cantharellus species as Q R &

26 IMA FUNGUS Afrocantharellus gen. nov. ARTICLE

X

X

Fig. 1. Map showing the distribution ?= & F< Q\R

16+JMX. F&(V +JM3RQ="&J.3F&(V F" 9' & +JM 7. set for Clavulina ' @ < Cantharellus and related genera selected for this +JM RQ="&J. Cl. cinerea with Clavulina J' @ F&(VXDacrymyces chrysospermus# ' & " # ! .MG # &FJ% QR 13333333&.1J6.W/7 by the results presented by Moncalvo et al..//V" LSU sampling, representatives of Craterellus, Hydnum, and Molecular study Multiclavula were included representing more remote relatives of CantharellusMulticlavula mucida 9' A concatenated data set included &N0F< +JM3RQ="&J.3F&(V .Q ' preferentially from the hymenium to avoid contamination, 6W L (@N0F< # X # K B"[@%0 N 6/-1M6/=G N0F < (E;&RR+JMRQ= 'RQ="&J.F&(V&F&(V "&J. F&(V ( 1 " F&(V RR+JM1+;G+;RB9P677/ however, no Craterellus ' # +;[; 1333 0PCantharellus species only two, vizC. 33µ+ ;0F .R=.QJ cibarius, and C. cinnabarinus were included. Considering that '+E61 C. cibarius' RRM@&EE@F@&&@&F@F&@F@MGRX '#@ RQJ="&J."&J&.XF&(V1 & F&(V=.F&(V=GK96777 ?#F&(V' !(E;+JMRQ="&J. #0PCantharellus species and F&(VF(®(E;(?< two Craterellus N† K G 3+ N0F " 6R3+6/3?#

VOLUME 3 · NO. 1 27 "#$%4"*%

Table 1. J'##@ 'X##[@"N ' No Species Voucher Locality Collection no. (UPS) LSU-GB 5.8-ITS2 GB ATP6-GB 1 <' NN&G6 &F0TF0"F1K &G6.//V JQ976959 ——

ARTICLE 2 < NN&2G &F0TF0"F1K &2G.//W JQ976965 —— 3 A. platyphyllus NN&VG &F0TF0"F1? &6/VG.//W JQ976970 —— cyanescens 4 A. platyphyllus NN&WQ &F0TF0"F1" &6/WQ.//W JQ976978 JQ976947 JQ976926 platyphyllus 5 A. platyphyllus NN&/G &F0TF0"F1? &6//G.//2 JQ976950 JQ976929 — platyphyllus 6 A. platyphyllus NN&26 &F0TF0"F1K &6/26.//V JQ976964 —— platyphyllus 7 A. splendens NN&RW &F0TF0"F1? &6/RW.//W JQ976967 JQ976937 JQ976916 8 A. splendens NN&6W &F0TF0"F1@ &6/6W.//R JQ976956 JQ976932 JQ976911 9 A. symoensii NN&GV &F0TF0"F1K &6/GV.//R JQ976961 JQ976934 JQ976914 10 A. symoensii NN&/2 &F0TF0"F1? &6//2.//R JQ976951 —— 11 A. symoensii NN&VV &F0TF0"F1" &6/VV.//W JQ976971 JQ976940 JQ976919 12 A. symoensii NN&66 &F0TF0"F1? &6/66.//R JQ976953 —— 13 A. symoensii NN&VW &F0TF0"F1" &6/VW.//W JQ976972 JQ976941 JQ976920 14 A. symoensii NN&62 &F0TF0"F1@ &6/62.//2 JQ976955 —— 15 Botryobasidium isabellinum F!G7G/2W — N‹RG2R7W6 16 C. appalachiensis N‹Q7QV7/ — — 17 C. appalachiensis P?WR/76V — — 18 C. cascadensis F/266R7 — — 19 C. cascadensis F/266RQ — — 20 C. cascadensis F/266V6 — — 21 C. cascadensis F/266V/ — — 22 C. cibarius # cibarius F/266RV — — 23 C. cibarius # cibarius F/266RR — — 24 C. cibarius # cibarius F/266RW — — 25 C. cibarius #roseocanus F/266R. — — 26 C. cibarius #roseocanus F/266RG — — 27 C. cibarius #roseocanus F/266R2 — — 28 C. cibarius #roseocanus F/266R6 — — 29 C. cibarius #multiramis P?WR/7./ — — 30 C. cibarius JJRW2 JL%N%01M [9! JQ976981 —— .//R3R/GWR. 31 C. cibarius %MR..Q.R — — 32 C. cibarius F>2/V2.Q — — 33 C. cibarius P?WR/7.W — — 34 C. cibarius FW2RW/Q 35 C. cibarius N‹Q7QV7G — — 36 C. cibarius # longipes P?WR/7.2 — — 37 C. cinnabarinus F/266VQ — — 38 C. cinnabarinus N‹Q7QV7. — — — — N‹6./722 — N‹Q7QV27 — 39 C. congolensis NN&WW &F0TF0"F1? &6/WW.//W JQ976977 JQ976946 JQ976925 40 C. congolensis NN&WV &F0TF0"F1" &6/WV.//W JQ976976 JQ976945 JQ976924

28 IMA FUNGUS Afrocantharellus gen. nov.

Table 1. E ARTICLE No Species Voucher Locality Collection no. (UPS) LSU-GB 5.8-ITS2 GB ATP6-GB 41 C. densifolius NN&2/ &F0TF0"F1K &6/2/.//V JQ976963 JQ976935 JQ976915 42 C. densifolius NN&RQ &F0TF0"F1? &6/RQ.//V JQ976968 JQ976938 JQ976917 43 @\ NN&GG &F0TF0"F1? &6/GG.//V JQ976960 — JQ976913 44 @\ NN&GQ &F0TF0"F1? &6/GQ.//R JQ976962 —— 45 C. formosus F/266VV — — 46 C. formosus F/266V2 — — 47 C. formosus F/266VR — — 48 C. garnierii FG7.WVW — — 49 C. garnierii FG7.WVQ — — 50 C. isabellinus P?WR/7G6 — — 51 C. isabellinus NN&G/ &F0TF0"F1? &6/G/.//V JQ976958 —— 52 C. isabellinus # parvisporus NN&6. &F0TF0"F1? &6/6..//2 JQ976954 JQ976931 JQ976910 53 C. isabellinus # parvisporus NN&.. &F0TF0"F1@ &6/...//R JQ976957 JQ976933 JQ976912 54 C. lateritius N‹Q7QV72 — — 55 C. minor N‹Q7QV76 56 C. minor P?WR/7.G — — 57 C. pallens JJRWW JL%N%01M N9[.//RJQ976984 —— R/GW.W

58 C. persicinus F/266V7 — — 59 C. pseudocibarius NN&/. &F0TF0"F1? &6//..//2 JQ976949 JQ976928 JQ976908 60 C. pseudocibarius NN&/R &F0TF0"F1@ &6//R.//2 JQ976952 JQ976929 JQ976909 61 C. pseudoformosus @M.GW/W6 — — 62 C. rhodophyllus P?WR/7.R — —

63 C. ruber NN&V/ &F0TF0"F1" &6/V/.//W JQ976969 JQ976939 JQ976918 64 C. ruber NN&2R &F0TF0"F1K &6/2R.//W JQ976966 JQ976936 — 65 C. subalbidus F/2662Q — — 66 C. subalbidus F/266R/ — — 67 C. subalbidus F/2662V — — 68 C. subalbidus F/2662W — — 69 C. subalbidus F/26627 — — 70 C. tomentosus NN&VQ &F0TF0"F1? &6/VQ.//W JQ976973 JQ976942 JQ976921 71 C. tomentosus NN&V7 &F0TF0"F1? &6/V7.//W JQ976974 JQ976943 JQ976922 72 Cantharellus P?WR/76W — — 73 Cantharellus P?WR/7.. — — 74 Cantharellus P?WR/7.Q — — 75 Cantharellus P?WR/7G/ — — 76 Cantharellus P?WR/7.V — — 77 Cantharellus P?WR/76Q — — 78 Cantharellus P?WR/7.6 — — 79 Cantharellus F>.W667. — — 80 Cantharellus F/266VW — — 81 Cantharellus P?WR/7.7 — — 82 Cantharellus. NN&W/ &F0TF0"F1? &6/W/.//W JQ976975 JQ976944 JQ976923 83 Cantharellus. NN&W7 &F0TF0"F1? &6/W7.//W JQ976979 JQ976948 JQ976927 84 Clavulina cinerea F?.R7.66 F!6QR7W2 — Clavulina N‹6./72W 85 Craterellus chantarellus. # P?WR/767 — — intermedius

VOLUME 3 · NO. 1 29 "#$%4"*%

Table 1. E No Species Voucher Locality Collection no. (UPS) LSU-GB 5.8-ITS2 GB ATP6-GB 86 Craterellus cornucopioides FW//6QQ — — >!7/W7VW 87 C. cornucopioides F>.W7RW. — — ARTICLE 88 C. lutescens JJRWR JL%N%01M [.//R JQ976982 —— R/GW/G 89 C. lutescens %MR..W2V — — 90 C. melanoxeros JJRWV JL%N%01M F.//Q JQ976983 —— 226QVR 91 C. sp. P?66GR.7 — — 92 C. tubaeformis F!.QWQR6 — — F!GQRVG. 93 C. tubaeformis JJRW. JL%N%01M +./6/ JQ976980 —— 94 C. tubaeformis N‹Q7QW26 — — 95 Dacrymyces chrysospermus F!.QWQRR — %MGG7.27 96 Hydnum rufescens F.7G6QW — — 97 Multiclavula mucida F!.QWQWR — —

./ 3+ ! +JM RQ="&J. (E; et al .//Q J#4 9 & .//7 < +JMF(E; F&(V M F"E >? # /66 K967771#GR72 ERR @ 9 @ .//G ( .//Q GW E6W. EG/GR72 E @&;¨@ RR2R E6W. E6/ +JM < RQ="&J. F&(VX @&;¨@¨" W. EF#/R +JM # ](E; # @&;¨@ ? ?J( ' (E;N#?FJ' +JM3RQ="&J.3F&(V= # \ ? F \ = < ?+ 1 W/ ] ?= @9K677V &#F = % <+JM = +JMRQJ="&J.F&(V ;F

Table 2. (RQJ="&J."&J

Primer Sequence forward "&JGE RRM@EF&E@F&@FF@FFE@EF@&MGR reverse Lcan RRM@&EE@F@&&@&F@F&@F@MGR forward RQJ RRME@F&@FF@FFE@EF@E@MGR forward R RRMEF&E@F@&E&&&@FFE@EFFFEMGR reverse +; RRMF&E@F@&E&&&@FFE@EFFFEMGR

30 IMA FUNGUS Afrocantharellus gen. nov.

AF287875 Multiclavula mucida ARTICLE AY293187 Hydnum rufescens DDT63 A. platyphyllus f. cyanescens DDT41 A. platyphyllus f. platyphyllus DDT03 A. platyphyllus f. platyphyllus DDT78 A. platyphyllus f. platyphyllus DDT17 A. splendens DDT57 A. splendens 1/100/98 DDT31 A. fistulosus DDT43 A. fistulosus DDT14 A. symoensii DDT66 A. symoensii DDT11 A. symoensii DDT04 A. symoensii DDT36 A. symoensii

DDT67 A. symoensii Afrocantharellus AY041169 C. persicinus AY041167 C. sp. HM750924 C. cibarius var. longipes DQ898693 C. cibarius AJ406428 C. cibarius SS577 C . pallens AY041159 C. cascadensis AY041158 C. cascadensis AY041161 C. cascadensis AY041160 C. cascadensis AY041152 C. cibarius var. roseocanus AY041154 C. cibarius var. roseocanus AY041153 C. cibarius var. roseocanus EU522825 C. cibarius AY745708 C. cibarius AY041151 C. cibarius var. roseocanus AY041150 C. subalbidus AY041146 C. subalbidus AY041148 C. subalbidus A AY041147 C. subalbidus * AY041149 C. subalbidus HM750926 C. sp. HM750918 C. sp. HM750927 C. cibarius HM750931 C. isabellinus HM750929 C. sp. AY041156 C. cibarius var. cibarius AY041157 C. cibarius var. cibarius SS574 C . cibarius AY041155 C. cibarius var. cibarius AY041166 C. formosus AY041164 C. formosus AY041165 C. formosus DQ898694 C. lateritius HM750919 Cr. cantharellus var. intermedius HM750930 C. sp. AJ271192 C. sp. HM750921 C. sp. GU237071 C. pseudoformosus HM750916 C. appalachiensis DDT76 C. congolensis DDT77 C. congolensis DQ898690 C. appalachiensis 1/80/93 HM750920 C. cibarius var. multiramis HM750917 C. sp. HM750922 C. sp. * 0.96/99/89 HM750925 C. rhodophyllus DQ898691 C. minor HM750928 C. sp. AY041168 C. cinnabarinus DQ898692 C. cinnabarinus HM750923 C. minor AY392767 C. garnierii AY392768 C. garnierii DDT30 C. isabellinus DDT40 C. densifolius DDT58 C. densifolius D DT12 C. isabellinus var. parvisporus D DT22 C. isabellinus var. parvisporus DDT68 C. tomentosus DDT69 C. tomentosus DDT02 C. pseudocibarius DDT05 C. pseudocibarius DDT33 C. floridulus DDT38 C. floridulus DDT45 C. ruber

DDT60 C. ruber Cantharellus s.str. SS572 C . tubaeformis AF287851 C . tubaeformis DQ898741 C . tubaeformis 1/100/96 SS576 C . melanoxeros EU522746 C. lutescens SS575 C. lutescens HM113529 C. sp. AY700188 C. cornucopioides AJ279572 C. cornucopioides Craterellus 0.1

Fig. 2(7.&6R2< +JM&Multiclavula mucida&# ((?(?+#((17R]?(?+1W/]F 6//]

VOLUME 3 · NO. 1 31 "#$%4"*%

Dacrymyces chrysospermus Botryobasidium isabellinum Clavulina sp. DDT36 Afrocantharellus symoensii DDT66 A. symoensii ARTICLE * 1/92 DDT67 A. symoensii 1/100/93 DDT57 A. splendens DDT17 A. splendens 1/97 DDT03 A. platyphyllus f. platyphyllus DDT78 A. platyphyllus f. platyphyllus Afrocantharellus DDT40 Cantharellus densifolius 1/90/100 1/91 DDT58 C. densifolius DDT 22 C. isabellinus var. parvisporus A 1/87 DDT 12 C. isabellinus var. parvisporus * 1 DDT68 C. tomentosus DDT69 C. tomentosus 1 1 1 DDT02 C. pseudocibarius DDT05 C. pseudocibarius DDT33 C. floridulus 0.85/- DDT70 C. sp. 2 1/94/100 * DDT79 C. sp. 2 1/97/96 DDT45 C. ruber * DDT60 C. ruber * DDT76 C. congolensis 0.81/- DDT77 C. congolensis

C. cinnabarinus Cantharellus s.str. * Craterellus tubaeformis 0.1 C. cornucopioides Craterellus

Fig. 3.(.Q'&66W< +JM3RQ="&J.3F&(V&Dacrymyces chrysospermus&# ((?(?+#((17R]?(?+1 W/]F6//]

<67/V 6///#;FJ&; F # Phylogenetic relationships were inferred separately +JM < ?+ +JM3RQ="&J.3F&(V ;F

32 IMA FUNGUS Afrocantharellus gen. nov.

Table 3. Morphological features of Afrocantharellus and Cantharellus. ARTICLE Afrocantharellus Cantharellus Basidiome colour always variegated Mostly uniformly coloured Hymenophore well-developed with thick diverging folds Poorly-developed, without folds or with thin folds but never with thick diverging folds Folds thick, blunt, always decurrent and distantly spaced ;# spaced Clamp connections Mostly absent Mostly present

supported branches, one containing only African species in conjunction with the rather distinctive morphological AfrocantharellusX((56/X?(56//X?+57Q characteristics of having a well-differentiated hymenophore 0 P F 0 with diverging folds, the variegated colour of the basidiomes Caledonian species, Cantharellus s. str.((5/7VX?(577X & G ! 2 ?+5Q7 J Cantharellus s. str. recognition of Afrocantharellu # and Afrocantharellus were mostly resolved, although with molecular evidence and morphological features, we suggest # emendation revised circumscription of Cantharellus to <C. symoensii, and the elevation of CantharellusAfrocantharellus to generic The combined data set phylogeny # & = ! G Craterellus F&(V < TAXONOMY ((56/X?(56//X?+56//= A, Cantharellus s. lat. ((56/X ?(56//X ?+56//F Afrocantharellus % 9 & gen. species in our sampling traditionally placed in Cantharellus stat. nov. Cantharellus s. lat. # ??R6QVQW Cantharellus s. str. and Afrocantharellus, with high support Basionym: CantharellusAfrocantharellus%9 # ((56// ?(57WX ?+57V ((56// Buyck, Docums Mycol1211RR.//6 ?(56//X?+57G# " Type: Cantharellus symoensii PBull. Jard. bot. État +JM +JM3RQ="&J.3 Brux361G2G67VV F&(V Cantharellus s. lat. includes two strongly supported subclades, Cantharellus s. str. and Afrocantharellus for all \### !.MG XGRM6Q Afrocantharellus, the sister-clade of Cantharellus s. with very well-differentiated, thick, blunt, distantly spaced and str #

Key to the species of Afrocantharellus

6 GRM6Q. 6RM.R 1. /##

.6 VM6QX=XX X2A. splendens =XGRM6.X=# XX G

G. ‹56VM.GXX= margin RA. symoensii ‹56.M6RX 2

2G JGA. platyphyllus f. cyanescens Stipe, cap margin and folds without glaucous or bluish tinges .A. platyphyllus f. platyphyllus

VOLUME 3 · NO. 1 33 "#$%4"*%

Species of Afrocantharellus " A. symoensii but lacks the pink tinge on 6 A!# ## & 9 the basidiomes of A. symoensiiX &comb. nov. basidiospores, rather than the ellipsoid ones of A. symoensii ??Q//.Q/

ARTICLE !2 Q P 67VV Basionym: @' & 9 Pet al.677R.//G Cryptogamie, Mycol. 2916GG.//Q X' Y Tanzania: Coast region: Kisarawe, Type: Tanzania: Coast regionK# /V /2RG.RR J /G7 6RR2QRR % Tibuhwa 1041.2006 M(J MNJ?X K /V /2RG.RR J /G7 6RRRVRR % [" " JMF # /V R.RG2RR J GW VWR.7R % by Brachystegia, Combretum and JulbernardiaF .//W Tibuhwa 1003.2004 M(J (E MNJ?X \" " Madibira Tibuhwa D 43.2007M(JMX1(EMNJ? /Q 6RR/QRRJGR 6WR.6RR%Tibuhwa 1078.2007M(JMNJ?X M Vigama village, Buyck 98.126 (E Buyck 98.127 (E Buyck 98.130(E Q &et al.//Q G Afrocantharellus platyphyllus f. cyanescens QK& &comb. nov. ??R6QV7G @& !2N Basionym: Cantharellus cyanescens Buyck, ]+ centre, pink hymenophore composed of widely spaced folds, Champ. Comest. l’Ouest Burundi (FG2‘1 and by the smooth hollow stipe, which is slightly twisted or 66.6772 Type: Burundi: 06772Buyck;M X' Y Tanzania: Coast region1 K # K /V /2RG.RR J /G7 6RRRVRR % Tibuhwa D Vernacular names: &P1LX 31.2006 M(J MNJ? Iringa region1 ? /Q 6RR/QRR 1 L K 1 J GR 6WR.6RR % 6Q2W M ? .//W (" Tibuhwa D 59.2007M(JMNJ? Basidiomata = . Cap RM6/ .Afrocantharellus platyphyllusP& comb. nov. f. platyphyllus the orange-red cap, margin and folds especially in young ??R6QV7G Folds deeply decurrent, thick, blunt, Basionym: Cantharellus platyphyllusPBull. Jard. bot. diverging, distantly spaced, strongly meshed, bright yellow État Brux 361G2.67VV Stipe GMV & /7M6G smooth, solid, cylindrical, the same colour as the folds in Type: Democratic Republic of Congo1%#67G. = De Loose 31;M Basidia#2R/MRR/MWR/&R/MW/MWR3‹5 VGM7Q.M2BasidiosporesWRM6//M6/V& Vernacular names1&1 R.MV6MVR3‹56GM6R LP1L SuprapellisQ/M6R3 Clamps Basidiomata = Cap GRM7R Folds well- Q 6772 & developed, yellow, thick and distantly spaced, forking or M#Stipe6RMVR&6M6R @&<\ Basidia#226MRR2MW//&R.MW.M7.3‹5 deep orange cap interrupted by blue or glaucous tinges and VVM7.BasidiosporesVGMWRMQV&R/M folds which are strongly meshed and not purely yellow but V.MW63‹566M6RSuprapellis6/M6.3 M & ' Clamps stipe and folds distinguish it from the otherwise very similar A. platyphyllus platyphyllus Q ; 6772 N;EP67VV& Q6772 P et al. 677R et al. ./// T J9L133#= X'Y Tanzania: [""M /V RRR66RR J /GW GRR./RR % Tibuhwa 1063.2007 M(J @&' MNJ?Tibuhwa 1056.2007M(JMNJ?X@" Kisarawe, to crimson colour, especially towards the cap centre, which /V /2RG.RRJ/G7 6RRRVRR%Tibuhwa 1034.2006M(JMNJ?

34 IMA FUNGUS Afrocantharellus gen. nov. ARTICLE

AB3 cm 2 cm

C 3 cm D 3 cm

E 3 cm F 2 cm

Fig. 4Afrocantharellus and Cantharellus species showing morphological differences of the hymenophores: A. Afrocantharellus symoensiiTibuhwa 1011.2005XM(J. B AC. A. splendensDDT 1053.2011XMNJ?D. A. platyphyllus cyanescens Tibuhwa 1063.2007XM(JE. Cantharellus congolensis Tibuhwa 1076.2007XMNJ?F. C. rufopunctatusTibuhwa 1010.2004XMNJ?F &NN&

2 Afrocantharellus splendens & Type: Burundi: under Brachystegia, Buyck 5518 ; M comb. nov. ??R6QV7. !2E Vernacular names1&01 Basionym: Cantharellus splendens Buyck, ]+@' K1( Comest. l’Ouest Burundi (FG2‘166.6772

VOLUME 3 · NO. 1 35 "#$%4"*%

Basidiomata Cap QM6Q = \= Folds thick, blunt diverging, distantly spaced, pale yellow towards the margin, and the bright yellow, distantly spaced, Stipe .RMW & 6.MGR "C. longisporus, solid, subcylindrical, slightly attenuated toward the base, of but differs in the differently shaped spores, and in lacking the same colour as the cap but paling to white toward the % 9 677Q et al.

ARTICLE Basidia M# 2//M27WM ./// RW2 & R2MVVMWW 3 ‹ 5 VWM76 Basidiospores Q6M77M6./&GWM2.M2W3‹5./M.W Q%9677Q# Suprapellis a detailed description of the holotype, and more descriptions RRMQ/3 Clamps 3 6772 P 67VVPet al.677R.//G Q 6772&et al../// X' Y Tanzania: Morogoro region: SUA forest #/V R6R..RRJGW G7R.GRR%Tibuhwa 1004.2005M(J(E @ & MNJ?XM/V RRR66RRJGW G2R./RR%Tibuhwa \ = 1011.2005 M(J (E MNJ?X @ " K which recall those of A. symoensii and A. platyphyllus& # J /V /2RG.RR J /G7 6RRRVRR % Tibuhwa 1007.2005 M(J pigmentation of the cap stains the hands upon handling, and (EMNJ?Tibuhwa 1036.2005M(JMNJ?Tibuhwa 1037.2006 microscopically a trichoderm pileipellis distinguishes it from M(J MNJ?X [+% " @=( # /. R.R.7RRJG. /WR.WRR%Tibuhwa 1014.2004M(J(EMNJ?X 9"?#/2 R7R..RRJ/G. 2/R./RR% Q6772 Tibuhwa 1021.2005 M(J MNJ?X \" " Madibira forest, Tibuhwa 1067.2007 M(J MNJ? Tibuhwa 1066.2007 M(J X'Y Tanzania: [""M MNJ?X Dar e Salaam District: bought in a market, Buyck 98.113 /V RRR66RR J /GW G2R./RR % Tibuhwa 1057.2007 M(J (E MNJ?X @ " Msanga area, near Chanika village, MNJ?X[+%"@=;#/G /7RR/RR Buyck 98.011(EMNJ? JG. /2RR.RR%Tibuhwa 1017.2005M(JMNJ?

R Afrocantharellus symoensii P & DISCUSSION comb. nov. ??R6QV76 & † !2F subclades in the LSU- phylogeny of Cantharellus s. str., Basionym: Cantharellus symoensiiPBull. Jard. bot. < = Cantharellus État. Brux. 361G2G67VV congolensis ((56//X ?(5Q/X ?+57G <# C. congolensis and C. garnierii Type: Democratic Republic of Congo1K67RQ 0 P Cantharellus Symoens 6037;M congolensis ! 2% Afrogomphus %9.//6@\, which was Vernacular names1 & 0 1 ? Rubrinus%9.//6# 1 L P 1 L #=& K1(( name of the generic type species, C. cibarius, is present on several subclades in the LSU analysis of Cantharellus s. Basidiomata = Cap GRMQ str. supports the opinion that this name may either embrace smooth, orange-red disrupted with pale pink and yellow several cryptic species, or that many morphologically similar Folds thick, blunt, < # # Stipe [ <# .RM2&/7M.< with critical morphological studies will further elucidate the maturity, cylindrical but slightly wider towards the cap, of the < Basidia#GQ.M2QWMR7G Afrocantharellus was a strongly supported clade in the & R/MVRMQ 3 ‹ 5 VGM6// Basidiospores W2M +JM!.# 7/M6/V & 2RM27MR. 3 ‹ 5 6VM.G +JM#Afrocantharellus SuprapellisWRM6/3Clamps is, however, strongly supported in the three-gene phylogeny !G# Distribution: ; 6772 < A. splendens. ! A. N;EP67VV& splendens NN&6W NN&RW et al. ./// P et al. 677R T +JM3RQ="&J.3F&(VF&(V %9677Q shorter in one, however, A. splendens is monophyletic in the +JM!. @&Afrocantharellus Afrocantharellus, as represented recently by C. F" platyphyllus and C. symoensii==

36 IMA FUNGUS Afrocantharellus gen. nov.

69P./66et al./66 N0F Molecular Phylogenetics and Evolution 22: ARTICLE " WVM7/ the same in both papers, the systematic arrangement follows ?PNJ+KP+%+%+@ %9.//6 .//R& & †=Homobasidiomycetes =6=##< Systematics and Biodiversity 3166GM6RW in comparison to RPB.?et al.//W"+JM 6772 ]+ ^ @'" @ ! . @ is within Afrocantharellus l’Ouest du Burundi ( F G2‘ <1 F&Cantharellus F@EN# & et al .//Q, and also morphologically 0677R% Parvocantharellus as Belgian Journal of Botany 12816G6M6GQ %9.//6 PB.//QF P# CantharellusInoculum 591.. molecular data place this species in Afrocantharellus, and P B ./66 & =6 ' thus the absence of clamp connection is not a synapomorphy to species delimitation in the Cantharellus cibarius < for Afrocantharellus & Afrocantharellus are MJFFungal Diversity 491GRM2V =&G E E E< F P B ./66 Cantharellus #< texensis # &< C. # " A. cinnabarinus#=6'Mycologia 103: symoensii A. platyphyllus platyphyllus, which in the 6/GWM6/2V A. symoensii[< %@K#F.///F# included are A. platyphyllus cyanescens, A. splendens, the genus CantharellusBasidiomycota, Cantharellaceae and<%9.//6 &Nova Hedwigia 711GM2 species to Cantharellus Afrocantharellus, < E ! @ ? > ( P=E > for <P#Afrocantharellus ? @ P .//Q ( # by having a well-differentiated hymenophore with diverging cryptic species and geographical structure in the cosmopolitan folds, and all species apart from < lack clamp Tricholoma scalpturatum Microbial Ecology 561R6GMR.2 ; N?N%J>L.///? < 9 of Craterellus1+JMN0F' P ./66 et al ./66 " # Mycological Research 1041GQQMG72 ' N J? K F ( ?% .//G E @Cantharellus cibarius ( Cantharellus formosus and C. minorE =Molecular Ecology 12: is clearly the best approach to make progress in the study 6V/WM6V6Q of genera with a rather uniform morphology where few %@.//GThe genus Cantharellus(N? #?# 0RP0( that we do not have clear morphological synapomorphies for % @ 677Q E  ' all monophylethic groups within former Cantharellus s. lat. genre Cantharellus F' 1 '' < Belgian Journal of Botany 13116G7M627 %@.//60 the genus CantharellusDocuments Mycologiques 1211RRMRV % @ E ; .//6 0 ACKNOWLEDGEMENTS and combinations in cuboid-spored Entoloma species from ?Mycological Research 10516622M662Q LJ=JF;%E"J %@JNL;BF.//70 Programme at Uppsala University and the Molecular Biology project of Cantharellus Basidiomycota, Cantharellaceae M#NJL ?Fungal Diversity 361RWMVW ; (E M(J !&((EL@6772+# L N0F < Mycological Research 981V62MV6Q # ! &( N ;+ E L@ >L 677W Phylogenetic relationships within the Cantharellaceae inferred ' N0F REFERENCES Mycological Research 101162.GM62G/ !=# &@ > &J +<= & K†=-; .//W ? E 67V7 ? # Amanita and phylogenetics and delimitation of species in Cortinarius section amonograph on its section LepidellaPersoonia 51.QRMRW7 CalochroiBasidiomycota, Agaricales % Molecular ?PNJ.//.P=# Phylogenetics and Evolution 441.6WM..W of Homobasidiomycetes= @ > + @F [R0 K 0 PE & N+ .//V

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#!F . 6 Amanita muscariaMolecular Ecology 151..RM.G7 Basidiomycota, FungiMolecular Phylogenetics and Evolution @J@[.//GF 4312G/MR6 <Systematic ?# >? 0 ;P K N J? & Biology 521V7VMW/2 ?(?+&?JL%?@J

ARTICLE P?0&?+677R% N%+@+%+%+KPB &Karstenia 35 J16M7. ;.//V&1 P?0&?+.//G&1 Mycologia 98: Norrlinia 1016M.// 7GWM72Q P ( 67RQ E E 0+@P>B#E(&./6.( ? @=! """ Cantharellineae Bulletin du delimitation in the genus Coprinellus with special emphasis on Jardin Botanique de l’Etat Bruxelles 281GQRM2GQ Mycologia 1041.R2M.WR P(67VVEKBulletin du Jardin [">?@=%<KJ".//7J Botanique de l’Etat Bruxelles 361GGRMGR. delimitation in the European species of ClavulinaCantharellales, P &L ? ; E F? K F .//R 0 Basidiomycota"&J Clavulina ( ? @ Mycological Research 11316.V6M6.W/ Mycolological Progress 41G2GMGR/ (N0#+N%?;.//GEcology and Management P>.//V[Trends in of Commercially Harvested Chanterelle Mushrooms. [USDA Ecology9Evolution 21122WM2R/ !J#&;(0L=@&;=RWV‘( PNJN?>.//6F [;1(0;JMJNF!J# among wood decay mechanisms, mating systems, and substrate (%?PNJN?>6777( Systematic Biology 501 .6RM of cantharelloid and clavarioid homobasidiomycetes based .2. N0F ' Mycologia 91: PNJ@+N?>.///%# 722M7VG Nature 407: ( N .//Q †?&1 ( ? F# R/VMR/Q Molecular Biology and Evolution 2516.RGM6.RV P NJ ( %? + % + @ N ?> 677W ;'!P>(.//RMrBayes. BG6 Evolution of gilled mushrooms and puffballs inferred from X N0F ' Proceedings of the National J#4 J & + .//7 &< Academy of Sciences, USA 9416.//.M6.//V Sporodictyon Verrucariaceae 0 % K K K K & P ? & .//R ?F!!& # R1 †FN # ' Taxon 581RQRMV/R Nucleic Acids Research 331R66MR6Q J J=E & \=!+ (=@ ./66 Cantharellus in southwestern K K ? K K K ? & .//. ?F!!&1 # E1Mycotaxon 11612GWM ' 22V !Nucleic Acids Research 301G/R7MG/VV J E L .//7 A preliminary list of the macrofungi KF?&N6777MV1 recorded in the Vumba.#= < Boletales Molecular Phylogenetics and J F .//V ;F .//Q F ; 10817QGM6//. ;F; P &L .//2 0 Clavulina from AgaricalesMolecular Phylogenetics and Evolution 3516M./ @Mycologia 961VR/MVRW ? ( L T ? E >? +L 0 ;P &NK#F&+.//Q@' PKLPBF>!JE++@% #&Cryptogamie Mycologie 2916.7M6GR ?+ N> LFL !=# & @ TL K ;L B ; P ? 677/ ; J>EB%E+T+&>!?PK N0F# ?KJ?&B>PNJ.//WE CryptococcusJournal of Bacteriology 17212.GQM2.2V

38 IMA FUNGUS