Research Article |
Corresponding author: Hyppolite L. Aïgnon ( hyppoliteaignon@yahoo.com ) Academic editor: Zai-Wei Ge
© 2021 Hyppolite L. Aïgnon, Sana Jabeen, Arooj Naseer, Nourou S. Yorou, Martin Ryberg.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Aïgnon HL, Jabeen S, Naseer A, Yorou NS, Ryberg M (2021) Three new species of Inosperma (Agaricales, Inocybaceae) from Tropical Africa. MycoKeys 77: 97-116. https://doi.org/10.3897/mycokeys.77.60084
|
Here, we describe three new species of Inosperma from Tropical Africa: Inosperma africanum, I. bulbomarginatum and I. flavobrunneum. Morphological and molecular data show that these species have not been described before, hence need to be described as new. The phylogenetic placements of these species were inferred, based on molecular evidence from sequences of 28S and RPB2. Additional analysis using ITS dataset shows interspecific variation between each species. Phylogenetic analyses resolve I. flavobrunneum in Old World Tropical clade 1 with weak support, I. bulbomarginatum is sister of Old World Tropical clade 1 and I. africanum is indicated as sister to the rest of Inosperma. Complete description and illustrations, including photographs and line drawings, are presented for each species. A new combination of Inocybe shawarensis into Inosperma is also proposed.
Ectomycorrhizal, molecular systematics, phylogeny, taxonomy, West Africa
Inocybaceae Jülich (Basidiomycota, Agaricales) is a family of ectomycorrhizal species, forming symbiotic association with more than 23 families of vascular plants (
Recently, Inocybaceae was revised to include seven genera, Auritella Matheny & Bougher, Inocybe (Fr.) Fr., Inosperma (Kühner) Matheny & Esteve-Rav., Mallocybe (Kuyper) Matheny, Vizzini & Esteve-Rav., Nothocybe Matheny & K.P.D. Latha, Pseudosperma Matheny & Esteve-Rav. and Tubariomyces Esteve-Rav. & Matheny (
In this study, we describe three new species of Inosperma from West Africa, based on morphological characters, as well as analysing their phylogenetic position using multigene molecular analysis of 28S and RPB2 sequences data.
Specimens were collected in Benin in Okpara Forest (9°15.13'N, 2°43.05'E), N’dali Forest Reserve (09°45.73'N, 2°19.93'E), Toui-Kilibo Forest Reserve (8°32.74'N, 2°40.42'E) and Alibori Superieur Forest Reserve (10°23.76'N, 2°5.15'E). Additionally, specimens were collected in, Burkina Faso in the Forest Reserve of Kou (10°55.86'N,4°51.83'W); Ivory Coast in Gbeke Region (7°40.52'N, 4°54.48'W), Guinea in National Park of Haut Niger (10°30.76'N, 9°57.68'W) and Togo in Central Region (09°20.38'N, 1°14.44'E).
The habitats are woodland dominated by Isoberlinia doka Craib & Stapf, I. tomentosa (Harms) Craib et Stapf, Uapaca togoensis Pax or gallery forest dominated by Berlinia grandiflora (Vahl) Hutch. Specimens were preserved by drying on an electric dryer (type Stöckli Dörrex) for 24 hours at 45 °C. All studied materials are deposited at the Mycological Herbarium of Parakou University (UNIPAR).
Specimens were photographed in the field with a digital camera Sony FE. Colour codes are described using
of specimens were rehydrated and examined directly in 3% potassium hydroxide (KOH) and Congo red. Drawings of microscopic characters were made with the aid of a drawing tube attached to a Leica DM2700. Microscopic characters were drawn at magnification 1000×. Spore measurements were made from 40 spores for each species. We measured length (L) and width (W) of the basidiospores and calculated the ratio Q = L / W. Measurements of basidiospores and basidia excluded the apiculus and sterigmata, respectively and are given as (a–)b–c(–d), where (a) = extreme minimum value, range b–c contains minimum of 90% of the calculated values and (d) = extreme maximum value as used in
Genomic DNA was extracted from dried specimens by QIAGEN® plant mini kit following the manufacturer’s instructions and PCR products were cleaned using ExoSAP-IT (
Nineteen new sequences were generated (Table
For phylogenetic analysis, the dataset of 28S and RPB2 was generated using Geneious 7.0.2 (
For Bayesian Inference (BI) analyses, GTR models with gamma-distributed rate heterogeneity and a proportion of invariant sites parameter were assigned to each partition as indicated above, using MrBayes 3.2.7 (
List of species, geographic origin and GenBank accession numbers of ITS, 28S and RPB2 sequences used in the molecular analysis; the new species and new combinations are in bold.
Species | Voucher | Country | ITS | 28S | RPB2 | References |
---|---|---|---|---|---|---|
Auritella brunnescens Matheny & Bougher | PBM3174 | Australia | KJ702344 | JQ313571 | KJ702349 |
|
Auritella dolichocystis Matheny, Trappe & Bougher | Trappe 24844 | New South Wales | AY380371 | AY337371 |
|
|
Auritella fulvella Matheny & Bougher | BRI:AQ669485 | Australia | KJ702355 | KJ702353 | KJ702357 |
|
Auritella hispida Matheny & T.W. Henkel | TH1009, TH10379 | Cameroon | KT378203 | KT378208 | KT378215 |
|
Auritella serpentinocystis Matheny, Trappe &Bougher ex Matheny & Bougher | PBM3188 | Australia | KJ729858 | JQ313559 | KJ756402 |
|
Auritella spiculosa Matheny & T.W. Henkel | MCA7031, TH9866 | Cameroon | MF374763 | KT378206 | KT378214 |
|
Inosperma adaequatum (Britzelm.) Matheny & Esteve-Rav. | JV 16501F, JV11290F | Finland | JQ801381 | JQ815407 | AY333771 |
|
I. africanum Aïgnon, Yorou & Ryberg | MR00387 | Togo | MN096189 | MN097881 | MT770739 | This study |
HLA0361 | Benin | MT534295 | MT560735 | |||
HLA0383 | Benin | MT534298 | MT560733 | |||
HLA0353 | Benin | MT534299 | ||||
BRF4157 | Benin | MK908843 | Unpublished | |||
I. akirnum (K.P.D. Latha & Manimohan) Matheny & Esteve-Rav. | CAL 1358 | India | NG_057279 | KY553236 |
|
|
I. apiosmotum (Grund& D.E. Stuntz) Matheny & Esteve-Rav. | AU10560, TENN:062779 | Canada, USA | HQ201336 | JN975022 | JQ846463 |
|
I. bongardii (Weinm.) Matheny & Esteve-Rav. | EL9406 | Sweden | FN550943 | FN550943 | Unpublished | |
I. bulbomarginatum Aïgnon, Yorou & Ryberg | MR00357 | Benin | MN096190 | MN097882 | MN200775 | This study |
HLA0373 | Benin | MT534301 | ||||
HLA0389 | Benin | MT534302 | ||||
HLA0417 | Benin | MT534300 | MT560734 | |||
PC96082 | Zambia | JQ801412 | JN975027 |
|
||
I. calamistratoides (E. Horak) Matheny & Esteve-Rav. | PBM3384 | Australia | JQ815415 | KJ729949 |
|
|
I. calamistratum (Fr.) Matheny & Esteve-Rav. | PBM1105 | USA | JQ801386 | JQ815409 | JQ846466 |
|
I. carnosibulbosum (C.K. Pradeep & Matheny) Matheny & Esteve-Rav. | TBGT:12047 | India | KT329448 | KT329454 | KT32944 |
|
I. cervicolor (Pers.) Matheny & Esteve-Rav. | SJ04024, TURA:4761 | Sweden, Finland | AM882939 | AM882939 | JQ846474 |
|
I. cookei (Bres.) Matheny & Esteve-Rav. | EL70A03 | Sweden | AM882953 | AM882953 |
|
|
I. cyanotrichium (Matheny, Bougher& G.M. Gates) Matheny & Esteve-Rav | TENN:065729 | Australia | JQ815418 | KJ729948 | Unpublished | |
I. flavobrunneum Aïgnon, Yorou & Ryberg | HLA0367 | Benin | MN096199 | MT536754 |
This study |
|
HLA0372 | Benin | MT534290 | MT536756 | |||
I. geraniodorum (J. Favre) Matheny & Esteve-Rav. | EL10606 | Sweden | FN550945 | FN550945 |
|
|
I. gregarium (K.P.D. Latha & Manimohan) Matheny & Esteve-Rav. | CAL 1309 | India | KX852305 | KX852306 | KX852307 |
|
I. lanatodiscum (Kauffman) Matheny & Esteve-Rav. | PBM2451 | USA | JQ408759 | JQ319688 | JQ846483 |
|
I. maculatum (Boud.) Matheny & Esteve-Rav. | MR00020 | Sweden | AM882958 | AM882958 |
|
|
I. maximum (A.H. Sm.) Matheny & Esteve-Rav. | PBM 2222,UBC F33244 | USA,Canada | MG953983 | EU569854 |
|
|
I. misakaense (Matheny & Watling) Matheny &Esteve-Rav. | 96234 (PC) | Zambia | JQ801409 | EU569874 | EU569873 |
|
I. mutatum (Peck) Matheny & Esteve-Rav. | PBM4108, PBM2953 | USA | MG773837 | JQ994476 | JQ846488 |
|
I. neobrunnescens (Grund & D.E. Stuntz) Matheny &Esteve-Rav. | PBM 2452 | USA | EU569868 | EU569867 |
|
|
I. quietiodor (Bon) Matheny & Esteve-Rav. | PAM01091310 | France | FJ936168 | FJ936168 |
|
|
I. rhodiolum (Bres.) Matheny & Esteve-Rav. | PAM00090117 | France | FJ904176 | FJ904176 |
|
|
I. rimosoides (Peck) Matheny & Esteve-Rav. | PBM 2459, PBM3311 | USA | JQ801414 | JQ815426 | DQ385884 |
|
I. rubricosum (Matheny & Bougher) Matheny & Esteve-Rav. | PBM3784 | Australia | NG_057260 | KM406230 |
|
|
I. shawarense (Naseer & Khalid) Aïgnon & Naseer | FLAS-FS9456 | Pakistan | KY616965 | KY616966 |
|
|
Inosperma sp. | DB166 | Democratic Republic of the Congo | KT461385 |
|
||
Inosperma sp. | PC 96013 | Zambia | JQ801383 | JQ815408 | EU600882 |
|
Inosperma sp. | BB3233 | Zambia | JQ801415 | EU600885 |
|
|
Inosperma sp. | G1842 | Zambia | MK278245 | Unpublished | ||
Inosperma sp. | TR220_06 | Papua New Guinea | JQ801416 | JN975017 | JQ846496 |
|
Inosperma sp. | L-GN3a | Papua New Guinea | JX316732 |
|
||
Inosperma sp. | Zam07 | Zambia | FR731653 |
|
||
Inosperma sp. | PBM3406 | Australia | JQ815431 | JQ846498 | Unpublished | |
Inosperma sp. | TJB10045 | Thailand | KT600658 | KT600659 | KT600660 |
|
Inosperma sp. | PC 96073 | Zambia | JQ801417 | EU600870 | EU600869 |
|
Inosperma sp. | PC:96080 | Zambia | JQ801382 | Unpublished | ||
I. vinaceobrunneum (Matheny, Ovrebo & Kudzma) Haelew. | TENN:062709, PBM 2951 | USA | FJ601813 | NG_067775 | JQ846478 |
|
I. viridipes (Matheny, Bougher & G.M. Gates) Matheny & Esteve-Rav. | PBM3767 | Australia | NR_153168 | KP171094 | KM656138 |
|
I. virosum (K.B. Vrinda, C.K. Pradeep, A.V. Joseph & T.K. Abraham ex C.K. Pradeep, K.B. Vrinda& Matheny) Matheny & Esteve-Rav. | TBGT:753 | India | KT329452 | KT329458 | KT329446 |
|
Mallocybe myriadophylla (Vauras & E. Larss.) Matheny & Esteve-Rav. | JV19652F | Finland | DQ221106 | AY700196 | AY803751 |
|
M. subdecurrens (Ellis & Everh.) Matheny & Esteve-Rav. | REH10168 | USA | MH024850 | MH024886 | MH577503 |
|
M. terrigena (Fr.) Matheny, Vizzini & Esteve-Rav. | EL11704, JV 16431 | Sweden | AM882864 | AY380401 | AY333309 |
|
M. tomentosula Matheny & Esteve-Rav. | PBM4138 | USA | MG773814 | MK421969 | MH577506 |
|
M. unicolor (Peck) Matheny & Esteve-Rav. | PBM 1481 | USA | AY380403 | AY337409 |
|
|
Pseudosperma lepidotellum (Matheny & Aime) Matheny & Esteve-Rav. | TENN066442 | Guyana | JN642233 | NG_042597 | MH577508 |
|
P. pluviorum (Matheny & Bougher) Matheny & Esteve-Rav. | BRI:AQ794010, PERTH:08556466 | Australia | NG_057259 | KM406221 |
|
|
Pseudosperma sp. | PBM3751 | Australia | KP636851 | KP171053 | KM555145 |
|
Pseudosperma sp. | TR194-02 (M) | Papua New Guinea | JQ408793 | JN975032 | JQ421080 |
|
Tubariomyces inexpectatus (M. Villarreal, Esteve-Rav., Heykoop & E. Horak) Esteve-Rav. & Matheny | AH25500 AH20390 | Spain | GU907095 | EU569855 | GU907088 |
|
T. similis Della Magg., Tolaini & Vizzini | RFS0805 | Spain | GU907096 | GU907092 | GU907089 |
|
T. hygrophoroides Esteve-Rav., P.-A. Moreau & C.E. Hermos. | P05112008 | France | GU907097 | GU907094 | GU907090 |
|
Inosperma is indicated as monophyletic with full bootstrap support. All three of the species described here, Inosperma africanum I. bulbomarginatum and I. flavobrunneum, are members of this genus. Phylogenetically, I. africanum is indicated as sister to the rest of Inosperma, with full support (99.9% SH-aLRT values, 100% ML ultrafast bootstrap, 1 BPP). The Old World Tropical clade 1 is retrieved with strong support (93.8% SH-aLRT values, 99% ML bootstrap, 1 BPP) and I. bulbomarginatum is indicated as the sister of Old World Tropical clade 1 with full bootstrap support (100% SH-aLRT values, 100% ML Ultrafast bootstrap, 1 BPP). The sequences of collection PC96082 are very similar to the sequences of I. bulbomarginata that we infer to be of the same species. Inosperma flavobrunneum is nested in Old World Tropical clade 1 as sister species to three undescribed collections, BB3233, G1842 and PC96013, all from Zambia with weak support.
Inosperma africanum is distinct from all species of Inosperma and truly outstanding by its vinaceous to red colouration.
Pileus 8.5–15 mm diam., convex to plane, uniform, surface fibrillose, vinaceous to red (8F8), surface rimose, dry. Lamellae moderately close, subventricose, narrowly attached, 0.5–1 mm deep; vinaceous, sometimes light pinkish (8F8), edges fimbriate, vinaceous (8B8). Stipe 15–23 × 0.5–1 mm, cylindrical, central, fibrillose, swollen, bulbous at the base, veil none with the lower two thirds pinkish-white (8A3) and the upper third light vinaceous (8A5). Odour and taste not distinctive. Basidiospores (6.2) 8–10 (10.3) × (3.8) 4–6.8 (7) μm, avl × avw = 8.3 × 5.3 μm, Q = (1.2) 1.1–2.1 (2.2), avQ = 1.6, smooth, (sub) globose to cylindrical, sometimes ellipsoid. Basidia 18–47 × 7–10 μm, clavate, 3–4 sterigmate, hyaline. Cheilocystidia 22–54 × 8–12 μm, cylindrical to clavate, thin-walled, hyaline. Pleurocystidia absent. Pileipellis a cutis with cylindrical, smooth, thin-walled hyphae, 6–20 μm diam., negative reaction of pileus surface in KOH. Stipitipellis a cutis radially arranged, hyphae 5–13 μm diam., parallel, sometimes septate, filamentous. Caulocystidia 22–63 × 8–13 μm, fusiform sometimes utriform, observed on the upper third of the stipe. Clamp connections present.
Currently known from Benin, Burkina Faso, Guinea, Ivory Coast, Togo.
Scattered in Tropical Woodlands dominated by Isoberlinia doka and I. tomentosa or gallery forests dominated by Berlina grandiflora.
africanum, referring to the distribution in Africa.
Benin, Borgou Province, N’dali Region: 8°32.74'N, 2°40.42'E, on soil in Woodland dominated by Isoberlina doka, 30 August 2017 in Forest Reserve of N’dali, Leg. Aïgnon HL., Voucher (HLA0461) GenBank accession: ITS (MT534297) and LSU (MT560732). Benin, Borgou Province, Tchaorou Region: 9°15.28'N, 2°43.38'E, on soil in forest of Okpara in woodland dominated by I. doka, 7 June 2017, leg. Aïgnon HL., Voucher (HLA0353) GenBank accession: ITS (MT534299). Benin, Borgou Province, N’dali Region: 8°45.73'N, 2°19.93'E, on soil in Woodland dominated by Isoberlina doka, 8 July 2013, leg. Ryberg M., Voucher (MR00361). Benin, Province, Boukoumbe, North Region: 10°14.45'N, 1°7.00'E, on soil in Woodland dominated by Isoberlina doka, 25 July 2020 in Koussoukouangou waterfall, Leg. Aïgnon HL., Voucher (HLA0736). Burkina Faso, Kenedougou Province, Toussiambandougou Region: 10°55.86'N, 4°51.83'W, on soil in gallery forest dominated by Berlina grandiflora, 27 June 2018, leg. Aïgnon HL., Voucher (HLA0353). Ivory Coast, Kekrekouakoukro Province, Bouake, Gbeke Region: 7°40.52'N, 4°54.48'W, on soil in Woodland dominated by B. grandiflora, 11 July 2018, leg. Aïgnon HL., Voucher (HLA0562). Guinea, Faranah Province, Upper Guinea Region, National Park of Haut Niger: 10°30.76'N, 9°57.68'W, on soil in Woodland dominated by B. grandiflora, 4 July 2018, leg. Aïgnon HL., Voucher (HLA0532). Togo, Central Region, Prefecture of Assoli, on the road between Bafilo and Aledjo: 09°20.38'N, 1°14.44'E in Woodlands dominated by I. tomentosa, 7 August 2013, leg. Martin Ryberg, Voucher (MR00387) GenBank accession: ITS (MN096189); LSU (MN097881), RPB2 (MT770739).
Inosperma africanum is nested in Inosperma and indicated as sister to the rest of the genus in our phylogenetic analyses and is very distinct by its small size and a vinaceous to red pileus. It has a wide distribution in West Africa.
Inosperma bulbomarginatum differs from I. flavobrunneum by the smaller size of its basidiomata and larger basidiospores. It is phylogenetically distinct from all other undescribed African Inosperma in Old World Tropical clade 2
Pileus 13–18 mm diam., undulating plane, fibrillose, margin rimose, orange-brown to somewhat cinnamon, greyish-white (8E5), splitting at edge. Lamellae 2–2.5 mm deep, moderately close, narrowly attached, pale grey brown (9B5) to dark brown (9D5), sinuate. Stipe 10–22 × 2–2.5 mm, central, equal, marginate bulb, white to pinkish-buff (7A2), velar remnants. Odour and taste not distinctive. Basidiospores (7.1) 8–12.1 (14) × (4) 4.2–6.7(7) μm, avl × avw = 9.6 × 5.4 μm, Q = (1.3) 1.2–2.3(2.6), avQ = 1.8, smooth, elongate, thick-walled. Basidia (25) 28–40 × 6–12 μm, tetrasporic. Cheilocystidia 20–25 × 10–12 μm, clavate, thin-walled hyaline. Pleurocystidia absent. Pileipellis a cutis, thin-walled hyphae, 3–12 μm diam., cylindrical. Stipitipellis a cutis with subparallel hyphae 3–15 μm diam., septate, filamentous, subhymenium of compact hyphae, any reaction of pileus surface in KOH not observed. Caulocystidia 25–60 × 7–20 μm, ovoid to obovoid, sometimes utriform, observed on the upper third of the stipe.
Currently known from Benin and Zambia.
Scattered in Woodland dominated by Isoberlinia doka and I. tomentosa.
bulbomarginatum referring to the presence of a marginate bulb at the base of the stipe.
Benin, Collines Province, Kilibo Region: 8°32.74'N, 2°40.42'E, on soil in Woodland dominated by Isoberlina doka, 22 June 2017 in the Forest Reserve of Toui-Kilibo, leg. Aïgnon HL., Voucher (HLA0389) GenBank accession: ITS (MT534302). Benin, Tchaorou, Borgou Prov, Okpara Forest: 9°15.28'N, 2°43.38'E, on soil in Woodland dominated by Isoberlina doka, 13 June 2017, leg. Aïgnon HL., Voucher (HLA0373) GenBank accession: ITS (MT534301). Benin, Alibori Borgou Prov, Alibori Superieur Forest Reserve: 10°23.76'N, 2°5.15'E on soil in Woodland dominated by Isoberlina doka, 11 July 2017, in Forest Reserve of Alibori Supérieur leg. Aïgnon HL., Voucher (HLA0417), GenBank accession: ITS (MT534300) and LSU (MT560734).
Inosperma bulbomarginatum is similar to Inosperma cervicolor (Pers.) Matheny & Esteve-Rav., by its orange-brown pileus, but differs from it by the smaller size of the basidiomata and basidiospores, as well as its ecological association with Fabaceae Lindley and/or Phyllanthaceae Martynov and extensive distribution in Tropical Africa. I. cervicolor is associated with Pinaceae Spreng. ex F. Rudolphi and distributed in Europe and North America.
Characterised by yellow to orange-brown pileus, 7–12 × 4–7 μm smooth, thick-walled, ellipsoid basidiospores with cheilocystidia measuring 23–41 × 7–10 μm, clavate, thin-walled.
Pileus 28–38 mm diam., umbonate, yellow (5A3) to orange brown (5C5), dark brown in middle, convex when young, plane at maturity, hard, surface rimose, dry. Lamellae emarginated, adnexed and decurrent, yellow brown (5B5). Stipe 27–39 × 5–6 mm, central, cylindrical, uniform; white, equal, solid, hard, base slightly swollen to bulbous, pruinose at the apex. Basidiospores (7.1) 9.2–11.2 (12) × (4.1) 5.7–7 (7.2) μm, avl × avw = 9.2 × 5.7 μm, Q = (1.2) 1.6–2.1 (2.5), avQ = 1.6, smooth, ellipsoid. Basidia 24–40 × 6–14 μm, clavate, 2–4 spored. Cheilocystidia 23–41 × 7–10 μm, clavate, thin walled. Pleurocystidia absent. Pileipellis a cutis thin-walled hyphae 4–16 μm diam., subparallel, compact hyphae, negative reaction of pileus surface in KOH. Stipitipellis a cutis hyphae 5–10 μm diam., septate, filamentous, thick, subparallel, compact. Caulocystidia 23–52 × 9–10 μm, utriform, rare, observed on the upper third of the stipe.
Currently known only from Benin in Soudano-Guinean zone.
Gregarious under Woodland dominated by Isoberlinia doka, I. tomentosa and Monotes kerstingii Gilg.
flavobrunneum referring to yellow to dark brown pileus.
In the phylogenetic tree (Figure
ML tree of 28S and RPB2 sequences showing the placement of Inosperma africanum, I. bulbomarginatum and I. flavobrunneum. Values above or below branches indicate bootstrap proportions SH-aLRT support ≥ 80% / ultrafast bootstrap support ≥ 95% / Bayesian posterior probabilities > 0.95 as shown. Origin of species is given after the name of each taxon. The new species are in red.
1 | Basidiomata large, pileus 28–38 mm diam., yellow to orange-brown, surface clearly rimose, lamellae adnexed and decurrent, subdistant | Inosperma flavobrunneum |
– | Basidiomata small, pileus 8.5–15 mm diam., fibrillose, lamellae close | 2 |
2 | Pileus vinaceous to red, basidiospores 8–10 × 4–7, (sub) globose to cylindrical, sometimes ellipsoid | I. africanum |
– | Pileus orange-brown to somewhat cinnamon, greyish-white, basidiospores 8–14 × 4–7 μm, elongate | I. bulbomarginatum |
For an evolutionarily-consistent taxonomy, we propose the following combination:
Inocybe shawarensis Naseer & Khalid, Mycotaxon 132: 912. 2018. Basionym.
This species is placed in the old Inosperma clade which became the genus Inosperma, but the combination is not made in the study of
The new species exhibit the overall characteristics often observed in Inosperma. These characters include; pileus radially rimose, fibrillose or squamulose and absence of pleurocystidia (
Phylogenetically, I. africanum is nested in Inosperma with full support (99% SH-aLRT values, 100% ML Ultrafast bootstrap, 1 BPP) and I. bulbomarginatum is indicated as the sister of Old World Tropical clade 1 with full support (100% SH-aLRT values, 100% ML bootstrap, 1 BPP). Sequences of Inosperma bulbomarginatum from West Africa and Zambia formed a subclade. Inosperma flavobrunneum is nested in Old World Tropical clade 1 and has sister species undescribed in a collection from Zambia, BB3233, G1842 and PC96013. ML and BI analysis, using 28S and RPB2 sequences data, shows most nodes well resolved; for example, the node uniting Old World Tropical clade 2 with the crown group of Inosperma is supported with 0.97 BPP, but with weak ML bootstrap as shown in
The position of each of these new species is confirmed by single data from ITS (Fig.
We are grateful to the Rufford Small Grants Foundation (grant n° 30738-2) which allowed us to collect some additional samples analysed in this paper, the Swedish Research Council for Environment, Agricultural Sciences and Spatial Planning (grant n° 226-2014-1109) for funding molecular analysis and the Deutscher Akademischer Austauschdienst (DAAD, grant n° PKZ 300499) for granting the University of Parakou with a microscope, type Leica DM5700, that enabled us to perform microscopic investigations. Anneli SVANHOLM, Bobby SULISTYO and Brandan FURNEAUX (Systematic Biology programme, Department of Organismal Biology, Uppsala University) are thanked for their assistance during molecular analyses. We also thank Kassim TCHAN ISSIFFOU and Evans CODJIA (MyTIPS Research Unit, University of Parakou) for their assistance during field data collection. P. Brandon MATHENY (Department of Ecology and Evolutionary Biology, University of Tennessee, USA) and an anonymous reviewer are thanked for their corrections and suggestions to improve our paper.
Partition for phylogeny analysis
Data type: phylogeny data