Comparative studies on microfungi in tropical ecosystems in Ivory Coast forest litter: behaviour on different substrata

Comparative studies on microfungi in tropical ecosystems in

Ivory Coast forest litter : behaviour on different substrata

Angelo RAMBELLI1*, Bonaria MULAS2 and Marcella PASQUALETTI1

1Dipartimento di Ecologia e Sviluppo Economico Sostenibile, Universita della Tuscia, 01100, Viterbo, Italy.2Dipartimento di Botanica ed Orto Botanico, Universita di Cagliari, 09100, Cagliari, Italy.E-mail : [email protected]

Received 1 October 2002; accepted 17 December 2003.

Fungal colonies sporulating on 71 kinds of leaves that fell during the dry season in the Tai National Park (Ivory Coast)

were analysed. A consistent connection between certain fungal species and their substrata was detected among the184 fungal species that were identified. Each fungal species was characterized according to morphological and ecologicalfeatures. Multidimensional scaling showed that certain ubiquitous and common species have morphological characters

distinguishing them from specialised species.

INTRODUCTION

Many studies have been published on the ecology offungi in tropical forest litter and the identification ofspecific fungal communities (Subramanian & Vittal1974, Rambelli et al. 1983, 1984, 1991, Mercado-Sierra1984, Bills & Polishook 1994, Lodge & Cantrell 1995,Læssøe et al. 1996, Matsushima 1971–96, Polishook,Bills & Lodge 1996, Lodge 1997, Calduch et al. 2002).Such studies often resulted in discovery of new fungaltaxa (Rambelli & Ciccarone 1985, Mercado-Sierra,Holubova-Jechova & Mena Portales 1997, Castaneda-Ruiz, Saikawa & Guarro 1999, Pasqualetti & Rambelli1999, Siboe, Kirk & Cannon 1999, Calduch et al.2002). Other investigations have focused on possiblespecialisation among saprotrophs (Pirozynski 1972,Subramanian & Vittal 1979, 1980, Kabi Ouanyou& Rambelli 1990, Læssøe & Lodge 1994, Lodge &Cantrell 1995, Lodge & Læssøe 1995, Mulas &Rambelli 1995, Lodge, Fisher & Sutton 1996, Polishooket al. 1996, Lodge 1997, Pascholati, Piccolo Grandi& Milanez 2001).

Recently, analysis of the microfungi colonising dif-ferent litter species in natural Mediterranean ecosys-tems has been extended to assess both the number ofcolonies per surface unit and the type of optimal andadaptive colonisation (Mulas, Pasqualetti & Rambelli1995, Pasqualetti, Ialongo & Rambelli 1995). This hasincreased our knowledge of ecological characters of

particular fungal species and has shed light on theirroles in differential colonisation and decomposition ofplant debris. In this study, we analysed microfungisporulating on incubated dead leaves in the TaiNational Park in the Ivory Coast in order to detectdifferential effects exerted by the litter of different plantspecies.

MATERIALS AND METHODS

Description of the study area

The study area is part of the Tai National Park locatedin the south-western region of the Ivory Coast. The360 000 ha Park is covered by native forest, a sub-hygrophilous forest representative of the Eremospato-Mabetum vegetation type (Huttel 1975). Rambelli et al.(1983) published an inventory of the plant familiesforming the Park’s vegetation. The park lies in thedrainage basin of the Cavally river that forms the bor-der between Ivory Coast and Liberia. The undulatingterrain reaches about 350 m a.s.l. at some points. Thesoils are mainly saturated ferrallithic with a fine sandysurface structure. The litter layer is thin (3–4 cm deep)and discontinuous, since wind and rain tend to con-centrate the litter into pockets where it mineralisesrapidly due to high temperature and humidity thatfavour microbial growth.

The climate of the area can be defined as humid,megathermic, with a very low moisture deficit overthe year (Rambelli et al. 1983). Annual temperature* Corresponding author.

Mycol. Res. 108 (3): 325–336 (March 2004). f The British Mycological Society 325

DOI: 10.1017/S0953756204009396 Printed in the United Kingdom.

variation is low. Daily temperatures range from amaximum of 27.9 xC in April to a minimum of 25.1 x inJuly–August, with a mean of 26.4 x. Yearly rainfallranges from 1500–2000 mm yrx1 and is seasonal. Thedry season runs from December to February, and therainy season comprises the remaining eight months ofthe year with the exception of two dry weeks in August.Relative atmospheric humidity ranges from 50–75%during the day, with peaks of over 90% at night.

Sampling

Samples were collected yearly in January 1992–95during the dry season (Dec.–Feb.). Neither senescentfreshly shed leaves nor highly decayed leaves of uncer-tain identity were included in the samples. This resultedin samples that were as homogeneous and comparableas possible. The litter samples for each plant specieswere placed in sterile paper bags, and were identifiedby Laurent Ake Assi (Director, Centre National deFloristique, Abidjan University, Ivory Coast). Fifteendamp chambers were set up for each substratum toallow direct observation, collection, and determinationof the fungal species. Voucher specimens are depositedin the ROHB herbarium in Rome.

Characterisation of the fungi

Some ecological and taxonomic characteristics wererecorded for each fungal species. The ecologicalcharacters were classified as follows: S, specialisedspecies present on one or two substrates ; C, commonspecies present on three to ten substrata; or U,ubiquitous species present on more than ten substrata.The fungi were also categorized according to thefollowing morphological features based on directobservations and confirmed by bibliographic data;HC, hyaline conidia; PG, pigmented conidia; UC,unicellular conidia; SC, septate conidia ; LP, low co-nidial production; AP, abundant conidial production;PC, phialidic conidiogenesis ; PE, percurrent coni-diogenesis ; SY, sympodial conidiogenesis ; SS, presenceof sterile setae; and SA, absence of sterile setae.

Data analysis

Data were analysed using an ordination method to re-duce the dimensionality, in which the original n vari-ables are replaced by artificial variables in an attemptto achieve a more efficient representation of data infew dimensions (Podani 1994). The multidimensionalscaling method (MDS) was employed, which does notassume linear relationships between variables ; an inputmatrix of normalised Euclidean distances was utilised(Wilkinson, Hill & Vang 1992). Chi-square tests wereperformed between all variables to determine if theMDS grouped characters were significantly correlated.The table showing the results was ordered using theblock clustering method. The rearrangement of the

data matrix is based on the assumption that therows and columns are classifiable into groups. Matrixrearrangement is useful in fungal ecology such as whenexplaining classification of fungal communities in termsof species groups and visa versa (Podani 1994).

The 23 ubiquitous fungal species (occurring on morethan ten leaf species) were analysed further. For eachspecies pair we determined whether the fungal speciesoccurred together randomly on the same substratum,or whether they were positively or negatively associatedwith each other. The Yule association index (Q) and theasymptotic standard error were calculated, followed bychi-square test to determine whether the Q values weresignificantly different from 0 at P<0.01 (Wardle &Parkinson 1991, Wilkinson, Hill & Vang 1992). Clusteranalysis (Euclidean distance, Ward’s method) was alsoperformed.

RESULTS

Leaves were collected from 71 plant species represent-ing 58 genera and 32 families (Table 1). The most fre-quent families included the Leguminosae (12 species),Euphorbiaceae (7), Ebenaceae (5), and Annonaceae (4)(Table 1). Overall, 184 fungal species belonging to 96genera were observed and identified (Table 2). Thespecies in Table 2 were ordered using the block clus-tering method. Five groups of fungal species (A–E) andsix groups of plant substrata (1–6) can be distinguished.The first fungal group (A) contains species found onmore than one leaf type: these species were present insubstratum groups 1–3 and were more sporadic in theothers. Fungal species with a more specialized behav-iour or those present on few substrata can be seen in thecentral part of the Table (B and C). B contains thefungal species predominantly associated with sector 1,and in particular leaves of Memecylon lateriflorum,Caloncoba brevipes, and Uapaca guineensis. C containsthe largest number of species where several specialisedassociations between groups of fungal species andcertain plant substrata can be distinguished. The high-est proportion of specialised associations (66) wasobserved in sectors 1 and 2, while sporadic special-ised relationships involving uncommon or rare fungiwere also detected in the other sectors. The greatestnumber of specialised species was found on Newtoniaduparquetiana, which can mainly be ascribed to theoccurrence of several species of Sporidesmium (Table2). This substratum hosts 21 specialised fungi. Anotherassociation with a particularly high number of specieswas recorded in sector 1 C on Diospyros sanza-minika,and in sector 2 C on Xylopia aethiopica and Allan-blackia floribunda. Only 16 fungal species with ubiqui-tous behaviour appeared in sectors D and E. In D theywere particularly abundant in substratum groups 1 and2, whereas the species in sector E showed a more uni-form pattern and were also present in 3, 4, 6 (Table 2).

Investigations were carried out to find a possiblecorrelation between the fungal species present on one

Microfungi in tropical forest litter 326

Table 1. List of substrata with symbols, year of collection, and voucher specimen reference numbers.

Year Leaf species Family Matrix symbols Voucher specimens

1992 Anthonotha fragrans Leguminosae aa ROHB 405

1992 Berlinia occidentalis Leguminosae by ROHB 406

1992 Dialium aubrevillei Leguminosae p ROHB 407

1992 Dichapetalum toxicarium Chailletiaceae m ROHB 408

1992 Diospyros cooperi Ebenaceae n ROHB 409

1992 Diospyros gabunensis Ebenaceae ae ROHB 410

1992 Duboscia viridiflora Tiliaceae bl ROHB 411

1992 Ficus vogeliana Moraceae u ROHB 412

1992 Gilbertiodendron limba Leguminose aw ROHB 413

1992 Memecylon lateriflorum Melastomataceae r ROHB 414

1992 Neuropeltis prevosteoides Convolvulaceae ao ROHB 415

1992 Scytopetalum tieghemii Scytopetalaceae o ROHB 416

1992 Tarrietia utilis Sterculiaceae am ROHB 417

1992 Terminalia superba Combretaceae j ROHB 418

1992 Trichoscypha chevalieri Anacardiaceae bn ROHB 419

1992 Uapaca esculenta Euphorbiaceae av ROHB 420

1992 Uvaria angolensis Annonaceae x ROHB 421

1992 Xylopia acutiflora Annonaceae ah ROHB 422

1993 Alchornea cordifolia Euphorbiaceae bu ROHB 423

1993 Bridelia grandis Euphorbiaceae t ROHB 424

1993 Cleistopholis patens Annonaceae ak ROHB 425

1993 Didelotia idae Leguminosae bg ROHB 426

1993 Diospyros mannii Ebenaceae bd ROHB 427

1993 Drypetes aylmeri Euphorbiaceae a ROHB 428

1993 Grewia barombiensis Tiliaceae s ROHB 429

1993 Harungana madagascariensis Hypericaceae bx ROHB 430

1993 Hypselodelphys violacea Marantaceae ad ROHB 431

1993 Landolphia owariensis Apocynaceae ap ROHB 432

1993 Memecylon afzelii Melastomataceae d ROHB 433

1993 Memecylon donianum Melastomataceae be ROHB 434

1993 Newtonia duparquetiana Leguminosae bt ROHB 435

1993 Sacoglottis gabonensis Humiriaceae aq ROHB 436

1994 Allanblackia floribunda Guttiferae ar ROHB 437

1994 Caloncoba brevipes Flacourtiaceae as ROHB 438

1994 Chrysophyllum taiense Sapotaceae g ROHB 439

1994 Diospyros kamerunensis Ebenaceae bp ROHB 440

1994 Irvingia gabonensis Simaroubaceae an ROHB 441

1994 Lophira alata Dipterocarpaceae at ROHB 442

1994 Lovoa trichilioides Meliaceae au ROHB 443

1994 Macaranga heterophylla Euphorbiaceae z ROHB 444

1994 Piptadeniastrum africanum Leguminosae aj ROHB 445

1994 Santalodes afzelii Connaraceae e ROHB 446

1994 Thaumatococcus daniellii Scitamineae bv ROHB 447

1994 Trichoscypha arborea Anacardiaceae al ROHB 448

1994 Xylopia aethiopica Annonaceae ac ROHB 449

1995 Bambusa vulgaris Gramineae bw ROHB 450

1995 Beilschmiedia mannii Lauraceae bi ROHB 451

1995 Calpocalyx aubrevillei Leguminosae l ROHB 452

1995 Calpocalyx brevibracteatus Leguminosae c ROHB 453

1995 Canthium rubens Rubiaceae ai ROHB 454

1995 Combretum dolichopetalum Combretaceae ag ROHB 455

1995 Corynanthe pachyceras Rubiaceae q ROHB 456

1995 Coula edulis Olacaceae f ROHB 457

1995 Decorsella paradoxa Urticaceae bf ROHB 458

1995 Dictyophleba leonensis Apocynaceae af ROHB 459

1995 Diospyros sanza-minika Ebenaceae i ROHB 460

1995 Ficus sagittifolia Moraceae br ROHB 461

1995 Guarea thompsonii Meliaceae bh ROHB 462

1995 Hannoa klaineana Simaroubaceae h ROHB 463

1995 Homalium aylmeri Flacourtiaceae az ROHB 464

1995 Landolphia hirsuta Apocynaceae bs ROHB 465

1995 Leptoderris cyclocarpa Leguminosae w ROHB 466

1995 Manniophyton fulvum Euphorbiaceae b ROHB 467

1995 Newtonia aubrevillei Leguminosae ay ROHB 468

1995 Parinarium excelsum Rosacea y ROHB 469

1995 Pentaclethra macrophylla Leguminosae bc ROHB 470

1995 Pseudospondias microcarpa Anacardiaceae bm ROHB 471

1995 Strephonema pseudo-cola Combretaceae ab ROHB 472

1995 Symphonia globulifera Guttiferae ba ROHB 473

1995 Tetracera potatoria Dilleniaceae k ROHB 474

1995 Uapaca guineensis Euphorbiaceae v ROHB 475

A. Rambelli, B. Mulas and M. Pasqualetti 327

Table 2. Presence of 184 fungal species on 71 substrata; the table is ordered by the block clustering method; principal groups of fungal species and substrata are separated.

Matrix groups Group 1 Group 2 Group 3 Group 4 Group 5 Group 6Symbols of matrices bt a bg aw aq r as v i ac ar bm bi br af j ak bl aa c g b p k bd bc ay d at f q be z ae w an y l m ao ad s ai h bn az e aj bu by am bf ah bx bw bs al bp ag av o ap n ba ab u x t bh au bvPseudobotrytis terrestris X X X X X X X X X X X X X X X X Group A 16Chaetosphaeria vermicularioides X X X X X X X X X X X X X X 14Chalara alabamensis X X X X X X X X X X X X X X X 15Cryptophiale kakombensis X X X X X X X X X X X X X X 14Cryptophiale udagawae X X X X X X X X X X X X 12Dictyochaeta assamica X X X X X X X X X X 10Phaeoramularia hachijoensis X X X X X X X 7Corynespora elaeidicola X X X X 4Sporidesmium parvum X X X X 4Brooksia tropicalis X X X X X X X 7Kylindria keitae X X X X X X X 7Hansfordia pulvinata X X X 3Pseudocochliobolus eragrostidis X X X X X 5Beltrania onirica X X X X X X X X X 9Sporidesmium sp. 1 X X X X 4Periconia minutissima X X X X X 5Melanopsammella chloroconica X X X X X X 6Periconia byssoides X X X X X X 6Zygosporium minus X X X X X 5Idriella tropicalis X X X X X 5Selenosporella curvispora X X X X X X X X X 9Rhinocladiella ellisii X X X X X X X X 8Gyrothrix magica X X X X X X X X X X X 11Idriella fertilis X X X X X X X X X X X X X 13Hansfordiellopsis lichenicola X X X X X X X X X X 10Kramasamuha sibika X X X X X X X X X X 10Stachybotrys kampalensis X Group B 1Hansfordiellopsis aburiensis X 1Anungitea fragilis X 1Gyrothrix grisea X 1Solosympodiella clavata X 1Ulocladium consortiale X 1Dictyochaetopsis intermedia X X X X X X X 7Sporidesmium tropicale X X X X 4Sporidesmium ghanaense X X 2Beltrania africana X 1Cordana pauciseptata X 1Alternaria alternata X X 2Kiliophora ubiensis X 1Ardhachandra cristaspora X X X X X X X 7Zanclospora indica X X X X X 5Chaetosphaeria innumera X X X X 4Tripospermum myrti X X X 3Torula herbarum X X 2Idriella lunata X X X 3Calcarisporium acerosum X X X X 4Sporidesmium njalaense X X X X 4Circinotrichum papakurae X X X 3Chaetopsina fulva X X X X 4Speiropsis pedatospora X X X X 4Parasympodiella laxa X X X X X X X 7Arthrinium phaeospermum X X X X X X X 7Stachybotrys parvispora X X X X X X X X X X 10Sporidesmium sp. 5 X Group C 1Sporidesmium jasminicola X 1Minimidochium setosum X 1Helicosporium MCF 1847 X 1Dictyopolyschema sp.1 X 1Acrodictys erecta X 1Deightoniella jabalpurensis X 1Gyrothrix circinata X 1Hyphodiscosia jaipurensis X 1Sporidesmium adscendens X 1Sporidesmium nodipes X 1Sporidesmium sp. 6 X 1Subulispora procurvata X 1Sporidesmium afrormosiae X 1Geotrichum candidum X 1Curvularia ovoidea X 1Chaetendophragmia triangularis var. africana X 1Articulospora foliicola X 1Blastophorum uniseptatum X 1Chalara microspora X 1Dactylaria sp. 1 X 1Pseudospiropes simplex X 1Zygosporium oscheoides X 1Venturia carpophila X X 2Spiropes clavatus X 1Spiropes guareicola X 1Spegazzinia tessarthra X X 2Flosculomyces floridaensis X X 2Phialocephala sp. 1 X X 2Sporidesmium penzigii X 1Bipolaris indica X 1Circinotrichum rigidum X 1Cladosporium chlorocephalum X 1Endophragmia brevis X 1Sporidesmium sp. 2 X 1

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or two substrata and the nature of these substrata.As few data are available on the chemical compositionof the leaf litter, the current morphologically basedbotanical classification was adopted which did notreveal any specific link between fungal species andplant family or genus (Table 2). Nevertheless, the find-ing that certain plants hosted only few fungi is note-worthy. Substrata colonised by not more than threefungal species included the following: Dichapetalumtoxicarium (Dichapetalaceae) which is extremely toxicas it contains fluoroacetic acid capable of destroyingthe tricarboxylic acids of the respiratory cycle ; Neuro-peltis prevosteoides (Convolvulaceae) which is a mega-phanerophyte liana belonging to a family in whichsome species contain hallucinogenic substances;Santalodes afzelii (Connaraceae), another mega-phanerophyte liana belonging to a family comprisingspecies with calcium oxalate crystals in their cell par-enchyma and the seeds and bark of which are extremelypoisonous, although the active compounds are notknown; Hannoa klaineana (Simaroubaceae), a meso-phanerophyte belonging to a family containing specieswith poisonous compounds that are used for theproduction of insecticides; Leptoderris cyclocarpa(Leguminosae), a micro-phanerophyte liana, but theproperties of this plant are unknown; Piptadeniastrumafricanum (Leguminosae), another mega-phanero-phyte ; Trichoscypha chevalieri (Anacardiaceae), amicro-phanerophyte with an edible fruit, but thefamily also contains species having allergenic resinsin the resin canals ; and Diospyros kamerunensis(Ebenaceae), a nano-phanerophyte, Diospyros speciesbeing among the plants most resistant to micro-organisms, with all parts containing poorly knownpoisonous substances that contribute to their durability(Mabberley 1997).

In contrast, certain other substrata were colonised bynumerous fungal species. D. sanza-minika, a meso-phanerophyte that can reach a height of over 20m,unlike the other Diospyros species, was colonised bymany leaf litter saprotrophs (47 fungal species werefound fruiting on incubated litter). Drypetes aylmeri(Euphorbiaceae) is a micro-phanerophyte on which 29microfungal species fruited. In addition, the dead leavesof the following native African meso-phanerophytetrees were colonised by numerous fungal species:Uapaca guineensis (Euphorbiaceae), Xylopia aethiopica(Annonaceae ; Guinea Pepper) which is used as a med-icinal plant, Sacoglottis gabonensis (Humiriaceae), andGilbertiodendron limba (Leguminosae) (Mabberley1997).

The 184 fungal species were grouped according toecological categories (S, C, U), revealing that 39.7%were associated with a single substratum, and 20.1%with two substrata. Therefore, the percentage of fungalspecies that may be regarded as specialised with regardto the substratum was 59.8%. Ubiquitous speciescomprised 12.5%, and common species 27.7%.

Classification of the morphological characters ofthe 184 species showed the following distribution: 70%had pigmented conidia, 59% non-septate conidia,58% produced few conidia, while 21% had phialidic,37% percurrent, and 42% sympodial conidiogenesis.Sterile setae were found in 24% of the species.

MDS was carried out on the ecological and mor-phological characters detected in the 184 species. In thisanalysis the common and ubiquitous species weregrouped into a single category. The MDS plot ofFig. 1 shows two associated groups, with a statisticalsignificance exceeding 90% (chi square test for eachpaired combination of grouped characters, P<0.1).The first group contains the common and ubiquitousspecies that were positively correlated with hyalineconidia, abundant spore production, phialidic coni-diogenesis, and the presence of sterile setae. The secondgroup comprises the specialized species having pig-mented and septate conidia and low spore production.Species with percurrent or sympodial conidiogenesiswere not associated with either group (Fig. 1).

Fungal species occurring on only one (73 species) ortwo substrata (36 species) were analysed (Table 2).Species detected on two substrata did not indicate anyassociations of groups of substrata. It was also evidentthat certain leaf litters selected high numbers ofspecialist fungal species: Newtonia duparquetiana52%, Xylopia aethiopica 30%, Drypetes aylmeri 29%,Corynanthe pachyceras 28%, Allanblackia floribunda26%, Diospyros sanza-minika 26%, Didelotia idae25%, and Uapaca guineensis 17%.

The ecological characters of the substrata hostingover ten fruiting species were compared. Fig. 2a showsthe percentage presence of single categories on leafsubstrata ordered using cluster analysis. Three groupsof substrata could be distinguished, characterisedby: (1) species with numerous ubiquitous fungi fruiting

DIM

EN

SIO

N 2

-+--------------+--------------+--------------+------------2 + | | | |

++

||

|

+||

|

+||

|

+

+

||

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−1 + | | |

−2 + -+--------------+--------------+--------------+------------

−2 2

DIMENSION 1

GED B

H

J

I

L A FCM

−1 0 1

A-Common and ubiquitous species (C-U)B-Specialized species (S)C-Pigmented conidia (PG)D-Hyaline conidia (HC)E-Presence of sterile setae (SS)F-Absence of sterile setae (SA)G-Phialidic conidiogenesis (PC)

H-Percurrent conidiogenesis (PE)I-Sympodial conidiogenesis (SY)J-Septate conidia (SC)K-Unicellular conidia (UC)L-Abundant conidial production (AP)M-Low conidial production (LP)

Fig. 1. MDS plot showing relationships between morpho-

logical and ecological characters of microfungi fruiting onincubated rain forest leaf litter.


Table 3. Yule coefficients of association (Q) and asymptotic standard errors calculated for fungal species present on over ten substrata; the underlined values were significant (P<0.01) in

chi-square tests.

Asterostomella Beltrania Beltraniella Chalara Chloridium Circinotrichum Cladosporium Cryptophiale Cryptophiale Grallomyces Gyrothrixsp.1 rhombica portoricensis alabamensis virescens maculiforme cladosporioides kakombensis udagawae portoricensis magica

Asterostomella sp. 1 -Beltrania rhombica 0.705±0.132 -Beltraniella portoricensis 0.542±0.176 0.743±0.120 -Chalara alabamensis 0.596±0.208 0.664±0.195 0.005±0.291 -Chloridium virescens 0.688±0.185 0.628±0.213 0.088±0.296 0.252±0.319 -Circinotrichum maculiforme 0.435±0.270 0.537±0.255 0.462±0.262 - 0.171±0.406 0.185±0.360 -Cladosporium cladosporioides 0.692±0.153 0.666±0.153 0.669±0.161 0.592±0.273 0.329±0.316 - 0.026±0.336 -Cryptophiale kakombensis 0.810±0.139 0.770±0.165 0.261±0.281 0.719±0.158 0.642±0.194 - 0.121±0.414 0.782±0.208 -Cryptophiale udagawae 0.890±0.112 0.886±0.134 0.628±0.217 0.798±0.127 0.729±0.162 - 0.432±0.446 0.735±0.248 0.990±0.012 -Grallomyces portoricensis 0.507±0.228 0.575±0.217 0.659±0.181 - 0.095±0.356 0.398±0.274 0.324±0.310 0.624±0.246 0.398±0.274 0.324±0.320 -Gyrothrix magica 0.726±0.195 0.481±0.289 - 0.024±0.329 0.436±0.288 0.251±0.354 0.360±0.334 0.416±0.342 0.481±0.275 0.370±0.334 0.391±0.299 -Idriella fertilis 0.650±0.205 0.743±0.182 0.357±0.274 0.500±0.250 0.546±0.237 0.471±0.277 0.760±0.207 0.546±0.237 0.241±0.354 0.260±0.318 0.802±0.130Chaetosphaeria vermicularioides 0.251±0.240 0.257±0.241 - 0.089±0.253 0.209±0.288 0.092±0.310 0.424±0.265 0.514±0.230 0.277±0.283 - 0.212±0.344 0.775±0.126 0.104±0.339Periconia cookei - 0.055±0.246 0.175±0.241 - 0.132±0.244 0.091±0.296 - 0.220±0.310 0.322±0.287 0.339±0.247 - 0.024±0.311 - 0.548±0.286 0.345±0.254 0.070±0.340Pestalotia sp. 1 0.719±0.135 0.642±0.155 0.610±0.168 0.680±0.217 0.648±0.234 0.346±0.316 0.880±0.071 0.832±0.165 0.793±0.200 0.862±0.138 0.525±0.298Pseudobotrytis terrestris 0.636±0.190 0.697±0.179 0.091±0.282 0.143±0.327 0.910±0.063 0.324±0.310 0.425±0.287 0.558±0.221 0.514±0.248 0.290±0.291 0.391±0.299Rhinocladiella selenoides 0.132±0.274 0.471±0.233 0.169±0.271 - 0.143±0.350 0.516±0.233 0.278±0.318 - 0.086±0.291 0.150±0.328 0.034±0.366 0.673±0.179 0.104±0.377Scolecobasidium constrictum 0.635±0.177 0.559±0.203 0.304±0.246 0.200±0.300 0.261±0.297 0.692±0.165 0.704±0.202 0.435±0.254 0.189±0.329 0.790±0.119 0.625±0.208Scolecobasidium tshawytschae 0.663±0.174 0.726±0.164 0.581±0.199 0.463±0.245 - 0.088±0.356 0.835±0.108 0.446±0.272 0.351±0.283 0.278±0.318 0.412±0.255 - 0.200±0.402Selenosporella sp. 1 0.228±0.285 0.628±0.213 0.271±0.281 0.011±0.364 0.066±0.365 0.420±0.290 0.556±0.281 0.076±0.375 0.121±0.414 0.782±0.129 - 0.059±0.421Zygosporium echinosporum 0.128±0.241 0.229±0.234 0.173±0.238 0.223±0.279 0.125±0.298 0.745±0.161 0.366±0.239 0.455±0.241 0.091±0.320 0.686±0.163 - 0.042±0.340Zygosporium gibbum 0.298±0.220 0.641±0.152 0.446±0.197 0.208±0.279 0.598±0.209 0.498±0.264 0.739±0.142 0.298±0.209 0.489±0.254 0.680±0.163 0.222±0.313Zygosporium masonii 0.316±0.234 0.561±0.192 0.102±0.254 0.410±0.249 0.607±0.196 0.608±0.207 0.481±0.241 0.316±0.277 0.065±0.336 0.713±0.150 0.356±0.292

Idriella Chaetosphaeria Periconia Pestalotia Pseudobotrytis Rhinocladiella Scolecobasidium Scolecobasidium Selenosporella Zygosporium Zygosporiumfertilis vermicularioides cookei sp.1 terrestris selenoides constrictum tshawytschae sp.1 echinosporum gibbum

Idriella fertilis -Chaetosphaeria vermicularioides 0.348±0.275 -Periconia cookei 0.293±0.293 0.726±0.131 -Pestalotia sp. 1 0.613±0.254 0.240±0.254 0.244±0.245 -Pseudobotrytis terrestris 0.455±0.272 - 0.034±0.305 0.024±0.294 0.541±0.246 -Rhinocladiella selenoides 0.212±0.324 0.249±0.270 0.126±0.280 0.245±0.282 0.412±0.256 -Scolecobasidium constrictum 0.831±0.108 0.729±0.137 0.157±0.267 0.497±0.237 - 0.059±0.325 0.265±0.279 -Scolecobasidium tshawytschae 0.697±0.169 0.396±0.242 0.280±0.260 0.577±0.231 0.239±0.297 0.186±0.302 0.553±0.205 -Selenosporella sp. 1 0.546±0.237 0.789±0.127 0.486±0.233 0.257±0.304 0.200±0.324 0.834±0.103 0.696±0.164 0.649±0.185 -Zygosporium echinosporum 0.378±0.266 0.412±0.216 0.378±0.218 0.523±0.201 - 0.014±0.293 0.513±0.211 0.392±0.231 0.724±0.146 0.455±0.241 -Zygosporium gibbum 0.690±0.185 0.547±0.186 0.034±0.247 0.670±0.153 0.429±0.239 0.335±0.248 0.771±0.129 0.606±0.190 0.725±0.167 0.700±0.137 -Zygosporium masonii 0.540±0.224 0.772±0.116 0.500±0.200 0.333±0.248 0.008±0.307 0.556±0.199 0.756±0.127 0.560±0.199 0.718±0.155 0.860±0.081 0.770±0.210

A.Rambelli,

B.MulasandM.Pasqualetti

331

(b)

(c)

(a)

S

U

0%

20%

40%

60%

80%

100%

S

C

U

0%

20%

40%

60%

80%

100%

PG

0%

20%

40%

60%

80%

UCSC

100%

HC

Fig. 2. (Cont.)


(d)

(e)

0%

20%

40%

60%

80%

100%

LPAP

0%

20%

40%

60%

80%

100%SYPEPC

( f )

0%

20%

40%

60%

80%

100%SASS

Fig. 2. (a) Percentages of single (S), common (C) and ubiquitous (U) colonizers on substrata hosting over ten fungal species ;the substrata are ordered by cluster analysis. Sporulation characters are coded as follows: (b) percentage of species withhyaline (HC) and pigmented (PG) conidia on substrata hosting over ten fungal species ; (c) percentage of species with

unicellular (UC) and septate (SC) conidia on substrata hosting over ten fungal species ; (d ) percentage of species with low(LP) and abundant conidial production (AP) on substrata hosting over ten fungal species ; (e) percentage of species withphialidic (PC), percurrent (PE) and sympodial (SY) conidiogenesis on substrata hosting over ten fungal species ; and ( f )percentage of species with (SS) and without sterile setae (SA) on substrata hosting over ten fungal species.


and few unique or common fungal species (from Lovoatrichilioides – au, to Calpocalyx brevibracteatus – c);(2) species with many common and ubiquitous fungifruiting (from Memecylon lateriflorum – r, to Land-olphia hirsuta – bs) ; and (3) species with a significantnumber of specialist species fruiting (from Xylopiaaethiopica – ac, to Newtonia duparquetiana – bt).Fungal species that occurred only once were morenumerous than common and ubiquitous species inNewtonia duparquetiana (bt).

Fungal species fruiting on more than ten substratawere analysed (Q association index) in order to detectsignificant association between them (Table 3). Regularassociations were recorded between Cryptophialeudagawae and C. kakombensis (0.99), and betweenChaetosphaeria vermicularioides and Pseudobotrytisterrestris (0.91), that were practically always associatedwith the same substrata. Equally high and significantassociations (Q>0.8) were found between the above-mentioned two species of Cryptophiale and Astero-stomella sp. 1, and also between Beltrania rhombicaand C. udagawae, Circinotrichum maculiforme andScolecobasidium tshawytschae, Cladosporium clado-sporioides and Pestalotia sp. 1, C. kakombensis andPestalotia sp. 1, Grallomyces portoricensis and Pestalo-tia sp. 1, Gyrothrix magica and Idriella fertilis, I. fertilisand Scolecobasidium constrictum, Rhinocladiella sele-noides and Selenosporella sp. 1, and Zygosporium echi-nosporum and Z. masonii. Moreover, Z. echinosporum,Z. masonii and Z. gibbum showed a significant corre-lation index of over 0.7.

Morphological characters of certain species occur-ring on substrata hosting over ten colonizers wereanalysed. In Fig. 2b conidial pigmentation was takenas the criterion, and the percentage of species withpigmented conidia was calculated for each leaf typewas compared with the mean percentage value ofall fungal species. In some substrata, the distributiondiverged greatly from the mean value (70% ofpigmented conidia). Over 80% of the fungi fruitingon litter of Xylopia aethiopica, Allanblackia floribunda,and Symphonia globulifera had pigmented spores ; inparticular, S. globulifera did not host any species withhyaline conidia. In contrast, less than 60% of thefruiting fungi had pigmented conidia on Calpocalyxbrevibracteatus, Chrysophyllum taiense, Tetracerapotatoria, Memecylon lateriflorum, Anthonotha fra-grans, Sacoglottis gabonensis, Lophira alata, Diospyrosmannii,Memecylon donianum, and Decorsella paradoxa(Fig. 2b).

Furthermore, in some substrata the distributionof fungi with septate conidia deviated from themean distribution (41%). In particular, Newtoniaduparquetiana hosted over 60% of fruiting specieswith septate conidia, whereas Calpocalyx brevi-bracteatus, Tetracera potatoria, Lophira alata, Dios-pyros mannii, Guarea thompsonii, and Landolphiahirsuta had over 80% of fruiting fungal species withone celled conidia (Fig. 2c). Finally, abundant conidial

production (mean value 42%) exceeded 60% inCoula edulis, Memecylon lateriflorum, Lophira alata,Newtonia aubrevillei, and Guarea thompsonii, while itwas below 30% in Lovoa trichilioides and Newtoniaduparquetiana (Fig. 2d ).

Fig. 2e shows the percentages of the various types ofconidiogenous cells. Phialidic conidiogenous cells oc-curred in about 20% the 184 species; in litter of manyspecies hosting more than ten fruiting species, thisnumber deviated greatly from this average. In particu-lar, litter of Manniophyton fulvum, Calpocalyx brevi-bracteatus, Chrysophyllum taiense, Dialium aubrevillei,Lophira alata, Lovoa trichilioides, Newtonia aubrevillei,Diospyros mannii, and Guarea thompsonii hosted over40% of phialidic-fruiting species, while Symphoniaglobulifera and Ficus sagittifolia had less than 10%.Fungal species with sympodial conidiogenesis (meanvalues 42%) often exceeded 50%, such as in Tetracerapotatoria, Diospyros cooperi, Corynanthe pachyceras,Memecylon lateriflorum, Xylopia aethiopica, Symphoniaglobulifera, Beilschmiedia mannii, and Ficus sagittifolia,while it was below 30% in Manniophyton fulvum andNewtonia duparquetiana. In particular, N. aubrevilleiand Ficus sagittifolia did not host any fruiting fungiwith percurrent conidiogenesis, and over 90% of thespecies had sympodial conidiogenesis in the latter(Fig. 2e). Plant species hosting fewer than 20% fruitingfungi with percurrent conidiogenesis (average 37%)included: Manniophyton fulvum, Calpocalyx brevi-bracteatus, Coula edulis, Chrysophyllum taiense, Dios-pyros sanza-minika, Tetracera potatoria, Diospyroscooperi, Corynanthe pachyceras, Lophira alata, Lovoatrichilioides, Diospyros mannii, and Beilschmiedia man-nii. In contrast, N. duparquetiana had more than 60%of the fruiting species with percurrent conidiogenesis(Fig. 2e).

Finally, some plant substrata were found wherethe presence of sterile setae in the fruiting structuresexceeded the mean value (24%). Presence of setaeexceeded 40% of the fruiting species in Calpocalyxbrevibracteatus, Chrysophyllum taiense, Tetracerapotatoria, Anthonotha fragrans, Lophira alata,Memecylon donianum, Ficus sagittifolia, and Land-olphia hirsuta (Fig. 2 f ).

DISCUSSION

The study focuses on relationships between plant hostand associated saprotrophic fungal communities, andbuilds on other comparative investigations on micro-fungi in the tropical environments of Tai National Park(Rambelli et al. 1983, 1984, 1991). Microfungal speciesthat fruited on incubated leaf litter samples wereidentified and characterised, and the characteristics ofthe fungal communities were then correlated with thehost plants. The detection on specialized plant-fungusspecies relationships is noteworthy as it confirms theexistence in tropical forest ecosystems of the previouslyobserved saprotrophic specialisation (Lodge & Cantrell


1995). In totally different environmental conditions,fungal communities supporting major environmentalstress were observed in the Mediterranean maquis(Mulas et al. 1995, Pasqualetti et al. 1999). These pre-liminary observations show that saprotrophic special-isation is not linked to particular environments andsubstrata, but is a natural phenomenon occurring todifferent degrees in all environments. It can be assumedthat in environments where conditions are optimal formicrofungal development (humidity and temperature),such as in tropical forests, saprotrophic specialisation ismainly related to nutritional factors and the secondarychemistry of the substrata.

The data presented were analysed for the distri-bution of 184 saprotrophic fungi on litter of 71 plantspecies. This enabled us to observe significant corre-lations between certain morphological and ecologicalparameters of hosts and fungal colonizers. The dataobtained revealed that ubiquitous-common and special-ist fungi have quite distinct characteristics, except forpercurrent and sympodial conidiogenesis which occursin both groups (Fig. 1).

Specialist species show limited production of oftenmulticellular, resistant, pigmented and scarcely vulner-able conidia. This may imply that these species aregreatly involved in vegetative propagation and are themain driving force behind the degradation of the sub-strata. In fact, it seems reasonable to suggest that ifthere is a high degree of fungal sepcialisation relatedto the substrata, the microfungi involved may haveadopted vital strategies whereby energy is mostly con-sumed for vegetative growth and not for reproduction.These species need not compete for the substratum andtheir highly resistant conidia permit colonisation ofadjacent similar substrata, not in competition withother fungi but as selected by the substratum. Thecommon or ubiquitous species showed a differentbehaviour with divergent morphological and ecologicalcharacters. These species may compensate for theirlimited nutritional specialisation by a greater sub-stratum colonisation capacity, as this colonisation mustbe achieved within a short time because of the vulner-ability of the hyaline ephemeral conidia. From anutritional point of view, the specialist species may actas primary colonisers capable of attacking the morerecalcitrant substrata and thus pave the way for sec-ondary colonisers.

ACKNOWLEDGEMENTS

We thankWalter Gams for critically reading the manuscript, Laurent

Ake Assi for identification of plant matrices, and the Italian Embassy

in Abidjan for help and assistance.

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Comparative studies on microfungi in tropical ecosystems in Ivory Coast forest litter: behaviour on different substrata