ORIGINAL ARTICLE

Coltriciella sonorensis sp. nov. (Basidiomycota, Hymenochaetales)from Mexico: evidence from morphologyand DNA sequence data.

Ricardo Valenzuela & Tania Raymundo &

Joaquín Cifuentes & Martín Esqueda & Mario Amalfi &Cony Decock

Received: 24 June 2010 /Revised: 10 November 2010 /Accepted: 26 November 2010 /Published online: 19 February 2011# German Mycological Society and Springer 2011

Abstract Coltriciella sonorensis is described here as a newspecies from Mexico. It is characterized by pleuropodal,flabelliform basidiomes, rounded to elongated or daedaloidpores, a well-developed sub-hymenium, and oblong tocylindrical basidiospores, slightly attenuated towards the apex.The specimen was collected on soil in an open Quercus standin mixed Quercus–tropical deciduous forest in the Sierra deÁlamos–Río Cuchujaqui Biosphere Reserve, Sonora, Mexico.From a phylogenetic perspective, the species appears to berelated to C. oblectabilis, also occurring in Mexico.

Keywords Basidiomycota . Hymenochaetales .

Hymenochaetaceae . Polyporoid fungi

Introduction

Coltriciella is a small genus of poroid Hymenochaetales, withabout seven recognized species (http://www.indexfungorum.org/Names/Names.asp.; Aime et al. 2003; Corner 1991;Ryvarden 2004). It includes species with resupinate, pendant,or stipitate basidiomes sharing coloured and ornamentedbasidiospores and a monomitic hyphal system (Corner 1991;Ryvarden 2004). The basidiomes are rather small, and arecommonly found in cryptic habitat such as woody debris, theunderside of fallen trunks, or in trunk cavities (Aime et al.2003); data on Coltriciella are therefore rather scarce.

From the ecological and nutritional strategy point ofview, although Coltriciella basidiomes are commonlyfound on wood (Aime et al. 2003; Corner 1991; Ryvarden2004), they are probably not lignicolous as such. Tedersooet al. (2007a, b) have demonstrated that some species,notably C. dependens s.l. and an unidentified Coltriciellaspecies, form ectomycorrhizal associations with angiospermtrees of various families (Caesalpiniaceae, Dipterocarpa-ceae, and Myrtaceae).

Coltriciella is closely related to Coltricia, whatever theperspective: morphologically, these genera differ mainly inhaving either ornamented or smooth basidiospores, all theother features (including the hyphal system) being shared(Ryvarden 1991, 2004); both genera contain species thatform mycorrhizal associations (Tedersoo et al. 2007a, b);phylogenetically, the two genera are resolved as sisterclades, in a lineage basal to the bulk of the poroidHymenochaetales (Larsson et al. 2006; Wagner and Fischer2002), but, according to Larsson et al. (2006), phylogeneticanalyses do not clearly support separation into two genera.

In Mexico, two species have been reported viz. C.dependens (Vásquez and Guzmán-Dávalos 1991; Romero-

R. Valenzuela : T. RaymundoLaboratorio de Micología, Departamento de Botánica,Escuela Nacional de Ciencias Biológicas, IPN,Apartado Postal 256, Centro Operativo Naranjo,Col. Santa María de la Rivera,México, D.F. C.P. 02600, México

J. CifuentesHerbario FCME, Facultad de Ciencias, UNAM,Apartado Postal 70–399, Coyoacán,México, D.F. C.P. 04510, México

M. EsquedaCentro de Investigación en Alimentación y Desarrollo, A.C.,Apartado Postal 1735, Hermosillo,Sonora C.P. 83000, México

M. Amalfi : C. Decock (*)Mycothèque de l’Université catholique de Louvain (MUCL,BCCM™), Earth and Life Institute – Mycology,Université catholique de Louvain,Croix du Sud 3,1348 Louvain-la-Neuve, Belgiume-mail: cony.decock@uclouvain.be

Mycol Progress (2012) 11:181–189DOI 10.1007/s11557-011-0740-7

Bautista et al. 2010), and C. oblectabilis (Raymundo andValenzuela 2003; as Coltricia pseudocinnamomea Burds.).

In the framework of a continuous survey of the poroidHymenochaetales occurring locally (Raymundo andValenzuela 2003; Raymundo et al. 2008, 2009; Rodríguez-Alcantar et al. 1996; Valenzuela et al. 1996, 2005, 2007), acollection of a Coltriciella species was found that could notbe attributed satisfactorily to any of the species described. Itsphylogenetic relationships, inferred from DNA sequencedata of the 5′ end of the nuclear ribosomal LSU (Larsson etal. 2006; Wagner and Fischer 2002) also show that thiscollection is distantly related to Coltriciella species for whichDNA sequence data are available (Larsson et al. 2006;Wagner and Fischer 2002, Tedersoo et al. 2007a, b). Wetherefore conclude that it represents an undescribed speciesthat we propose as Coltriciella sonorensis sp. nov.

Materials and methods

Collection localities The specimen examined was collectedin the Sierra de Álamos-Río Cuchujaqui Biosphere Reserve,Sonora State. The local ecosystem consists of a transition areabetween tropical deciduous forest and the Sierra Madreevergreen pine–oak forest (Rzedowski 2006). Locally, Quer-cus oblongifolia Torr. is the dominant species, and is mixedwith some elements of the tropical caducifolious forest.

Morphology and anatomy Morphological examinationswere conducted using protocols outlined by Ryvarden(1991) and Cifuentes et al. (1986). Basidiome colours aredescribed according to Kornerup and Wanscher (1981).Measurements of anatomical characteristics were takenfrom rehydrated tissues in 5% aqueous KOH, whileamyloid/dextrinoid reactions were taken with Melzer’sreagent. For the SEM studies, the sample was preparedaccording to the critical-point-drying method outlined inMoreno et al. (1995) and examined with a Zeiss DSM-950scanning electron microscope. Voucher specimens aredeposited in ENCB with duplicates in MUCL and CESUES[Herbarium acronyms are according to Thiers 2011;continuously updated).

Sequencing DNA extraction, amplification, and sequencingof the nuclear ribosomal 5′ end of the LSU are as describedin Decock et al. (2007). The primers LROR and LR5 wereused for PCR amplifications. Successful PCR reactionsresulted in a single band observed on an 0.8% agarose gel,corresponding to approximately 900 bp. Sequencing reac-tions were performed using CEQ DTCS Quick Start Kit®

(Beckman Coulter), according to the manufacturer’s rec-ommendations, with the primers LROR, LR3, LR3R, LR5(http://biology.duke.edu/fungi/mycolab/primers.htm).

Phylogenetic analysis Thirty-four specimens and culturesrepresenting 26 taxa were included in the nuc-LSU-basedphylogenetic analysis. The materials and sequences used inthis study are listed in Table 1. Nucleotide sequences wereautomatically aligned with Clustal X (version 2.0.1;Thompson et al. 1997) and manually adjusted as necessarywith the text editor in PAUP* (version 4.0b10). Trichaptumabietinum AY059063 and Pyrrhoderma scaurumAY059030 were used as the outgroup (Larsson et al.2006; Tedersoo et al. 2007a). Alignment is deposited atTreeBASE (http://www.treebase.org/treebase/index.html).

Phylogenetic analyses were performed using maximumparsimony (MP) as implemented in PAUP* version 4.0b10(Swofford 2003) and Bayesian inference (BI) as imple-mented in MrBayes v3.1.2 (Huelsenbeck and Ronquist2001). Evolution models for Bayesian inference wereestimated using the AIC (Akaike Information Criterion) asimplemented in Modeltest 3.7 (Posada and Crandall 1998).Bayesian analyses were implemented with two independentruns, each with four simultaneous independent chains forfour million generations, starting from random trees, andkeeping one tree every 1,000th generation. All treessampled after convergence (average standard deviation ofsplit frequencies <0.01 and confirmed using Tracer v1.4;Rambaut and Drummond 2007) were used to reconstruct a50% majority-rule consensus tree (BC) and to estimateposterior probabilities. The posterior probability (BPP) ofeach node was estimated based on the frequency at which thenode was resolved among the trees sampled with theconsensus option of 50% majority-rule (Simmons et al.2004). A BPP above 0.95 was considered a significant value.

For MP analyses, gaps were treated as missing data. Themost parsimonious trees (MPT) for the nucLSU data wereidentified, using heuristic searches with 1,000 randomaddition sequences, evaluated further by bootstrap analysis,retaining clades compatible with the 50% majority-rule inthe bootstrap consensus tree. Analysis conditions were treebisection addition branch swapping (tbr), starting treeobtained via stepwise addition, steepest descent not ineffect, MulTrees effective. A bootstrap support value (BS)above 70% was considered significant. The final alignmentcomprised 919 positions including gap.

Results

Phylogenetic analysis

Within Coltriciella, the length of the LSU fragmentanalyzed ranged from 865 (C. pusilla AY059060) to 869(C. sonorensis) bp. The final alignment of the 34sequences resulted in 905 positions including gaps, of

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which 257 were variable and 174 characters parsimonyinformative. The best-fit model for the nuc LSU data wasGTR+I+G with unequal base frequencies (A=0.2632, C=0.1861, G=0.2928, T=0.2579), a gamma distributionshape parameter of 1.5922, and a proportion of invariablesites of 0.5749.

The MP analysis produced 183 most parsimonious trees(522 steps in length, consistency index 0.600, retentionindex 0.745, and rescaled consistency index 0.447). Thestrict consensus of the 183 most parsimonious trees and theBC tree were mostly congruent. The Bayesian consensustree is represented in Fig 1.

The general topology of the BC tree, especially thecomposition and relationships of the different clades withinColtriciella, are in accordance with Tedersoo et al. (2007b).The Coltriciella clade is well supported by the Bayesiananalysis (BPP 1.0). With regard to our collection, theseanalyses resolved it as a distinct terminal branch (Fig. 1),related to C. oblectabilis, both forming sister clades (BS=70%, BPP=1.0). The C. oblectabilis sequence and ourcollection differ in 13 positions.

The analysis also revealed that sequences referred to asC. dependens do not form a monophyletic clade, as alreadyhighlighted by Tedersoo et al. (2007b); none is identical to

Table 1 List of species, collections, and sequences used in the phylogenetic analyses

Species name Specimen reference Origin Accession number

Coltricia cinnamomea (Jacq.) Murrill Dai 2464 Finland AF311003

Coltricia confluens P.-J. Keizer TAA 181460 Estonia AM412241

Coltricia montagnei (Fr.) Murrill 96-96 USA AY039683

Coltricia cf. oblectans (Berk.) G. Cunn. TAA 195036 Seychelles AM412245

TU 103621 Seychelles AM412246

Coltricia perennis (L.) Murrill DSH 93-198 ND AF287854

92-96 Germany AF311004

CH08-526 China HQ534102

GEL3915 ND AJ406472

Coltriciella dependens (Berk. & M.A. Curtis) Murrill LR 39016 Puerto Rico AY059059

TU103378 China AM412252

TU103611 Seychelles AM412253

TAA195099 Seychelles AM412254

TU100518 Australia AM412251

MEL2096512 Australia AM412248

MEL2292349 Australia AM412249

TU100506 Australia AM412250

Coltriciella oblectabilis (Lloyd) Kotl., Pouzar & Ryvarden RC 11835 USA AY059061

Coltriciella navispora T.W. Henkel, Aime & Ryvarden C. Aime Guyana AY059062

Coltriciella pusilla (Imazeki & Kobayasi) J.E. Wright M. Núñez 26.7.95 Japan AY059060

Coltriciella sonorensis R. Valenz., Esqueda & Decock (T) ENCB RV13144 Mexico HQ439179

Coltriciella tasmanica (Cleland & Rodway) D.A. Reid TAA 159956 Australia AM412247

Hyphodontia quercina (Pers.) J. Erikss. GEL 3790 ND AY059065

Hyphodontia radula (Pers.) Langer & Vesterh. GEL 3798 ND AJ406466

Hyphodontia sambuci (Pers.) J. Erikss. GEL 2414 ND AJ406461

Pyrrhoderma scaurum (Lloyd) Ryvarden Dai 2272 ND AY059030

Schizopora paradoxa (Schrad.) Donk TW 4.6.98b ND AY059067

Trichaptum abietinum (Dicks.) Ryvarden TW 172 ND AY059063

Uncultured ectomycorrhizal fungus L2367 Seychelles AM412255

L2528_C.dep Seychelles AM412256

L2410_C.obl Seychelles AM412257

L2368_Coltr Seychelles AM412258

L2450_Coltr Seychelles AM412259

L2480_Coltr2 Seychelles AM412260

T, PT = type, paratype.

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the “reference” sequence AY059059 used in Larsson et al.(2006) and Wagner and Fischer (2002) and are scattered inseveral clades. The closest sequence (AM412254, originat-ing from the Seychelles) differs in 8 positions fromAY059059, suggesting the existence of species diversitylarger than presumed.

Taxonomy

Coltriciella sonorensis R. Valenz., Esqueda & Decock sp. nov.Figs. 2 and 3

MycoBank: MB 515460Basidiomata annua, stipitata, flabelliformia. Pileus 8–

12 mm latus, orbicularis vel reniformis, flavo-brunneus velrubro-brunneus, zonatus. Hymenophorum porosum, poriscircularibus vel elongatis, 2–3 per mm, daedaloideis inaliquot partibus hymenophori. Stipites laterales, brunnei.Systema hypharum monomiticum, hyphis generatoriis sep-tis simplicibus. Basidiosporae 8–10.5×4.0–5.0 μm, oblon-gae vel cylindricae, in proportione minore naviculares,pallide flavo-brunneae vel rubro-brunneae, nonamyloideae,verrucosae, crassitunicatae.

Fig. 1 The 50% majority ruleconsensus tree from Bayesianinference based on nucLSUsequences. Thickened branchesin boldface indicate bootstrapsupport greater than 70% andBayesian posterior probabilitygreater than 0.95. Bayesianposterior probabilities andParsimony bootstrap supportvalues are shown only whennot concordant

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Holotypus MEXICO, SONORA STATE: Municipium Ála-mos, La Cañita, 26º 59′ 32.6″N, 108º 38′ 59.52″W, 657 masl., ad terram in sylvis mixtis Quercus et caducifolioussylvis, 15 sep 2006, R. Valenzuela 13144, ENCB, IsotypusCESUES et MUCL.

Description Basidiome annual, stipitate, pleuropodal, withan overall coriaceous texture; stipe lateral, cylindrical,tapering to the apex, 4–8 mm long×1 mm diam, lightbrown (6D8) to brown (7D8, 7E8), velutinate, with agranulate base consisting of organic matter, soil and

mycelium; pilei flabelliform, semicircular to kidney-shaped, projecting upwards 8–12 mm, 15–20 mm wide,up to up to 3 mm thick, in brown shade [(6–7(D8–F8, up to8 F8), light brown, rust brown, reddish brown to darkbrown], with darker concentric lines (6 F4,3), finelyappressed velutinate to slightly fibrillose, smooth in parts,concentrically zonate; margin acute, slightly velutinate,fimbriate to wavy; pore surface brown (6E8,7D8–E8) toreddish brown (8E8), the edges darker; pores rounded toelongate, daedaloid in some parts, entire, 2–3 per mm(radially measured), with thick dissepiments; tubes up to

Fig. 2 Coltriciella sonorensis. 2Basidiospores. 3 Basidia. 4Subhymenium. 5 Hymenophoraltrama. 6 Pileipellis. 7 Trama ofthe pileus context

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2 mm deep, concolorous with the pores surface; contextvery thin, less than 1 mm thick, homogeneous, brown (6E8,7D8), fibrous.

Hyphal system monomitic; generative hyphae simple-septate, simple to branched, with many bifurcate hyphae,thin- to thick-walled, golden yellow to reddish brown inKOH, negative in Melzer's reagent, hyphae of the hyme-

nophoral trama 3.2–6.4 μm diam., slightly interwoven,golden yellow to yellowish brown in KOH, simple tobifurcated, thick-walled; contextual hyphae slightly inter-woven, yellowish brown to dark brown in KOH, simple tobranched, mainly bifurcate, thick-walled, 4–7 μm diam;pileipellis a cutis, up to 50 μm thick, hyphae 3–5 μm diam,prostrate to semi-erect, sub-parallel to slightly interwoven,

Fig. 3 Coltriciella sonorensis. 8 Basidiome. 9 Pileus surface. 10 Context. 11 Subhymenium.;12 Basidiospores, 13 Hymenophoral trama. 14Scanning electron microscope image of Basidiospores

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hyaline, yellowish to yellowish brown in KOH, simple toscarcely branched, some bifurcated, thick-walled; stipehyphae parallel to sub-parallel, yellowish brown to reddishbrown in KOH, simple to scarcely branched, somebifurcated, thick-walled, 4–7 μm diam; stipitipellis atrichoderm, up to 240 μm thick, hyphae 4–5.6 μm diam.,erect, semi-erect to prostrate, laxly interwoven, yellowishbrown to brown in KOH, branched, some bifurcate, thick-walled; subhymenium very developed, up to 32 μm thick,ramose to ramose-inflated, with two layers, one near thetrama, laxly, hyaline, with bifurcated hyphae, 4–8 μm diam,hyaline to yellowish in KOH, thin- to thick-walled; anothernear the hymenial layer, compacted, brown, with inflatedhyphae and cells, bifurcated, 6–8 μm diam, pale brown toyellowish brown in KOH, thin to thick-walled; basidiaclavate, hyaline in KOH, base simple septate, 16–24×6–10 μm, with 4 sterigmata, 4–6 μm long; basidiosporesoblong to cylindrical, somewhat attenuated towards theapex, some navicular, pale yellowish brown, rust brown toreddish brown in KOH, inamyloid, finely verrucose, thick-walled, uniguttulate, 8.0–10.5×4–5 μm; cystidia and othersterile hymenial elements absent.

Ecology, range, and distribution Solitary, growing on soilin open areas, in the vicinity of living Quercus oblongifoliatrees, in Q. oblongifolia dominant areas, mixed with sometropical caducifolious elements. Know only from the typelocality in Sonora.

Specimen examined MEXICO, SONORA STATE: Muni<cipality of Álamos, La Cañita, 26º59′32.6″N, 108º38′59.52″W, 660 m asl, on soil in open areas near Quercusoblongifolia., in oak forest mixed with elements of thecaducifolious tropical forest, 15 Sep 2006, R. Valenzuela13144, Holotype (ENCB), Isotype (CESUES, MUCL).

Discussion

The small, pleuropodal basisiome with flabelliform, semicir-cular to kidney-shaped, zonated pilei, the irregular, rounded todaedaloid pores, the well-developed subhymenium, and thebasidiospores features characterize this species. It can becompared to C. oblectabilis and C. dependens, both occur-ring also in Mexico (Raymundo and Valenzuela 2003;Vásquez and Guzmán-Dávalos 1991). Coltriciella oblecta-bilis is phylogenetically related to C. sonorensis (Fig. 1) butis easily distinguished by its mesopodal basidiomes, infun-dibuliform pilei, and oblong ellipsoid spores (Gilbertson andRyvarden 1986). Coltriciella dependens differs in havingpendant basidiomes, attached by a small stipe at the vertex.

The pleuropodal basidiomes and the basidiospores of C.sonorensis also call to mind C. navispora Aime et al.

described from Guyana (Aime et al. 2003). The latter hasslightly infundibuliform pilei, regular, angular, slightly largerpores (1–2 per mm), and possibly, slightly larger basidio-spores, 10–12 × 4–5 μm. Both species also differ in theirecological requirements: C. navispora was found growingscattered in sheltered areas in hollowed, well-decorticatedtrees or on the underside of fallen trunks in forestsdominated by Dicymbe corymbosa Spruce ex Benth. inGuyana, while C. sonorensis was found on soil in open areasin a Q. oblongifolia dominant area, mixed with tropicalcaducifolious elements. The two species are also distantlyrelated phylogenetically (Fig. 1). Coltriciella navisporaclusters within a clade formed of collections identified asC. dependens, all originating from Australia (Tedersoo et al.2007b), while C. sonorensis is related to C. oblectabilis.

Coltriciella pusilla, known so far exclusively from EastAsia (Núñez and Ryvarden 2000), develops similar basi-diomes. It is distinguished by its ellipsoid basidiospores andis reported on the rotten stumps of Castanopsis (Fagaceae)(Imazeki and Kobayashi 1966; Núñez and Ryvarden 2000).Phylogenetically, C. pusilla and C. sonorensis are alsodistantly related.

Coltriciella sonorensis was found on soil in a Q.oblongifolia dominant area. Considering the mycorrhizalnature of Coltriciella species (Tedersoo et al. 2007a), asymbiotic association of C. sonorensis with Quercus couldbe advisably searched.

The phylogenetic analyses also reveal that the circum-scription of C. dependens is vague. Sequences gatheredfrom specimens of mycorrhizal sheets from differentregions of the world are distributed among several clades,and no sequence is 100% identical with the referencesequence AY059059. The latter was obtained from aspecimen originating in Puerto Rico. The circumscriptionof C. dependens should be re-evaluated, taking into accountits autecology, among other features its potential botanicalhost association (Tedersoo et al. 2007a).

Key to species of Coltriciella

1. Basidiomata resupinate to effuse-reflexed..................21. Basidiomata pileate-stipitate.......................................3

2. Basidiomata resupinate; pores 2–3/mm; basidio-spores ellipsoid/subcylindrical, apically slightlytruncate, 7.2–10.5 × 4.2–5.5 μm (Buchanan andRyvarden 1993).................................C. tasmanica(Cleland & Rodway) D.A. Reid

2. Basidiomata effuse-reflexed; pileus subtomen-tose; pores 3–5 per mm (150–300 μm wide);basidiospores mango-shaped, subacute to sub-truncate, 9–11.5 × 4–5.5 μm....................C.corticicola Corner

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3. Basidiomata pendant; pileus velutinate; dorsal stipehairy; pores 2–3 per mm, angular; spores ellipsoid7–10 × 4.5–5.5 μm (Gilbertson and Ryvarden1986)..................................................C. dependens(Berk. & M.A. Curtis) Murrill s. l.

3. Basidiomata erected, pleuropodal or mesopodal.........44. Stipe central..........................................................54. Stipe lateral...........................................................6

5. Basidiospores broadly ellipsoid, 6–8 × 5–6.5 μm;pores 150–300 μm wide, enlarging to 600 μm(about 3–5 per mm); pileus plano-umbilicate,appressedly fibrillose, centre often strigose, cinna-mon ferruginous, fuscous ferruginous in age, faintlyzoned........................................................C. subpicta(Lloyd) Corner

5. Basidiospores oblong ellipsoid with rounded base andsomewhat tapering at the other end, 7–10 × 4–5 μm(Ryvarden 2004); pileus infundibiliform, slightlyshiny, appressed tomentose, dark cinnamon to rusty-brown, weakly concentrically zonate; pores 1–2 (–3)per mm, angular, shallow.......................C. oblectabilis(Lloyd) Kotl., Pouzar & Ryvarden6. Basidiospores navicular, 10–12 × 4–5 μm; pileus

infundibuliform, dull, finely appressed velutinate,rusty brown, weakly concentrically zonate; pores 1–2 per mm, angular.................................C. navisporaT.W. Henkel, Aime & Ryvarden

6. Spores less 10 μm long.........................................77. Pores regular, 2–4 per mm; basidiospores ellipsoid,

yellowish brown, 7.5–8.5 × 4.5–5 μm; basidiomataspathulate; pileus shiny, glabrous; species known sofar only from East Asia..................................C. pusilla(Imazeki & Kobayasi) Corner

7. Pores rounded, elongated to daedaloid, 2–3 per mm;basidiospores oblong to cylindrical, somewhat atten-uated towards the apex, some navicular, 8–10.5 × 4–5 μm; basidiomata flabelliform; pileus appressedvelutinate to slightly fibrillose, dull; species knownso far only from Mexico..........................C. sonorensis

Acknowledgements The authors acknowledge the financial contribu-tion received from the CONACyT (Mexico) and the FNRS (Belgium), inthe framework of their bilateral cooperation agreement that allowedfieldwork in Mexico, and a research visit of R. Valenzuela and T.Raymundo at MUCL. Ricardo Valenzuela, Martín Esqueda, and JoaquínCifuentes thank COFAA and IPN (project SIP-20100942), theSEMARNAT-CONACYT (Grant 2002-C01-0409), and UNAM (projectPAPIIT IN218008), respectively. Mario Amalfi gratefully acknowledgesthe financial support received from the Belgian State – Belgian FederalScience Policy through an Interuniversity Attraction Poles Program(phase VI, contract P6/06) and the logistic support received fromMUCL.Aldo Gutierrez (CIAD) kindly prepared the plates. Cony Decockgratefully acknowledges the financial support received from the BelgianState – Belgian Federal Science Policy (contract BCCMC3/10/003). Theauthors also warmly thank Stéphanie Huret and Céline Bivort for their

help with the sequencing program and Felipe Barredo Pool for taking thephotographs in the Scanning Electron Microscope. The authors thankalsoDr. Fernando Chiang of the Instituto de Biología, UNAM for helpingus with the Latin diagnosis.

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