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Kurtziana

versión On-line ISSN 1852-5962

Kurtziana vol.37 no.1 Córdoba ene./jun. 2012

 

ARTÍCULOS ORIGINALES

On Neotropical Daedalea species: Daedalea ryvardenica sp. nov.

 

Elisandro R. Drechsler-Santos 1*, María A. de Queiroz Cavalcanti 2, Clarice Loguercio-Leite 1 & Gerardo L. Robledo 3*

1 Laboratório de Micologia, Departamento de Botânica, Universidade Federal de Santa Catarina, CEP: 88040-900 Florianópolis, SC, Brazil.
2 Departamento de Micologia, Universidade Federal de Pernambuco, CEP: 50760-420 Recife, PE, Brazil.
3 Laboratorio de Micología, Instituto Multidisciplinario de Biología Vegetal, CONICET - Universidad Nacional de Córdoba, CC 495, CP 5000 Córdoba, Argentina.
* Corresponding authors: drechslersantos@yahoo.com.br and glrobledo@yahoo.com

 


Summary

The new species, Daedalea ryvardenica, presents irregular hymenophore, generative hyphae with irregular thickened walls, skeletal hyphal endings forming an irregular palisade within the hymenium, and presents the largest basidiospores in the genus, with a ventral concavity next to the apiculous and tapering apex. A taxonomic discussion of the morphological characters of Daedalea, the hyphal system and the shape of the basidiospores, and a key of Neotropical Daedalea species are presented.

Key words: Caatinga; Cerrado; Polypores; Taxonomy.

Resumen

Sobre las especies Neotropicales de Daedalea: Daedalea ryvardenica sp. nov.

La nueva especie, Daedalea ryvardenica, presenta himenoforo irregular, hifas generativas con paredes irregularmente engrosadas, terminaciones de hifas esqueletales formando una empalizada irregular en el himenio y por presentar las basidiosporas más grandes en el género con una concavidad ventral cerca del apículo y ápice que se adelgaza. Se presenta una discusión taxonómica sobre los caracteres morfológicos de Daedalea, sistema hifal y forma de las basidiosporas, y una clave para las especies Neotropicales de Daedalea.

Palabras clave: Caatinga; Cerrado; Políporos; Taxonomía.


 

Introduction

During studies of polypores from Caatinga and Cerrado Biomes of Brazil (IBGE, 2004), several collections of a brown-rot species from subxerophytic to dry ecosystems deserve special taxonomic attention. One species with thin and flexible pilei and an irregular hymenophore was at first identified as Daedalea stereoides Fr. However, the distinct large spores of the Brazilian specimens require the proposal a new taxon of Daedalea Pers.

In the Neotropics, the genus has been poorly studied and there are few species of Daedalea occurring from Belize to Argentina (Wright & Deschamps, 1975; Rajchenberg, 1986; Lindblad & Ryvarden, 1999; Gugliotta et al., 2010; Lindner et al., 2011). The most representative species is Daedalea aethalodes (Mont.) Rajchenb., originally described from Brazil as Trametes aethalodes Mont., abundantly recorded in most of the ecosystems (from semiarid to rain forests) of N Argentina and Brazil (Rajchenberg, 1986; Gugliotta et al., 2010).

Molecular phylogenetic works show that Daedaelea species cluster in the"Antrodia clade" (Hibbett & Binder, 2002; Kim et al., 2003; Binder et al., 2005; Rajchenberg et al., 2011). Recently, a synopsis of the genus Daedalea sensu stricto has been provided by Lindner et al. (2011) based on morphological characteristics and analyses of the rDNA ITS regions from the genus type species, D. quercina (L.) Pers., and five others. According to Lindner et al. (2011) Daedalea s.s. includes brown rot species with a trimitic hyphal system, clamped, thin-walled generative hyphae; a cork-coloured context, small, hyaline, oblong-ellipsoid to cylindrical spores, and a distinct catahymenium. However, there are other views concerning the key characteristics of the Daedalea genera in the literature. For example, Rajchenberg (1986) describes the noteworthy morphological characteristics of Daedalea as being the presence of generative hyphae with irregularly thickened walls, the nature of the hyphal system, and the tramal cystidia that protrude into the hymenium, forming a distinct catahymenium of widened skeletal ends.

In this work we describe a new Neotropical species of Daedalea with distinct large spores, and discuss some of the main morphological characteristics of the genus. In addition, a key for Neotropical species of Daedalea is presented.

Material and Methods

Collecting was performed in the Caatinga (Ceará, Paraíba and Pernambuco States) and Cerrado (Mato Grosso State) biomes of Brazil and collections were deposited at CORD, FCHBS, FLOR, O, and URM Herbaria. Herbaria acronyms are according to Holmgren et al. (1990). Basidiomata were analyzed macro- and micro-morphologically. Microscopic observations were made from slide preparations stained with 3-5% KOH and Phloxine, or Melzer's reagent (Ryvarden, 1991). Color designations of basidiomata follow Watling (1969).

Taxonomy

Daedalea ryvardenica Drechsler-Santos& Robledo sp. nov. Figs. 1A-D, 2A-D, 3A-E.


Fig. 1. Daedalea ryvardenica: microscopic features of holotype. A. basidiospores; B. generative hyphae with irregular thickened walls; C. skeletal hyphae dominating in the basidiome; D. short skeletal hyphae that end in the hymenium.


Fig. 2. Daedalea ryvardenica: microscopic features of holotype. A-D. generative hyphae with irregular thickened walls (photos by Marco Antonio Borba da Silva). Bars=10 µm.


Fig. 3. Macroscopical features of Neotropical Daedalea species. A-E. D. ryvardenica (type specimen). A. general view in situ. B. upper view with details of brown rot in the wood. C. detail of the pilear surface. D. section showing context and tube layer. E. detail of the pore surface. F-H. D. stereoides (Robledo 983). F. section showing context and tube layer . G. detail of the pilear surface. H. detail of the pore surface. I-L. D. aethalodes. I. general view in situ of fresh specimen (Robledo 1563). J. detail of the pilear surface (Robledo 1824). K. detail of the pore surface (Robledo 1389). L. section showing context and tube layer (Robledo 1988).

Mycobank: MB 563588.

BRAZIL. Mato Grosso. Chapada dos Guimarães: Parque Nacional da Chapada dos Guimarães, Véu da Noiva (15°24'25"S, 55°50'17"W), Ferreira-Lopes VFL001, 12- VI-2011. Holotypus specei (FLOR41052). Figs. 1A-D, 2A-D, 3A-E

Etymology: in honor of Dr. Leif Ryvarden.

Basidiome annual, effuse-reflexed to pileate, thin, 0.3 to 1.5 cm thick, flexible and coriaceous, semicircular to flabelliform, forming clusters or imbricate, 3.0-9.0 X 2.5-5.0 cm when solitary, generally concrescent up to 28 cm wide; upper surface vinaceus buff (31), buff (32) to fulvous (12), glabrous, slightly concentrically zonate and radially scrupose; margin thinning out and curved; context homogenous, whitish to vinaceous buff (31), up to 0.7 cm thick.

Hymenophore poroid, regular with angular pores at the base to completely irregular, with daedaloid, semi-labyrinthic to lamellae-like portions, the transversal dissepiments become lacerate with age and some portions are almost irpicoid, 1-3/mm; hymenial surface purplish chestnut (21) to vinaceous buff (31); tubes up to 0.8 cm deep.

Hyphal system dimitic. Generative hyphae with clamps; hyaline and thin-walled hyphae restricted to the sub-hymenium, 2-3 µm diam.; throughout the basidiome generative hyphae are pale yellowish to hyaline, notoriously thickwalled with irregular thickened walls, showing a sinuous and undulating lumen, 4-6 µm diam. Skeletal hyphae dominating in the basidiome, with a basal, more or less sinuous and geniculate portion, generally with 1-3 short branches, aborted branch process can often be observed, followed by a main part that is straight, thick walled, and with a straight lumen. Sometimes these main parts present thick walls with a capillary lumen or are almost solid, in the apex walls thin and secondary septa are present, up to 130 µm; in the trama of the tubes skeletal hyphae are shorter and those that end protruding into the hymenium are thick-walled right up to the rounded apex.

Cystidia absent, but with abundant skeletal hyphae that end protruding into the hymenium among the basidia. Basidia clavate, 15-23 x 5-8 µm. Basidiospores ellipsoid, with a distinct ventral concavity next to the apiculous, and a tapering apex, (7.0-) 7.5-11.0 x 2.5-3.5 µm, X= 9.1 x 2.9 µm (n= 40), smooth, thin-walled, hyaline, IKI-.

Specimens examined: BRAZIL. Ceará: Graças: Cachoeira do Belizário (4º06'29"S, 40º45'47"W, 435 m. a. s. l.), Drechsler- Santos DS003CE, 13-VI-2007. (URM80481, O); Reriutaba: Trilha d'olho da água da bananeira (4º02'15"S, 40º39'32"W, 240 m. a. s. l.), Drechsler-Santos DS013CE, 14-VI -2007. (URM80383) ; Mato Grosso: Chapada dos Guimarães: Parque Nacional de Chapada dos Guimarães, Vale do Rio Claro (15°31'08" S, 55° 08'44" W), Costa-Rezende CR001, 15-V-2011. (FLOR41053, FCHBS); Ibid (15°31'08" S, 55°84'43" W), Costa-Rezende CR003, 15- V-2011. (FCHBS); Chapada dos Guimarães: Ibid, Costa-Rezende CR004, 15-V-2011. (FCHBS); Ibid (15°31'45,7" S, 55°85'27" W), Costa-Rezende CR37, 11-IX-2011. (FCHBS); Ibid, Costa-Rezende CR38, 11- IX-2011. (FCHBS); Ibid, Costa-Rezende CR39, 11-IX-2011. (FCHBS); Chapada dos Guimarães: Parque Nacional de Chapada dos Guimarães, Véu da Noiva (15°24'25" S, 55°50'17" W), Ferreira-Lopes VFL001, 12-VI-2011. (FLOR41052, FCHBS); Alves- Silva GAS001, 14-VIII-2011. (FLOR41054, FCHBS);, Alves-Silva GAS005, 14-VIII- 2011. (FCHBS); Paraíba: Sumé: Fazenda Almas (7º28'16"S, 36º53'53"W, 623 m. a. s. l.), Rajchenberg & Baltazar 014PB, 10- VIII-2008. (URM80492); Ibid, Rajchenberg et al. 023PB, 10-VIII-2008. (URM80510); Ibid (7º28'27"S, 36º53'27"W, 700 m. a. s. l.), Drechsler-Santos et al. DS054PB, 11- VIII-2008. (URM80518); Ibid, Rajchenberg et al. 020PB, 10.VIII.2008. (URM80513); Ibid (7º28'16"S, 36º53'53"W, 623 m. a. s. l.), Rajchenberg et al. 026PB, 10-VIII- 2008. (URM80497); Ibid, Nogueira-Melo et al. 033PB, 10-VIII-2008 (URM80515); Pernambuco: Parque Nacional do Catimbau: Trilha da Batinga (8º30'39"S, 37º16'54"W, 971 m. a. s. l.), Drechsler-Santos & B.T. Goto DS070, 28-IX-2006. (URM80700, O); Ibid, Trilha das Torres (8º34'20"S, 37º14'45"W, 774 m. a. s. l.), Drechsler-Santos DS105, 08-XII- 2006. (URM80771, O); Serra Talhada: Estação Experimental do IPA (7º53'63"S, 38º18'34"W, 501 m. a. s. l.), Drechsler- Santos DS019PE, 12-IX-2007 (URM80528, O).

Additional specimens examined: Daedalea aethalodes - ARGENTINA. Jujuy: Dpto. Ledesma: Parque Nacional Calilegua, Mesada de las colmenas, sendero La cascada (23º42'1,5"S, 64º51'56,8"O, alt. 1082 m. a. s. l.), Robledo 1563, 24-V-2007. (CORD); Ibid, Robledo 1255, 24-V-2007. (CORD); Ibid (23º45'18"S, 64º51'13"O, alt. 720 m. a. s. l.), Robledo 1824, 01-IV-2008. (CORD). Salta: Dpto. Sta. Victoria Oeste: Parque Nacional Baritú, Camino a Campo Grande (22º26'10,8"S, 64º43'40.2"O, alt. 1990 m. a. s. l.), Robledo 1389, 09-V-2007. (CORD); Ibid (22º26'21,6"S, 64º44'7,8"O, alt. 1222 m. a. s. l.), Robledo 1264, 06-V- 2007. (CORD). BRASIL. Pernambuco: Triunfo (7º52'53"S, 38º06'15"W, 550 m. a. s. l.), Robledo 1988, 26-V-2009. (CORD). Daedalea stereoides - ARGENTINA. Jujuy: Dpto. Ledesma: Parque Nacional Calilegua, Sendero Tataupá (23º44'33,3"S, 64º51'8,5"O, alt. 745 m. a. s. l.), Robledo 1618, 23.V.2007. (CORD); Ibid, sendero La cascada (23º42'1,5"S, 64º51'56,8"O, alt. 1082 m. a. s. l.), Robledo 1558, 22-V-2007. (CORD); Salta: Dpto. Anta: Parque Nacional El Rey (25º00'3"S, 64º35'38,3"O, alt. 810 m. a. s. l.), Robledo 983, 23-III-2007. (CORD).

Ecology and distribution: Saprobe, found on dead fallen branches and logs of angiosperms, causing a brown-rot; currently this species is known from the subxerophytic to dry areas of Caatinga (Ceará, Paraíba, and Pernambuco States) and Cerrado ecosystems (from Mato Grosso State).

Discussion

The identification of this new species in Daedalea is based on the combination of an irregular hymenophore and an associated brown rot. In addition, the generative hyphae with irregular thickened walls forming a sinuous lumen (Figs. 2A-D), the skeletal hyphal endings forming an irregular palisade into the hymenium, the presence of purplish chestnut to vinaceous buff hymenial surface, and the particular basidiospore shape are characters that warrant this placement.
The inclusion of a mostly poroid species (i.e. Daedalea aethalodes) into Daedalea was proposed by Rajchenberg (1986). Besides the type of rot, Rajchenberg (1986) also based his classification on micro morphological characters, such as the irregular thickening of generative hyphae walls and the endings of skeletal hyphae protruding into the hymenium. As previously mentioned, all of these characters are present
in Daedalea ryvardenica. Rajchenberg (1986) also discussed the nature of the hyphal system showing that D. aethalodes does not present a typical trimitic hyphal system [i.e. as in Trametes versicolor (L.) Lloyd], because binding hyphae always present a prolongation in one of their branches. We confirm that D. aethalodes, D. stereoides and D. ryvardenica have a dimitic hyphal system with skeletal hyphae presenting a sinuous geniculate base that is occasionally branched, with a distinctly straight apical portion. The hyphae are long, with thin walled widened apices and, gradually towards the tubes, they become shorter, with thick walled apices into the hymenium. We did not study Daedalea hydnoides I. Lindblad & Ryvarden, however, the presence of skeletal endings as skeletocystidia bending into the hymenium has been reported by Lindblad & Ryvarden (1999).

Regarding macroscopical features, the pinkish, purplish chestnut to vinaceous buff colors of the basidiome is a character present in all Neotropical species of the genus. Daedalea stereoides and D. ryvardenica show a very similar pigmentation (Figs. 3A-G) and Daedalea neotropica presents a characteristic violet pigmentation (Lindner et al., 2011). We report here that specimens of D. aethalodes present pinkish vinaceous tints when fresh (Figs. 3I-J), that typically turn brown to dark brown when the basidiome matures or is dried (Rajchenberg, 1986). A violent black reaction with KOH is present in the context and pileus surface of D. aethalodes, D. stereoides and D. ryvardenica, but it later fades to a dark gray or a grayish spot (Fig 3A). This reaction was not reported for D. neotropica (Lindner et al., 2011).

Daedalea ryvardenica basidiospores have a particular morphology, with a ventral concavity next to the apiculum and a tapering apex. This morphological pattern was observed in other related brown rot taxa such as Antrodia serialis (Fr.) Donk, Antrodia malicola (Berk. & M.A. Curtis) Donk, and Fomitopsis nivosa (Berk.) Gilb. & Ryvarden (Gilbertson & Ryvarden, 1986; Rajchenberg, 1986, 2006). In spite of the fact that basidiospore descriptions of Daedalea species are generally presented as cylindrical to ellipsoid, spore drawings suggest that the above mentioned morphological pattern could be consistent for the Daedalea and related genera. A critical description of the spore's bilateral symmetry must be taken into account to evaluate its potential as a defining morphological characteristic in Daedalea.

Molecular phylogenetic analyses that include Daedealea species suggest that the limits with other brown rot genera (i.e. Fomitopsis and Antrodia) are not clear (Lindner et al., 2011; Rajchenberg et al., 2011). Therefore, for the moment, we prefer to place the new species in Daedalea until new molecular studies including more Neotropical taxa are developed. Furthermore, preliminary phylogenetic studies of Daedalea (Lindner et al., 2011) light the way for future investigations and raise new systematic and taxonomic hypothesis, such as the existence of distinct geographical lineages (i.e. North temperate, Asian and Neotropical species) as previously demonstrated for other polypore genera, for example Fomitiporia Murrill by Decock et al. (2007). Among the Neotropical species, another taxonomic problem is that of Daedalea microsticta Cooke, which was described from Rio de Janeiro State (Brazil), but the material from Costa Rica studied by Lindner et al. (2011) fell in the core polyporoid clade (white-rot species) in the phylogenetic analysis, raising questions about whether this species belongs to Daedalea s.s.

Some characters of the traditional morphological circumscription of Daedalea, i.e. a trimitic hyphal system, thin-walled generative hyphae, presence of binding hyphae sensu stricto and basidiospore morphology (Gilbertson & Ryvarden, 1986; Núnez& Ryvarden, 2001; Lindner et al., 2011) should be re-evaluated. A critical analysis of the above mentioned morphological characters could be very important in order to create a phylogeny that integrates molecular data, biological features (i.e. nuclear behavior, mating systems), decaying abilities and ecological strategies as recently proposed by Rajchenberg (2011).

Key to Neotropical Daedalea species

1. Basidiospores (7.0-) 7.5-11.0 µm long. ............................................................................... D. ryvardenica
1'. Basidiospores shorter, up to 7.0 µm long. .................................................................................................. 2

2. Hymenophore poroid ..................................................................................................... D. aethalodes
2'. Hymenophore irregular, daedaloid, irpicoid to hydnoid ..................................................................... 3

3. Basidiospores 6.0-7.0 x 2.0-2.5 µm, pilear surface rust brown to dark brown ....... D. hydnoides
3'. Basidiospores shorter, up to 5.5 µm long, pilear surface paler (brown to whitish) .................... 4

4. Basidiospores cylindrical, 5.0-5.5 x 2.0-3.0 µm, violaceous coloration that occurs in irregular patches on the pileus present, on Quercus ...................................................................................................... D. neotropica
4'. Basidiospores ellipsoid, 3.5 - 5.5 X 2.0 - 2.5 µm, pilear surface homogenoulsy pale brown, violaceous coloration absent, on other substrates ............................................................................................. D. stereoides

Acknowledgments

We thank the curators of CORD, FCHBS, FLOR, O and URM Herbaria for access to collections under their keeping; the responsible for the collecting areas for access and permitions concerned and the IBAMA/ICMBio; D. Bousfield for expert review of English and valuable contribution with this work; D.H. Costa-Rezende, G. Alves-Silva and V. Ferreira- Lopes for kind support in the field work, M.A. Borba-Silva for photos of figure 2, and Andrei Mello for important contribution with this work; Conselho Nacional de Desenvolvimento Científico (CNPq) and Coordenação de Aperfeiçoamento Pessoal de Nível Superior- CAPES and Ministerio de Ciência, Tecnologia e Innovación Productiva - MINCYT for partially financing this research (CNPq Projects: Universal 478973/2006-3 and 479961/2007- 7, Pesquisa em Biodiversidade do Semi-árido 010105.00/2004/PPBio, and Base de Dados Consolidada das Plantas e Fungos do Nordeste 552615/05-6); CAPES-MINCYT Project: 161/09 - BR/08/13 - Cooperação Brasil- Argentina). GLR is grateful to G. Bertone and A. Bringas (CPA CONICET-UNC) for their technical support, to Idea Wild for their support with technical equipament and to A. Gil for their support in developing this work. CNPq, MINCYT and CAPES also provided PhD (modalities: GD 141072/2006-7 and SWE 201847/2008-6) and Pos-doctoral scholarships (CAPES-MINCYT and CAPES/REUNI/UFSC) to ERDS. This work is part of project "Fungos poliporóides (Agaricomycetes) do PARNA Chapada dos Guimarães, Mato Grosso - Políporos PNCG-MT" (Depto. Botânica/UFSC nº 2011.0421).

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Original recibido el 1 de Diciembre de 2011;
primera decisión: 11 de Enero de 2012;
aceptado el 15 de Febrero de 2012.

Editor responsable: Carlos Urcelay.