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Bojamyces Transfuga Sp. Nov. and New Records of Trichomycetes from Mayfly Larvae in Spain

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Bojamyces Transfuga Sp. Nov. and New Records of Trichomycetes from Mayfly Larvae in Spain Mycologia, 96(6), 2004, pp. 1386±1392. q 2004 by The Mycological Society of America, Lawrence, KS 66044-8897 Bojamyces transfuga sp. nov. and new records of Trichomycetes from may¯y larvae in Spain Laia G. Valle1 lishing the generic limits. Their relative position on Sergi Santamaria zygosporophore is the basis to designate the zygo- Unitat de BotaÁnica, Departament de Biologia Animal, spore type (Moss et al 1975). The shape of the hold- de Biologia Vegetal i d'Ecologia, Facultat de CieÁncies, fast and the rami®cation pattern of the thalli are oth- Universitat AutoÁnoma de Barcelona, 08193-Bellaterra er interesting features to consider in classi®cation (Barcelona), Spain (Lichtwardt et al 2001). Longcore (1989) described the genus Bojamyces Abstract: The new species Bojamyces transfuga is de- with the single species B. repens from nymphs and scribed. It was found growing in the hindgut of may- molts of Leptophlebia intermedia Traver (Ephemer- ¯y nymphs and in their shed abdominal exoskeleton optera, Leptophlebiidae) in the United States. Licht- lumen. Bojamyces transfuga shows single appendaged wardt and Williams (1992) reported immature thalli trichospores and zygospores of type I, unlike Boja- from Australasian may¯ies that resemble B. repens. myces repens, where appendages and zygospores are No further occurrence of the taxon has been report- unknown. The generic description of Bojamyces is ed since. emended to accommodate this new taxon. Bojamyces We have found the new species of Bojamyces grow- repens, known only from the United States, has been ing and infecting Caenis luctuosa Burmeister found for the ®rst time in the Iberian Peninsula on (Ephemeroptera, Caenidae) molts and larvae. Both may¯y molts. We also note the presence of Legerio- trichospores and zygospores were found, allowing us myces rarus in Europe, on the Iberian Peninsula and to study and consider diagnostic characters that are Balearic Islands. Previously it was known only in West- different enough to describe a new taxon. The new ern Australia. records of Bojamyces repens have been found in molts Key words: Ephemeroptera, exuvia, gut fungi, of Habroleptoides confusa Sartori & Jacobs (Ephem- Legeriomycetaceae, scanning electron microscopy, eroptera, Leptophlebiidae) with the typical charac- species distribution, taxonomy ters of the species but without zygospores. Longcore (1989) provided a generic diagnosis based on the single species, B. repens, but we believe that the new species must be included also in the INTRODUCTION genus Bojamyces by morphological and ecological The species we describe are included in the order traits. The generic description should be emended to Harpellales, with species having branched or un- account for characters discovered in the new species. branched thalli separated into two families, respec- Legeriomyces rarus Lichtw. was known only by its tively the Legeriomycetaceae and the Harpellaceae. original description from Western Australia in an en- Legeriomycetaceae develop within the hindgut lining demic genus of Caenidae, Tasmanocoenis (Williams of their larval hosts, mainly ¯ies (Diptera), stone¯ies and Lichtwardt 1993). In Spain, this species was (Plecoptera), may¯ies (Ephemeroptera) (Lichtwardt found together with the new Bojamyces species. et al 2001) and very rarely, beetles (Coleoptera) (Lichtwardt et al 1999), caddis worms (Trichoptera) and isopods (White 1999). Harpellaceae grow within MATERIALS AND METHODS the peritrophic matrix of Diptera larvae (Lichtwardt The description of the new species is based on material et al 2001). found in localities from Northeast Spain and Balearic Is- Trichospores provide weighty characteristics used lands. Collections of ephemeropteran nymphs were made for the classi®cation of genera and species, including by hand picking from the stream substrate with entomolog- their size and shape, the number of appendages and ical forceps and by dragging aquatic nets over rocks and their arrangement on fertile branches. Likewise, zy- sand in streams. Nymph's molts were collected between gospores, when found, are very important for estab- leaves and organic debris on the river shores and trans- ported in a small cooler to the laboratory. Insect nymphs Accepted for publication April 13, 2004. were dissected to extract and clean the guts under a stereo- 1 Corresponding author. E-mail: [email protected] microscope using ®ne-pointed forceps and entomological 1386 VALLE AND SANTAMARIA:TRICHOMYCETES ON MAYFLY LARVAE FROM SPAIN 1387 needles. Molts containing the ectodermic hindgut remains imities of these bridges, from one of the conjugants also were examined. Fungi were placed on a slide with water (FIGS. 2, and 5, arrows). as mounting medium and photomicrographed using phase Etymology. L., transfuga 5 transgressor, deserter; contrast and interference contrast optics (DIC). Slides were referring to its habitat, outside the living larval host. ®xed and stained with lactophenol cotton blue to be pre- Specimens examined. SPAIN. BARCELONA: Marganell, served in the herbarium (BCB-Mycotheca, at the institu- Riera de Marganell, prepared from Caenis luctuosa larvae tional address of the authors). and molts, 7-V-2001, L. G. Valle, BCB-Tr0729 (HOLO- Some Caenidae individuals, collected from the locality TY PE), BCB-Tr0707 to BCB-Tr0750 (ISOTY PES); same lo- where the infestation percentage was greatest (Marganell), cality, 6-VI-2001, BCB-Tr0830, BCB-Tr0831; Cerdanyola del were placed in an aquarium until molts were obtained. ValleÁs, Riera de St. Iscle, prepared from Caenis luctuosa lar- Some of the living larvae were dissected just before molting. vae and molts, 7-VI-2001, L. G. Valle, BCB-Tr0832±0834. For SEM, Bojamyces material together with the hindgut The genus Bojamyces is well-de®ned by its thallial molt was ®xed with 2.5% glutaraldehyde in cacodylate buff- er 2M (pH 7). The thalli were washed with the same caco- morphology, development and ecology. Characteris- dylate solution, and dehydrated in an ethanol series (15 tics of appendages in trichospores, important for de- min in each 10, 20, 40, 60, 80%) to absolute ethanol before limitation of many genera in Harpellales, is consid- the thalli were critical point dried and gold metallized. Sam- ered here as a secondary trait, useful for speci®c sep- ples were examined in a Hitachi S-570. aration purposes. Accordingly, the new species has been placed in this genus because the thallial fea- tures, such as the scattered branching, the extremely TAXONOMY long hypha which are irregularly septated in gener- Bojamyces Longcore, emend. L.G. Valle & Santam. ative cells and the disarranged trichospores, are co- Thallus with sparse and indeterminate branching, incident with those described for Bojamyces repens, attached to the host hindgut cuticle. Generative cells the type species of the genus. Nonetheless, the new scattered on the thallus, interspersed with vegetative species reveals the presence of characteristics not in- cells. Trichospores elongate-ellipsoidal, with or with- cluded in the original diagnosis of the genus (Long- out a single appendage and with a ¯ared collar. Zy- core 1989), as trichospores bearing one appendage gospores perpendicularly and medially attached to and the occurrence of zygospores, as well as a differ- the zygosporophore (type I [Moss et al 1975]), with ent ecology in a new host family. a collar. In ephemeropteran nymphs. It is not unusual within Harpellales that different Typus generis: Bojamyces repens Longcore. species of the same genus live in different insect gen- era, even belonging to unrelated insect families. Moreover, Longcore (1989) reported the presence of Bojamyces transfuga L.G. Valle & Santam., sp. nov. a Bojamyces-like fungus in Siphlonurus sp. (Ephem- FIGS. 1±15 eroptera, Siphlonuridae). Several genera of Harpel- 3 m 3 Trichosporae (24±)30(±36) 5±6 m, collare 1.5±2.5 lales display, at least, narrow generic host speci®city, m 2±2.5 m et appendicem, conspicuam in cellula genitali, but some are not so speci®c, like those found mostly ferentes. Trichosporae post liberatas appendicem confertim in Dipteran larvae such as Smittium, Stachylina and plicatam praebentes. Zygosporae (27±)42(±50) 3 8±9 mm, basaliter in¯ato zygosporophoro ad perpendiculum et in Simuliomyces, which also include species in Plecop- medio af®xae. Zygosporae post liberatas cum plane cam- teran nymphs (Lichtwardt et al 2001). Species of Zyg- paniformi collari 7±7.5 mm longitudine et ®liformi append- opolaris grow also in different Ephemeropteran fam- ice. ilies (Lichtwardt and Williams 1984, Moss et al 1975). Trichospores (24±)30(±36) 3 5±6 mm, with a 1.5± Therefore, it is not so unusual that a species of Bo- 2.5 mm long and 2±2.5 mm broad collar and a single jamyces could be present in Caenidae rather than appendage (FIG. 6), visible inside the generative cell Leptophlebiidae or Siphlonuridae. before the trichospore release (FIG. 1, arrows). After Trichospores of B. transfuga are smaller and nar- trichospore release, the appendage appears com- rower than those of B. repens, and show an append- pactly folded (FIG. 7, arrow). Trichospores arise from age (FIGS. 6, 7) that can be seen even inside the gen- generative cells without a de®nite direction. Zygo- erative cell (FIG. 1, arrow) and when just released, it spores (27±)42(±50) 3 8±9 mm, perpendicularly and is folded compactly (FIG. 7, arrow). It is not easy to medially attached to the zygosporophore, which is in- observe the appendage completely unfolded after tri- ¯ated at the lower part (FIGS. 10, 11, arrows). Once chospore release. released, a 7±7.5 mm long ¯ared collar and a single The use of SEM has revealed the presence of a net- ®liform appendage can be observed. Later, typical like skirt of an unknown substance covering the low- scalariform conjugation bridges develop (FIGS.2,ar- er part of the zygosporophores (FIG. 10, arrow), and rows, and 3, 5), and zygospores grow from the prox- also located just at the lower margins of the tricho- 1388 MYCOLOGIA FIGS. 1±15. Bojamyces transfuga. 1. Fertile branches with intercalary trichospores showing the appendage sac inside the generative cell (arrows) and free trichospores [BCB-Tr1832].
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