Mycologia, 95(4), 2003, pp. 685–687. ! 2003 by The Mycological Society of America, Lawrence, KS 66044-8897

Another fossil agaric from Dominican amber

David S. Hibbett1 Manfred Binder Zheng Wang

Etymology. The generic name means ‘‘golden mushroom’’; the epithet refers to the collection locality. Pileus 3 mm broad, convex, with a broad raised center, glabrous or minutely textured, yellow-brown; margin incurved, striated. Lamellae subdistant; margins smooth; lamellulae or anastomoses absent; attachment not observed. Stipe central, 0.8 ! 7 mm, cylindric, smooth; annulus, volva and rhizoids absent. Basidiospores 3.5–4.4 ! 3.0–3.5 "m, broadly elliptic. HOLOTY PE: DOMINICAN REPUBLIC: El Valle, Yanigua mine. In the private collection of Yale Goldman (s.n.). Aureofungus yaniguaensis is an agaricoid homobasidiomycete. The vast majority of such forms occur in the euagarics clade (# Agaricales pro parte), which contains about 8500 described species (Hawksworth et al 1995, Hibbett and Thorn 2001). The shape and stature of the fruiting body suggest that A. yaniguaensis might be related to the smaller palespored genera traditionally classified as Tricholomataceae (e.g., Collybia (Fr.) Staude, Mycena (Pers.) Roussel, Marasmius Fr., Marasmiellus Murrill), or perhaps certain non-deliquescent, dusky-spored taxa (e.g., Coprinellus disseminatus (Pers. : Fr.) J. E. Lange). Unfortunately, the anatomical characters that would be needed to resolve the placement of A. yaniguaensis could not be observed. Because of the thickness of the amber it was not possible to view the specimen with greater than a 10 ! objective, which made it difficult to assess the ornamentation of the spores and the surface textures of the pileus and stipe. The spores appear to be pigmented (FIG. 3), which would suggest a relationship to extant chromosporic groups, but this could be an artifact of preservation or an optical effect of the amber. Lacking evidence of its precise taxonomic placement, we suggest that A. yaniguaensis should be classified as incertae sedis among the Agaricales, euagarics clade. Although it is not possible to precisely place A. yaniguaensis, sufficient characters can be seen to distinguish it from Coprinites dominicana, Protomycena electra and Archaeomarasmius leggeti. The pileus of Coprinites dominicana (Poinar and Singer 1990) was described as squamulose-pectinate with a pleated or grooved margin and a small depression in the center, whereas that of A. yaniguaensis is smooth or glabrescent, with faint striae and a broad raised central area.

Department of Biology, Clark University, 950 Main St., Worcester, Massachusetts 01610

Yale Goldman 86 Dunne Ave., Collinsville, Connecticut 06019

Abstract: We report the discovery of a fossil agaricoid homobasidiomycete from Dominican amber (ca 15–20 Ma). Aureofungus yaniguaensis appears to be a member of the euagarics clade, but its precise taxonomic placement is obscure. This is the fourth known fossil agaric and the third from Dominican amber. Key words: Agaricales, Aureofungus yaniguaensis, euagarics clade, paleomycology Three fossil agarics are known to science. Coprinites dominicana Poinar and Singer and Protomycena electra Hibbett, Grimaldi and Donoghue are both from Dominican amber and are 15–20 million years old (Ma), while Archaeomarasmius leggeti Hibbett, Grimaldi and Donoghue is from Atlantic Coastal Plain amber and is 90–94 Ma (Grimaldi, Beck and Boon 1989, Poinar and Singer 1990, Hibbett et al 1995, 1997, Iturralde-Vinent and MacPhee 1996). Here, we report the discovery of a fourth fossil agaric (FIGS. 1–5), which was collected in the Yanigua mine in the eastern Dominican Republic and purchased from an amber dealer living in the town of El Valle in August 2000. A single fruiting body is present (FIGS. 1–2, 4), as well as many basidiospores that are laid down in masses, suggesting that they were produced by the fruiting body in the amber (FIGS. 3, 5). Based on comparisons to Coprinites Poinar and Singer, Protomycena Hibbett, Grimaldi and Donoghue and Archaeomarasmius Hibbett, Grimaldi and Donoghue, we conclude that the fossil represents a previously undescribed genus and species. Aureofungus yaniguaensis Hibbett, Binder and Wang, gen. et sp. nov. Accepted for publication September 8, 2002. 1 Corresponding author. E-mail: [email protected]

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FIGS. 4–5. Aureofungus yaniguaensis. 4. Fruiting body. 5. Basidiospores. Scale bars: 4 # 100 "m, 5 # 4 "m.

FIGS. 1–2. Aureofungus yaniguaensis fruiting body. Scale bar # 100 "m.

In addition, the hymenophore of Coprinites is reported to have lamellulae, which are absent in A. yaniguaensis. Finally, the spores of Coprinites differ from those of A. yaniguaensis in being larger (6–7 "m long) and oblong-elliptic. Protomycena electra (Hibbett et al 1997) differs from A. yaniguaensis in having a more campanulate pileus with a reflexed margin, and more distant lamellae with lamellulae and anastomoses. Archaeomarasmius leggeti (Hibbett et al 1995, 1997) differs from A. yaniguaensis in having a more strongly sulcate pileus and larger spores (6.5– 8.3 ! 4.0–5.2 "m). The differences described above certainly warrant recognition of A. yaniguaensis as a new species. While we cannot rule out the possibility that A. yaniguaensis is closely related to any of the

previously described fossil agarics, there are no characters (synapomorphies) that provide positive support for such a relationship. Therefore, we think it is appropriate to classify A. yaniguaensis in a new genus. Fossil evidence (Archaeomarasmius) indicates that the euagarics clade is at least 90 million years old (Hibbett et al 1995), and molecular clock studies suggest that the basidiomycetes as a whole could be anywhere from 500 million years to 1.2 billion years old (Berbee and Taylor 2001, Heckman et al 2001). Iturralde-Vinent and MacPhee (1996) suggested that all Dominican amber was formed in a single sedimentary basin during a 5 million year interval in the Miocene (15–20 Ma; contra Poinar and Singer 1990). Thus, A. yaniguaensis does not affect the minimum age estimates for either the euagarics clade or the basidiomycetes. Nevertheless, fossil fungi in Dominican amber, such as A. yaniguaensis, are of value for understanding the ecology of Miocene ecosystems in the Caribbean and could help evaluate certain historical biogeographic hypotheses (see Hibbett 2001 for an example). The discovery of a third fossil agaric from Dominican amber suggests that many more such finds might lie in store. ACKNOWLEDGMENTS

We are grateful to David Grimaldi, who examined the specimen and commented on its authenticity, and Scott Redhead, who made valuable editorial suggestions. LITERATURE CITED

FIG. 3. Aureofungus yaniguaensis basidiospores. Scale bar # 10 "m.

Berbee ML, Taylor JW. 2001. Fungal molecular evolution: gene trees and geologic time. In: McLaughlin DJ, McLaughlin EG, Lemke PA, eds. The mycota VII systematics and evolution. Part B. Berlin: Springer-Verlag. p 229–245. Grimaldi D, Beck CW, Boon JJ. 1989. Occurrence, chemical characteristics and paleontology of the fossil resins

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from New Jersey. American Museum Novitates 2948:1– 27. Hawksworth DL, Kirk PM, Sutton BC, Pegler DN. 1995. Dictionary of the fungi. 8th ed. Wallingford: CAB International. 616 p. Heckman DS, Geiser DM, Eidell BR, Stauffer RL, Kardos NL, Hedges SB. 2001. Molecular evidence for the early colonization of land by fungi and plants. Science 293: 1129–1133. Hibbett DS. 2001. Shiitake mushrooms and molecular clocks: historical biogeography of Lentinula. J Biogeogr 28:231–241. ———, Thorn RG. 2001. Basidiomycota: Homobasidiomy-

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cetes. In: McLaughlin DJ, McLaughlin EG, Lemke PA, eds. The mycota VII systematics and evolution. Part B. Berlin: Springer-Verlag. p 121–168. ———, Grimaldi D, Donoghue MJ. 1995. Cretaceous mushrooms in amber. Nature 377:487. ———, ———, ———. 1997. Fossil mushrooms from Miocene and Cretaceous ambers and the evolution of homobasidiomycetes. Am J Bot 84:981–991. Iturralde-Vinent MA, MacPhee RDE. 1996. Age and paleogeographical origin of Dominican amber. Science 273: 1850–1852. Poinar GO Jr., Singer R. 1990. Upper Eocene gilled mushroom from the Dominican Republic. Science 248: 1099–1101.

Another fossil agaric from Dominican amber - Clark University

Abstract: We report the discovery of a fossil agari- coid homobasidiomycete from Dominican amber (ca. 15–20 Ma). Aureofungus yaniguaensis appears to be a.

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