- Email: [email protected]
A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber
f u n g a l b i o l o g y x x x ( 2 0 1 6 ) 1 e4
journal homepage: www.elsevier.com/locate/funbio
A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber George POINAR Jr* Department of Integrative Biology, Oregon State University, Corvallis, OR 97331, USA
article info
abstract
Article history:
A densely scaled small mushroom in Baltic amber is described as Gerontomyces lepidotus
Received 17 May 2016
gen. et sp. nov. and is characterized by a convex pileus 1.0 mm in diameter, distant to sub-
Received in revised form
distant lamellae with smooth margins and a centrally inserted cylindrical, solid stipe. Its
8 June 2016
taxonomic placement is uncertain. This is the first mushroom described from Baltic amber.
Accepted 9 June 2016
ª 2016 British Mycological Society. Published by Elsevier Ltd. All rights reserved.
Corresponding Editor: Nicholas P Money Keywords: Eocene amber Cenozoic Agaricales Fossil mushroom
Introduction Considering their fragile and transitory nature, it is not surprising that the great majority of fossil mushrooms (using the term in a general sense that includes macroscopic fungal fruiting bodies) are known from Tertiary and Cretaceous amber deposits. Aside from gilled mushrooms in amber (Poinar & Singer 1990; Hibbett et al. 1995, Hibbett et al. 1997, 2003; Poinar & Buckley 2007), there are also examples of puffballs (Poinar 2001) and earthballs (Poinar et al. 2014). While a wide diversity of angiosperms (5 families of monocots and 38 families of dicots) have been described from Eocene Baltic amber, there are no previous accounts of fossil mushrooms from these extensive deposits (Bachofen-Echt 1949; Czeczott 1961; Larsson 1978; Weitschat & Wichard 2002).
The present study describes a new genus and species of gilled mushroom in Baltic amber. Fossiliferous amber from the Baltic region was formed in tropical and subtropical forests that covered a large part of northern Europe for some 10 million years during the early Cenozoic (Bachofen-Echt 1949; Larsson 1978; Weitschat & Wichard 2002). The amber was eroded from the original accumulation and redeposited in marine sediments referred to as the ‘blue earth’ layers in the Samland Peninsula. Original age estimates of amber from this region ranged from 35 to 55 million years (Larsson 1978; Weitschat & Wichard 2002). The latest estimate, based on geochemical tests performed on amberbearing sediments, was between 45 and 55 million years (Wolfe et al. 2016).
* Corresponding author. Tel.: þ1541 760 7319; fax: þ1541 757 1639. E-mail address: [email protected] http://dx.doi.org/10.1016/j.funbio.2016.06.008 1878-6146/ª 2016 British Mycological Society. Published by Elsevier Ltd. All rights reserved.
Please cite this article in press as: Poinar G, Jr, A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber, Fungal Biology (2016), http://dx.doi.org/10.1016/j.funbio.2016.06.008
2
G. Poinar
Materials and methods The specimen originated from the Samland Peninsula in the Kalinin District of the Russian Federation (54.7104 N: 20.4522 E). The amber piece was reshaped and polished to a final length of 9 mm, width of 3 mm, and depth of 3 mm. Observations and photographs were made with a Nikon SMZ-10 R stereoscopic microscope and Nikon Optiphot compound microscope with magnification up to 800. Helicon Focus Pro X64 (D-Studio, 22 Krakovska St. Kiev, Ukraine 02094) was used to stack photos for better clarity and depth of field.
Results The specimen is small, but well preserved and includes a complete pileus and attached stipe that is broken at the base. An insect nymph and strand of mammalian hair are associated with the specimen. Phylum Basidiomycota Order Agaricales Gerontomyces lepidotus Poinar, gen. et sp. nov. (Figs 1e4) Description Total length of pileus and remaining stipe ¼ 1.8 mm; pileus brown, bell-shaped, convex, dry, thick, with straight to slightly incurved margin; surface densely scaly with decumbent to erect scales extending outward to 35 mm; pileus diameter, 1.0 mm; lamellae (12e14 in number) white, broad, flat, distant to subdistant, with smooth uniform margins, attached to stipe apex: lamellulae and spores absent; stipe brown, base broken (remaining portion 1.5 mm long and 0.3 mm wide), dry, solid, insertion central, covered
Fig 1 e Gerontomyces lepidotus gen. et sp. nov. with adjacent insect exuviae (arrow) and mammalian hair in Baltic amber. Bar [ 0.5 mm.
Fig 2 e Detail of pileus and stipe of Gerontomyces lepidotus gen. et sp. nov. in Baltic amber. Bar [ 0.2 mm.
Fig 3 e Pileus of Gerontomyces lepidotus gen. et sp. nov. viewed from above showing slightly prolonged area with attached water droplet (arrow). Bar [ 0.3 mm.
Please cite this article in press as: Poinar G, Jr, A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber, Fungal Biology (2016), http://dx.doi.org/10.1016/j.funbio.2016.06.008
Gilled mushroom in Baltic amber
3
Discussion
Fig 4 e Ventral (under) view of Gerontomyces lepidotus gen. et sp. nov. in Baltic amber showing lamellae. Bar [ 0.5 mm.
with decumbent to erect scales extending outward to 30 mm; veil and annulus absent. Holotype: Accession # Myc-12, deposited at the Bone Room, 1573 Solano Ave., Berkeley, CA 94707. Etymology: Generic name from Greek ‘geraios’ ¼ old and ‘mykes’ ¼ mushroom. Specific epithet is from the Greek ‘lepidotos’ ¼ scaly in reference to the dense scaly covering of the fossil. Type locality: Samland Peninsula of the Baltic Sea in the Kalinin District of the Russian Federation. Diagnosis: The combination of small size, dry, brownish, densely scaly surface, bell-shaped, convex, dry pileus, broad, distant to subdistant lamellae, the absence of lamellulae, and a straight, solid stipe lacking a veil and annulus characterize the fossil and distinguish it from previously described fossil gilled mushrooms. While the top of the pileus was originally probably round, it was preserved slightly asymmetrical with one margin extended (Fig 3). This is probably due to moisture that had collected on the extended side, some of which collected as an attached water droplet (Fig 3). Moisture is often represented as a gray film on the surface of fossils in Baltic amber. Adjacent to the mushroom is the exuvia of a nymph of a Phasmatodea (Fig 1). Measuring only 0.6 mm in body length, the nymph has features of Pseudoperla lineata Pictet & Hagen, which is fairly common in Baltic amber (Bachofen-Echt 1949; Larsson 1978). In the same piece of amber is a strand of mammalian hair. The strand is several millimetres in length with a diameter reaching 140 mm.
The evolutionary history of gilled mushrooms is restricted to the relatively few fossils from Cenozoic and Cretaceous amber, with the oldest specimen, Palaeoagaricites antiquus, in Myanmar amber showing an affinity to members of the family Tricholomataceae (Taylor et al. 2015). Due to the absence of spores and the basal portion of the stipe, it is difficult to align Gerontomyces lepidotus gen. et sp. nov. with any modern genus or even family. Thus it is not possible to use the fossil to provide minimum age estimates for the origins of any particular agaric lineage. The combination of a small size, dry, brownish, scaly surface, broad, distant to subdistant lamellae and a straight, solid stipe are not common characters in gilled mushrooms (Lincoff 1981; Arora 1986; McKnight & McKnight 1987). However, some of these features do occur in a few genera in the family Tricholomataceae. Members of the genus Squamanita are small woodland mushrooms that have a scaly pileus and stipe with some erect scales as in the fossil (Arora 1986). However species of Squamanita have a distinct veil. The adjacent nymphal exuviae of a phasmatoid could imply a feeding association since phasmatoids are herbivorous and there are records of extant phasmatoids feeding on mushrooms. Thus there is a possibility that the insect was exploring the pileus of the mushroom just as the latter was covered with resin. Finding the empty exuviae suggests that the nymph made a narrow escape.
Conclusions The Baltic amber mushroom reported here adds an additional degree of diversity among gilled Agaricales that existed in the Cenozoic.
Acknowledgements I thank Diana Mansfield of the Bone Room (1573 Solano Avenue, Berkeley, California 94707) and Alex Brown for facilitating the loan of the present specimen and Roberta Poinar for reading earlier drafts of this manuscript.
references
Arora D, 1986. Mushrooms Demystified, 2nd edn. Ten Speed Press, Berkeley, 959 pp.. € sse. Springer Bachofen-Echt A, 1949. Der Bernstein und seine Einschlu Verlag, Vienna, 204 pp.. Czeczott H, 1961. The flora of the Baltic amber and its age. Prace Muzeum Ziemi (Paleobotaniczne) Warszawa 4: 119e145. Hibbett DS, Grimaldi D, Donoghue MJ, 1995. Cretaceous mushrooms in amber. Nature 377: 487. Hibbett DS, Grimaldi D, Donoghue MJ, 1997. Fossil mushrooms from Miocene and Cretaceous ambers and the evolution of Homobasidiomycetes. American Journal of Botany 84: 981e991.
Please cite this article in press as: Poinar G, Jr, A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber, Fungal Biology (2016), http://dx.doi.org/10.1016/j.funbio.2016.06.008
4
Hibbett DS, Binder M, Wang Z, Goldman Y, 2003. Another fossil agaric from Dominican amber. Mycologia 95: 685e687. Larsson SG, 1978. Baltic Amber e a palaeobiological study. Scandinavian Science Press, LTD, Klampenborg. Lincoff GH, 1981. The Audubon Society Field Guide to North American Mushrooms. Alfred A. Knopf, New York, 926 pp.. McKnight KH, McKnight VB, 1987. A Field Guide to Mushrooms. Houghton Mifflin Company, Boston, 429 pp.. Poinar Jr GO, 2001. Fossil puffballs (Gasteromycetes: Lycoperdales) in Mexican amber. Historical Biology 15: 219e221. Poinar Jr GO, Singer R, 1990. Upper Eocene gilled mushroom from the Dominican Republic. Science 248: 1099e1101. Poinar Jr GO, Buckley R, 2007. Evidence of mycoparasitism and hypermycoparasitism in Early Cretaceous amber. Mycological Research 111: 503e506.
G. Poinar
Poinar Jr GO, Donis da Silva A, Goulart Baseia I, 2014. A gasteroid fungus, Palaeogaster micromorpha gen. & sp. nov. (Boletales) in Cretaceous Myanmar amber. Journal of the Botanical Research Institute of Texas 8: 139e143. Taylor TN, Krings M, Taylor EL, 2015. Fossil Fungi. Elsevier, Inc., Amsterdam, 382 pp.. Weitschat W, Wichard W, 2002. Atlas of Plants and Animals in Baltic amber. Verlag Dr. Friedrich Pfeil, Munich, 256 pp.. Wolfe AP, McKellar RC, Tappert R, Sodhi RNS, Muehlenbachs K, 2016. Bitterfeld amber is not Baltic amber: three geochemical tests and further constraints on the botanical affinities of succinite. Review of Palaeobotany and Palynology 225: 21e32.
Please cite this article in press as: Poinar G, Jr, A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber, Fungal Biology (2016), http://dx.doi.org/10.1016/j.funbio.2016.06.008
journal homepage: www.elsevier.com/locate/funbio
A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber George POINAR Jr* Department of Integrative Biology, Oregon State University, Corvallis, OR 97331, USA
article info
abstract
Article history:
A densely scaled small mushroom in Baltic amber is described as Gerontomyces lepidotus
Received 17 May 2016
gen. et sp. nov. and is characterized by a convex pileus 1.0 mm in diameter, distant to sub-
Received in revised form
distant lamellae with smooth margins and a centrally inserted cylindrical, solid stipe. Its
8 June 2016
taxonomic placement is uncertain. This is the first mushroom described from Baltic amber.
Accepted 9 June 2016
ª 2016 British Mycological Society. Published by Elsevier Ltd. All rights reserved.
Corresponding Editor: Nicholas P Money Keywords: Eocene amber Cenozoic Agaricales Fossil mushroom
Introduction Considering their fragile and transitory nature, it is not surprising that the great majority of fossil mushrooms (using the term in a general sense that includes macroscopic fungal fruiting bodies) are known from Tertiary and Cretaceous amber deposits. Aside from gilled mushrooms in amber (Poinar & Singer 1990; Hibbett et al. 1995, Hibbett et al. 1997, 2003; Poinar & Buckley 2007), there are also examples of puffballs (Poinar 2001) and earthballs (Poinar et al. 2014). While a wide diversity of angiosperms (5 families of monocots and 38 families of dicots) have been described from Eocene Baltic amber, there are no previous accounts of fossil mushrooms from these extensive deposits (Bachofen-Echt 1949; Czeczott 1961; Larsson 1978; Weitschat & Wichard 2002).
The present study describes a new genus and species of gilled mushroom in Baltic amber. Fossiliferous amber from the Baltic region was formed in tropical and subtropical forests that covered a large part of northern Europe for some 10 million years during the early Cenozoic (Bachofen-Echt 1949; Larsson 1978; Weitschat & Wichard 2002). The amber was eroded from the original accumulation and redeposited in marine sediments referred to as the ‘blue earth’ layers in the Samland Peninsula. Original age estimates of amber from this region ranged from 35 to 55 million years (Larsson 1978; Weitschat & Wichard 2002). The latest estimate, based on geochemical tests performed on amberbearing sediments, was between 45 and 55 million years (Wolfe et al. 2016).
* Corresponding author. Tel.: þ1541 760 7319; fax: þ1541 757 1639. E-mail address: [email protected] http://dx.doi.org/10.1016/j.funbio.2016.06.008 1878-6146/ª 2016 British Mycological Society. Published by Elsevier Ltd. All rights reserved.
Please cite this article in press as: Poinar G, Jr, A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber, Fungal Biology (2016), http://dx.doi.org/10.1016/j.funbio.2016.06.008
2
G. Poinar
Materials and methods The specimen originated from the Samland Peninsula in the Kalinin District of the Russian Federation (54.7104 N: 20.4522 E). The amber piece was reshaped and polished to a final length of 9 mm, width of 3 mm, and depth of 3 mm. Observations and photographs were made with a Nikon SMZ-10 R stereoscopic microscope and Nikon Optiphot compound microscope with magnification up to 800. Helicon Focus Pro X64 (D-Studio, 22 Krakovska St. Kiev, Ukraine 02094) was used to stack photos for better clarity and depth of field.
Results The specimen is small, but well preserved and includes a complete pileus and attached stipe that is broken at the base. An insect nymph and strand of mammalian hair are associated with the specimen. Phylum Basidiomycota Order Agaricales Gerontomyces lepidotus Poinar, gen. et sp. nov. (Figs 1e4) Description Total length of pileus and remaining stipe ¼ 1.8 mm; pileus brown, bell-shaped, convex, dry, thick, with straight to slightly incurved margin; surface densely scaly with decumbent to erect scales extending outward to 35 mm; pileus diameter, 1.0 mm; lamellae (12e14 in number) white, broad, flat, distant to subdistant, with smooth uniform margins, attached to stipe apex: lamellulae and spores absent; stipe brown, base broken (remaining portion 1.5 mm long and 0.3 mm wide), dry, solid, insertion central, covered
Fig 1 e Gerontomyces lepidotus gen. et sp. nov. with adjacent insect exuviae (arrow) and mammalian hair in Baltic amber. Bar [ 0.5 mm.
Fig 2 e Detail of pileus and stipe of Gerontomyces lepidotus gen. et sp. nov. in Baltic amber. Bar [ 0.2 mm.
Fig 3 e Pileus of Gerontomyces lepidotus gen. et sp. nov. viewed from above showing slightly prolonged area with attached water droplet (arrow). Bar [ 0.3 mm.
Please cite this article in press as: Poinar G, Jr, A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber, Fungal Biology (2016), http://dx.doi.org/10.1016/j.funbio.2016.06.008
Gilled mushroom in Baltic amber
3
Discussion
Fig 4 e Ventral (under) view of Gerontomyces lepidotus gen. et sp. nov. in Baltic amber showing lamellae. Bar [ 0.5 mm.
with decumbent to erect scales extending outward to 30 mm; veil and annulus absent. Holotype: Accession # Myc-12, deposited at the Bone Room, 1573 Solano Ave., Berkeley, CA 94707. Etymology: Generic name from Greek ‘geraios’ ¼ old and ‘mykes’ ¼ mushroom. Specific epithet is from the Greek ‘lepidotos’ ¼ scaly in reference to the dense scaly covering of the fossil. Type locality: Samland Peninsula of the Baltic Sea in the Kalinin District of the Russian Federation. Diagnosis: The combination of small size, dry, brownish, densely scaly surface, bell-shaped, convex, dry pileus, broad, distant to subdistant lamellae, the absence of lamellulae, and a straight, solid stipe lacking a veil and annulus characterize the fossil and distinguish it from previously described fossil gilled mushrooms. While the top of the pileus was originally probably round, it was preserved slightly asymmetrical with one margin extended (Fig 3). This is probably due to moisture that had collected on the extended side, some of which collected as an attached water droplet (Fig 3). Moisture is often represented as a gray film on the surface of fossils in Baltic amber. Adjacent to the mushroom is the exuvia of a nymph of a Phasmatodea (Fig 1). Measuring only 0.6 mm in body length, the nymph has features of Pseudoperla lineata Pictet & Hagen, which is fairly common in Baltic amber (Bachofen-Echt 1949; Larsson 1978). In the same piece of amber is a strand of mammalian hair. The strand is several millimetres in length with a diameter reaching 140 mm.
The evolutionary history of gilled mushrooms is restricted to the relatively few fossils from Cenozoic and Cretaceous amber, with the oldest specimen, Palaeoagaricites antiquus, in Myanmar amber showing an affinity to members of the family Tricholomataceae (Taylor et al. 2015). Due to the absence of spores and the basal portion of the stipe, it is difficult to align Gerontomyces lepidotus gen. et sp. nov. with any modern genus or even family. Thus it is not possible to use the fossil to provide minimum age estimates for the origins of any particular agaric lineage. The combination of a small size, dry, brownish, scaly surface, broad, distant to subdistant lamellae and a straight, solid stipe are not common characters in gilled mushrooms (Lincoff 1981; Arora 1986; McKnight & McKnight 1987). However, some of these features do occur in a few genera in the family Tricholomataceae. Members of the genus Squamanita are small woodland mushrooms that have a scaly pileus and stipe with some erect scales as in the fossil (Arora 1986). However species of Squamanita have a distinct veil. The adjacent nymphal exuviae of a phasmatoid could imply a feeding association since phasmatoids are herbivorous and there are records of extant phasmatoids feeding on mushrooms. Thus there is a possibility that the insect was exploring the pileus of the mushroom just as the latter was covered with resin. Finding the empty exuviae suggests that the nymph made a narrow escape.
Conclusions The Baltic amber mushroom reported here adds an additional degree of diversity among gilled Agaricales that existed in the Cenozoic.
Acknowledgements I thank Diana Mansfield of the Bone Room (1573 Solano Avenue, Berkeley, California 94707) and Alex Brown for facilitating the loan of the present specimen and Roberta Poinar for reading earlier drafts of this manuscript.
references
Arora D, 1986. Mushrooms Demystified, 2nd edn. Ten Speed Press, Berkeley, 959 pp.. € sse. Springer Bachofen-Echt A, 1949. Der Bernstein und seine Einschlu Verlag, Vienna, 204 pp.. Czeczott H, 1961. The flora of the Baltic amber and its age. Prace Muzeum Ziemi (Paleobotaniczne) Warszawa 4: 119e145. Hibbett DS, Grimaldi D, Donoghue MJ, 1995. Cretaceous mushrooms in amber. Nature 377: 487. Hibbett DS, Grimaldi D, Donoghue MJ, 1997. Fossil mushrooms from Miocene and Cretaceous ambers and the evolution of Homobasidiomycetes. American Journal of Botany 84: 981e991.
Please cite this article in press as: Poinar G, Jr, A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber, Fungal Biology (2016), http://dx.doi.org/10.1016/j.funbio.2016.06.008
4
Hibbett DS, Binder M, Wang Z, Goldman Y, 2003. Another fossil agaric from Dominican amber. Mycologia 95: 685e687. Larsson SG, 1978. Baltic Amber e a palaeobiological study. Scandinavian Science Press, LTD, Klampenborg. Lincoff GH, 1981. The Audubon Society Field Guide to North American Mushrooms. Alfred A. Knopf, New York, 926 pp.. McKnight KH, McKnight VB, 1987. A Field Guide to Mushrooms. Houghton Mifflin Company, Boston, 429 pp.. Poinar Jr GO, 2001. Fossil puffballs (Gasteromycetes: Lycoperdales) in Mexican amber. Historical Biology 15: 219e221. Poinar Jr GO, Singer R, 1990. Upper Eocene gilled mushroom from the Dominican Republic. Science 248: 1099e1101. Poinar Jr GO, Buckley R, 2007. Evidence of mycoparasitism and hypermycoparasitism in Early Cretaceous amber. Mycological Research 111: 503e506.
G. Poinar
Poinar Jr GO, Donis da Silva A, Goulart Baseia I, 2014. A gasteroid fungus, Palaeogaster micromorpha gen. & sp. nov. (Boletales) in Cretaceous Myanmar amber. Journal of the Botanical Research Institute of Texas 8: 139e143. Taylor TN, Krings M, Taylor EL, 2015. Fossil Fungi. Elsevier, Inc., Amsterdam, 382 pp.. Weitschat W, Wichard W, 2002. Atlas of Plants and Animals in Baltic amber. Verlag Dr. Friedrich Pfeil, Munich, 256 pp.. Wolfe AP, McKellar RC, Tappert R, Sodhi RNS, Muehlenbachs K, 2016. Bitterfeld amber is not Baltic amber: three geochemical tests and further constraints on the botanical affinities of succinite. Review of Palaeobotany and Palynology 225: 21e32.
Please cite this article in press as: Poinar G, Jr, A gilled mushroom, Gerontomyces lepidotus gen. et sp. nov. (Basidiomycota: Agaricales), in Baltic amber, Fungal Biology (2016), http://dx.doi.org/10.1016/j.funbio.2016.06.008