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Notes on the early ouralensis de Verneuil, 1845

Anthony J. Wright University of Wollongong, [email protected]

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Recommended Citation Wright, Anthony J., "Notes on the early devonian brachiopod Leptaena ouralensis de Verneuil, 1845" (2013). Faculty of Science, Medicine and Health - Papers: part A. 1198. https://ro.uow.edu.au/smhpapers/1198

Research Online is the open access institutional repository for the University of Wollongong. For further information contact the UOW Library: [email protected] Notes on the early devonian brachiopod Leptaena ouralensis de Verneuil, 1845

Abstract On the basis of the type material, an attempt is made to review the widely reported brachiopod Leptaena ouralensis de Verneuil, 1845, much more commonly known as Megastrophia uralensis (de Verneuil, 1845). Assignment of this to Megastrophia is still tentative, and many reports of the species, especially those based on crack-out material, must remain doubtful. The species has an established narrow geographic and stratigraphic range to the upper (upper Lower Devonian) of mostly Siberia, Mongolia and China.

Keywords Megastrophia uralensis, ouralensis, brachiopod, stropheodontid, biogeography, Emsian, Devonian, Siberia, China, GeoQuest

Disciplines Medicine and Health Sciences | Social and Behavioral Sciences

Publication Details Wright, A. J. (2013). Notes on the early devonian brachiopod Leptaena ouralensis de Verneuil, 1845. Memoirs of the Association of Australasian Palaeontologists, 44 87-93.

This journal article is available at Research Online: https://ro.uow.edu.au/smhpapers/1198 Notes on the Early Devonian brachiopod Leptaena ouralensis de Verneuil, 1845

ANTHONY J. WRIGHT

WRIGHT, A.J., 2013:04:26. Notes on the Early Devonian brachiopod Leptaena ouralensis de Verneuil, 1845. Memoirs of the Association of Australasian Palaeontologists 44, 87-93. ISSN 0810-8889.

On the basis of the type material, an attempt is made to review the widely reported brachiopod Leptaena ouralensis de Verneuil, 1845, much more commonly known as Megastrophia uralensis (de Verneuil, 1845). Assignment of this species to Megastrophia is still tentative, and many reports of the species, especially those based on crack-out material, must remain doubtful. The species has an established narrow geographic and stratigraphic range to the upper Emsian (upper Lower Devonian) of mostly Siberia, Mongolia and China.

A.J. Wright ([email protected]), School of Earth and Environmental Sciences, University of Wollongong, Wollongong, New South Wales 2522, Australia. Received 8 October 2012.

Keywords: Megastrophia uralensis, ouralensis, brachiopod, stropheodontid, biogeography, Emsian, Devonian, Siberia, China.

THE ORIGINAL type material of Leptaena ouralensis de Phylum BRACHIOPODA Duméril, 1806 Verneuil, 1845 (p. 220, pl. XIV, fig. 1a-b-c-d) was collected Class RHYNCHONELLATA Williams et al., 1996 from Isvioska, near Nizhnij Turinsk, on the east side of the Order Öpik, 1934 Ural Mountains of Russia by the expedition of Roderick Impey Murchison, Édouard de Verneuil and Le Comte Megastrophia Caster, 1939 Alexandre Keyserling in 1841 (see Collie & Diemer 2004). De Verneuil (1845) stated that the first fragmental material Remarks. This stropheodontid was discussed by of the species was collected on the banks of the Serebrianka Williams (1953) who recognised the nominotypical subgenus River, a tributary of the Tchusovaya River, and more material and Megastrophia (Protomegastrophia) Caster, 1939. Harper was collected on the Isvioska River. & Boucot (1978a, b) also discussed the genus, erecting During a brief visit to the palaeontological museum of the the new subgenus M. (Megastrophiella) and recognising Université Claude Bernard Lyon 1 in Villeurbanne, France, in Protomegastrophia as a subgenus of Brachyprion. As the September 2011, I found four de Verneuil specimens (2052, latter authors differentiated the genera on the basis of features 2070 and 2077; on the board with this last number were of the cardinal process lobes, the present type material does two ventral valves) in addition to the four I had previously not permit confident subgeneric assignment. borrowed and photographed (2066, 2067, 2068, 2069). These four digit numbers are those cited in the priceless original de Megastrophia (Megastrophia) Caster, 1939 Verneuil fossil catalogue held by the museum; these eight rock specimens have now been renumbered and catalogued. Type species. (Stropheodonta) concava Hall, The de Verneuil catalogue lists eight specimens from the 1857. Isvioska, but none from the Serebrianka. Most specimens still bear a distinctive white paper label with a four digit Megastrophia (Megastrophia?) uralensis (de Verneuil, number and the words ‘coll. de Verneuil’ as well as a white 1845) (Figs 1-2) paper label with the word ‘Isvioska’. The eight shells occur on eight pieces of rock, including one which bears two 1845 Leptaena ouralensis de Verneuil, p. 220-222, pl. 14, shells (2069A-B), and two other rocks (2066, 2070) which figs 1a-d. are counterparts. All are crack-out specimens retaining little 1887 Strophomena uralensis Verneuil; Chernyshev, p. 108. internal morphology. 1930 Stropheodonta uralense Verneuil; Nalivkin, p. 69, pl. 1, figs 1a-c, 12a-c, 15a-b. SYSTEMATICS 1947 Stropheodonta uralensis Verneuil, 1845; Nalivkin, pl. All this de Verneuil type material is held in the palaeontology 14, figs 3-4. museum at the Université Claude Bernard Lyon 1 at 1955 Stropheodonta uralensis (Verneuil); Rzhonsnitskaya, Villeurbanne, France; the prefix EM pertains to this p. 244, pl. 54, fig. 3a-b. collection. 1961 Megastrophia uralensis (Verneuil, 1845); Kul’kov in Gratsianova et al., p. 437, table D-67, fig. 3. 88 AAP Memoir 44 (2013)

Figure 1. Type material of Megastrophia uralensis (de Verneuil, 1845). A, view of two specimens EM 25053A-B, x1.5. B-C, lectotype dorsal valve EM 25053A in dorsal and posterior views respectively, showing thin median ridge, small subcircular muscle field, a linear arrangement of nodes anteriorly and parvicostellate sculpture, x 1.5. D, exterior of ventral valve 25053 B showing nervostrophid sculpture, x 2.25. E, EM 25050, shell broken largely along concave external surface of dorsal valve, but showing interarea of ventral valve, x 1.4. F, EM 25052, shell broken largely along convex internal surface of dorsal valve showing median ridge and interarea of ventral valve, x 1.8. G, lateral view of deformed ventral valve EM 25051, x 0.9. AAP Memoir 44 (2013) 89

1962 Megastrophia uralensis (Verneuil); Tyazheva, p. 41, tendencies). EM 25052, the fourth specimen (not illustrated pl. 9, fig. 6-7. by de Verneuil), is a dorsal valve broken so as to show some ?1962 Megastrophia uralensis sibirica subsp. nov.; Krilova, external sculpture and a slender median ridge internally. p. 47, pl. 2, fig. 8, pl. 3, figs 4-5. Specimens photographed by Dr Abel Prieur are EM 25054 1974 Megastrophia uralensis (Verneuil); Wang et al., pl. (ventral valve), EM 25055 (ventral valve) and EM 25056 11, figs 13-25. (large ventral valve). 1975 Megastrophia uralensis (Verneuil, 1845); Gratsianova, p. 26-27, pl. 4, figs1-8. Description. Strongly concavo-convex, thin, rather alate 1977 Megastrophia uralensis (Verneuil, 1845); Gratsianova shells up to about 75 mm wide and 40 mm long. Sculpture & Schischkina, pl. 21, figs 1-5; pl. 22, figs 104; pl. 23, on both valves parvicostellate with 4-6 fine costellae figs 1-7. between the major ribs, which tend to be discontinuous 1977 Leptaena uralensis; Nalivkin, p. 266. on one specimen (i.e. the nervate condition shown by EM ?1977 Megastrophia semispheroidea Hou & Xian; Yang et 25053B, see Fig. 1D). Apsacline ventral interarea several mm al., p. 316. high, weakly curved, and strongly denticulate and striated; 1978 Megastrophia uralensis (Verneuil); Gratsianova, pl. pseudodeltidium apparently simple. No information on 53, fig. 15; pl. 54, figs 9-10; pl. 56, fig. 16; pl. 57, fig. 1. ventral valve interior. Dorsal valve with a slender median 1979 Megastrophia uralensis (Verneuil, 1845); Wang & Zhu, ridge which extends to about 70% of valve length and a small p. 18, pl. 1, figs 18-20; pl. 20, figs 13-17; pl. 34, fig. 5. subcircular adductor field with low rounded bounding ridges 1981 Megastrophia uralensis (Vern.); Sapelnikov & Mizens, located well posterior of midlength. An irregular oblique p. 13, pl. 2, fig. 12. row of nodes, flanking the median ridge, arises posterior 1981 Megastrophia uralensis (Vern.); Sapelnikov et al., of midlength and extends to at least 70% of dorsal valve pp. 8, 37. length. Cardinal process lobes not seen, but apparently small 1991 Megastrophia uralensis (Verneuil, 1845); Menakova (cf. Gratsianova 1978, pl. 56, fig. 16). Marginally, dorsal in Dshalilov, p. 182, pl. 92, fig. 1. valve has low, linear, rounded ribs internally, as illustrated 1996 Megastrophia uralensis (Verneuil); Alekseeva et al., by Alekseeva et al. (1996, pl. 1, fig. 5). Pseudopunctae p. 31, pl. 1, figs 5-7. abundant. 2001 Megastrophia uralensis (de Verneuil); Talent et al., p. 38, 40, 55-56, 71, 77, 110, 143. Selection of lectotype. In response to my questions about this 2003 Megastrophia uralensis (Verneuil, 1845); Afanasjeva species, the late Dr Rimma Gratsianova sent me in 1981 inter et al., p. 91, pl. 33, figs 1-2. alia copies of two plates and some text that appeared to be 2007 Megastrophia uralensis (Verneuil, 1845); Kim et al., from a publication by Breivel & Breivel (1978) sent to her by p. 255, pl. 85, figs 28-32. the authors. In this text, Breivel & Breivel selected a neotype 2011 Megastrophia uralensis (Verneuil, 1845); Kim, fig. 3, for M. uralensis; this was presumably deemed necessary as pl. 3, figs 28-29. Nalivkin (1977) had reported that he was unable to identify 2012 Megastrophia uralensis; Talent et al., fig. 2, p. 824. the original type material (see below for translation of parts of the Nalivkin paper). However, Dr Larissa Mizens (personal Type material. EM 25053A (de Verneuil number 2069) is communication, June 2010) has informed me that the 1978 here selected as lectotype from the type series of de Verneuil; publication never appeared. A lectotype is selected here it is a dorsal valve (Fig. 1A-C) broken largely along the (see above) from the de Verneuil type series, even though external surface, but retaining some vestiges of internal the specimen is less than ideally preserved and shows little structures. The rock bearing this specimen also bears a distinctive morphology. ventral valve, now numbered EM 25053B. Nalivkin (1977, p. 263) commented on the original The type series has been renumbered as follows: 2052 = (figured?) de Verneuil material as follows: Unfortunately,“ EM 25056; 2066 = EM 25050A, 2067 = EM 25052; 2068 = a part of Verneuil’s materials [have been] sometimes taken EM 25051; 2069 = EM 25053A-B (one dorsal valve and one out of cabinets, some examples appear to be lost, although ventral valve, not of the same shell); 2070 = EM 25050B; there is a hope of finding them in other parts of the same 2077A = EM 25054; 2077B = EM 25055. store or even in other museums. Specimens illustrated by de Verneuil appear to be: The originals on the plates were not selected by Verneuil” EM 25056 (his fig. 1a-b, a large ventral valve with some [and] “Leptaena uralensis. Many (examples), typical, but I remaining pieces of the dorsal valve of the same shell), could not detect originals. Locality ‘Izviostka’.” (kindly perhaps with additions from EM 25055; EM 25050A (his translated by Dr Larissa Mizens, Yekaterinburg). fig. 1c; 2066, counterpart of 2070; this specimen bears inked lines accentuating radial sculpture); and EM 25054 Species name. The original spelling of the species name (his fig. 1d). The illustrations of de Verneuil appear to be (ouralensis) has, as far as I am aware, been used only by the diagrammatic, so confident identification of actual specimens original author. The specific name uralensis has been used from his illustrations (no catalogue numbers being cited in exclusively ever since and is retained here. The original the caption) is not possible. It seems possible that figures spelling could be retained here in accordance with ICZN incorporate features from more than one specimen; further, Article 32. However, according to ICZN Article 33.2.3.1, the some damage or preparation over the years may have subsequent usage is treated as a justified emendation and the substantially altered the appearance of specimens. specific nameuralensis is accepted here by me. The four specimens photographed by me are: the lectotype dorsal valve EM 25053A, but also showing the ventral valve Generic assignment. Assignment of uralensis to Megastrophia interarea); EM 25051 (a large, deformed ventral valve); or (more recently) Megastrophia (Megastrophia) has been EM 25053B (a ventral valve broken along the external accepted without question for at least 50 years, being first surface and showing well preserved sculpture with nervate suggested by Gratsianova et al. (1961). Megastrophia was 90 AAP Memoir 44 (2013)

Figure 2A-E, Megastrophia uralensis (de Verneuil, 1845). A-B, plan (x 1.8) and posterior (x 2.1) views of exterior of ventral valve EM 25054. C, plan view of exterior of ventral valve EM 25055, x 1.3. D-E, plan and posterior views of exterior of EM 25056, both x 1.2. Photography by A. Prieur, Collections of Geology, UCBL1. extensively discussed by Harper & Boucot (1978b) and it all belongs to uralensis, further characters of uralensis seems possible that uralensis belongs to the nominotypical include a short, flabellate ventral muscle field with a subgenus in being large and (see below) in having cardinal prominent median ridge and the distinctive structure of the process lobes joined basally. As no critical internal details cardinal process lobes (see below). Material illustrated by are preserved (although preparation of some type material of Alekseeva et al. (1996, pl. 33, fig. 5) from the NE part of the uralensis might expose these) this must await resolution by USSR includes a ventral valve with internal ribbing similar further study. The slight nervate tendency in sculpture does to that of the uralensis types, but confident comparison not really resemble any of the nervostrophiids illustrated of the type uralensis with the illustrated dorsal valve by Harper & Boucot (1978a) and much more information (Alekseeva et al. 1996, fig. 6) with a subcircular muscle field is needed on this aspect of the morphology of uralensis. is inconclusive due to the poor preservation of the lectotype of uralensis; their illustrated specimen does, however, show Remarks. It is clear that de Verneuil (1845, p. 222) erred in cardinal process lobes joined basally, suggesting assignment identifying the specimens illustrated in his plate 14, figs 1a-b to M. (Megastrophia). and d as dorsal valves, and his plate 14, fig. 1c as a ventral valve. His fig. 1c clearly shows the ventral interarea with Discussion. It is appropriate to place here a cautionary delthyrium, but most of the specimen shows the (concave) note about the above synonymy. Much crack-out material exterior of the dorsal valve. His fig. 1d clearly shows the assigned to this species for more than 130 years (excluding parvicostellate sculpture; he remarked on the fine costellae the original description) is probably indeterminate to both between the major ribs. species and genus, showing neither external sculpture nor If the material illustrated by Gratsianova (1975), internal features. The status of the Russian subspecies M. Gratsianova & Shischkina (1977 and Alekseeva et al. (1996) uralensis siberica and the Chinese species M. semispheroidea AAP Memoir 44 (2013) 91 are yet to be determined. seems questionable. Talent et al. (1987, pp. 230-231) Notwithstanding the very limited information that can be reported Megastrophia pseudointerstrialis (Hou) from Nei gleaned from the type material, there is a lot of other material Monggol. Harper & Boucot (1978b) referred manchurica to that has been identified asMegastrophia uralensis. Notable ?Strophodonta. Xian (1988) described M. ertaiziensis from exceptions to this largely indeterminate crack-out material the Longmenshan Mountains. Mendbajar (1994) described are the internals illustrated by Alekseeva et al. (1996, pl. 1, the new species M. zhogtensis from the Lower Devonian figs 5-7) and Gratsianova (1975, pl. 4, figs 1, 3-5: ventral (Emsian) of eastern Mongolia (for details see Afanasjeva et valves only). Gratsianova & Schischkina (1977, pls 21-22) al. 2003, p. 91-92). One interesting report is “Megastrophia illustrated “M. (M.) uralensis (Vern.)” material with good ex. gr. concava (Hall)” from Vietnam (Nguyen et al. 1980, parvicostellate sculpture and ventral internal moulds, and p. 108, pl. 41, figs 1, 3, 5), based on a ventral internal mould Gratsianova & Schischkina (1977, pl. 34) illustrated M. with a good muscle field; elsewhere in the same volume (p. (M.) bobilevi Schischkina sp. nov. with internal moulds of 277), “Megatrophia (sic.) ex. gr. dubrovensis Rzhon.” is both valves. reported from the Miale Suite, to which a late Early to early Middle Devonian age is assigned. Palaeogeographic distribution. The invaluable compilation Reports of Megastrophia from Australia are yet to be of Devonian brachiopod distribution by Talent et al. (2001) fully documented but the genus, along with the brachiopod and followed up by Talent et al. (2012) has greatly clarified Zdimir and a serotinus conodont fauna, has been reported the geographic distribution of reports of M. uralensis; from Mount Podge, Queensland (Talent & Yolkin 1987, relevant references for the following comments are given Talent & Mawson 1994) and Mount Frome, New South by Talent et al. (2001). They reported the species from Wales (Wright in Percival et al. 2010). numerous regions, almost entirely from the late Emsian of Siberia, Mongolia and China, as follows: ACKNOWLEDGEMENTS 1. Laurussia: Biya Gorizont and the Afonin Gorizont, west I thank Dr Abel Prieur (Villeurbanne) for allowing me to of the Urals; borrow some of de Verneuil’s type material and, in September 2. Kazakhstania: East of the Urals, Karpinsk Gorizont and 2011, to access the collections in Villeurbanne and locate the Tal’tiy Gorizont; other type materials of this species; all this type material is 3. Siberia: Salair, Shanda Gorizont; Gornyi Altai, Kuvash the basis of this paper. Dr Prieur also allowed me access to Beds; the de Verneuil hand-written catalogue, catalogued all these 4. South China; de Verneuil specimens and most graciously photographed 5. Karakorum; and the specimens discovered in 2011 for me. I particularly 6. South Fergana. thank Dr Larissa Mizens (Yekaterinburg) for very helpful According to Alekseeva (1992), the maximum correspondence; establishing that the Breivel & Breivel palaeolatitude for the above continental plates was less paper was never published, and that, therefore, no neotype than 45° N. Pedder & Oliver (1990, p. 267) concluded, with was selected; and for translating parts of Nalivkin (1977). I reference to the reconstruction of Scotese (2001) that “it is am greatly indebted to other colleagues who have assisted questionable that the original latitude of any large northern with discussion or literature: Firuza Salimova and Aleksey hemisphere Devonian coral fauna would have exceeded Kim (Tashkent), Olga Obut and the late Rimma Gratsianova 45°”. Further comments on Devonian biogeography, based (Novosibirsk), Tatjana Grunt and Anne Madison (Moscow), on fish, were made by Young (1990). Chen Xiqin (Nanjing), and John Talent (Sydney); without There are many important reports and illustrations their assistance this study could not have been made. This of megastrophiid from China. Hou (1981) is a contribution to IGCP Project 591 (The Early to Middle recognised the Zdimir community with Megastrophia Paleozoic Revolution) and IGCP Project 596 (Climate uralensis in what he considered Middle Devonian strata in his change and biodiversity patterns in the mid-Palaeozoic). I am South China Province, but it is clear that such megastrophiid grateful to Professor Colin Murray-Wallace for provision of forms occur in Mongolia and, indeed across western Siberia, facilities in the School of Earth and Environmental Sciences, even occurring on the west side of the Urals (Chernyshev University of Wollongong where my research has been 1885). Hou & Wang (1996) considered that the Inner supported by the Geoquest Research Centre and the Linnean Mongolian Devonian fauna was close to that of North China Macleay Fellowship. but made no mention of Megastrophia. Limited material from South China was assigned to uralensis by Wang et al. 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