Nomenclatural Notes on the Eurypterid Family Carcinosomatidae

Zoosyst. Evol. 88 (1) 2012, 19–24 / DOI 10.1002/zoos.201200003

Nomenclatural notes on the eurypterid family Carcinosomatidae

Jason A. Dunlop*,1 and James C. Lamsdell2

1 Museum fçr Naturkunde Berlin, Leibniz-Institut fçr Evolutions- und Biodiversitåtsforschung an der Humboldt-Universitåt zu Berlin, Invalidenstraße 43, 10115 Berlin, Germany 2 Department of Geology and Paleontological Institute, University of Kansas, 1475 Jayhawk Boulevard, Lawrence, KS 66045, U.S.A.

Abstract

Received 19 August 2011 The genus level systematics of the eurypterid family Carcinosomatidae (Chelicerata Accepted 6 September 2011 Eurypterida) is briefly reviewed, with particular reference to some nomenclatural prob- Published 16 March 2012 lems associated with homonyms and their replacement names. Eusarcus scorpionis Grote & Pitt, 1875 is the type species of both Eusarcana Strand, 1942 and Paracarci- Key Words nosoma Caster & Kjellesvig-Waering, 1964. Although Strand’s name has been completely overlooked in the eurypterid literature, it remains the oldest available (and valid) Eurypterida name associated with Eusarcus scorpionis. It should be noted that Strand replaced a Fossil homonym under circumstances when this was not really necessary, but the Principle of Nomenclature Priority means that we are forced to reclassify three Paracarcinosoma species under his Homonyms genus name as Eusarcana scorpionis (Grote & Pitt, 1875), Eusarcana acrocephala Embrik Strand (Semper, 1898) and Eusarcana obesa (Woodward, 1868); all comb. nov.

Introduction Systematics

The eurypterid family Carcinosomatidae (Chelicerata: Family Carcinosomatidae Størmer, 1934 Eurypterida) includes a number of quite large and 1934 Carsinosomidae Størmer, p. 104. highly distinctive fossil arthropods. Its constituent spe- 1955 Carcinosomatidae: Størmer, p. 35 [nom. correct.] cies are characterised by spiny forelegs, large paddles, a broad and rounded preabdomen, and a slender postab- Remarks. The family group name was introduced, as domen ending in a curved telson. In this sense, they Carcinosomidae, by Størmer (1934) for the genera Car- appear quite scorpion-like (Fig. 1) and were presumably cinosoma Claypole, 1890 [including in this sense Eu- one of the groups which contributed to the eurypterids’ sarcus Grote & Pitt 1875; see below], Mixopterus Rue- common name of ‘sea scorpions’. Within Carcinosoma- demann, 1921, Echinognathus Walcott, 1882 and tidae, a number of genera have been proposed; some of Megalograptus Miller, 1874. The latter three genera them unfortunately representing preoccupied names were subsequently transferred to their own families, (see below). This has led to a degree of nomenclatural whereby family authorship should effectively be cred- confusion and inconsistencies in the literature. In parti- ited to Caster & Kjellesvig-Waering in Størmer (1955). cular, we were recently made aware of the fact that one In detail, the new family names were first mentioned in of the oldest published names, Eusarcus scorpionis the Treatise on Invertebrate Paleontology, albeit draw- Grote & Pitt, 1875, is ostensibly the type species of ing on a manuscript by Kenneth Caster and Erik Kjel- two separate genera: the widely overlooked Eusarcana lesvig-Waering which was supposedly in press at the Strand, 1942 and the more familiar Paracarcinosoma time of writing, but in fact came out much later as Cas- Caster & Kjellesvig-Waering, 1964. Here, we attempt ter & Kjellesvig-Waering (1964). Thus in 1955 Mixop- to resolve this issue as part of a brief review the basic terus was assigned to Mixopteridae; Echinognathus and systematic structure of the family Carcinosomatidae. Megalograptus to Megalograptidae. For completeness, the species composition of all four currently recognised carcinosomatid genera is listed below.

* Corresponding author, e-mail: [email protected]

# 2012 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 20 Dunlop, J. A. & Lamsdell, J. C.: Nomenclatural notes on Carcinosomatidae

Figure 1. Specimen (non-type) of Eusarcus (now Eusarcana) scorpionis (Grote & Pitt, 1875) from the Silurian ‘Bertie Waterlime’ of New York, USA; courtesy of the American Museum of Natural History, New York (image taken by Steve Thurston).

Carcinosoma Claypole, 1890 Type species. Eurysoma newlini Claypole, 1890a [= C. ingens Clay- pole, 1894]. 1890a Eurysoma Claypole, p. 259 [preoccupied by Eurysoma Dejean, 1831 (Coleoptera)] Other included species. C. harleyi Kjellesvig-Waering, 1961b, C. liber- 1890b Carcinosoma Claypole, p. 400 [replacement name] tyi Copeland & Bolton, 1960, ?C. punctatum (Salter in Huxley &

museum-zoosyst.evol.wiley-vch.de # 2012 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 88 (1) 2012, 19–24 21

Salter, 1859); C. scorpioides (Woodward, 1868) [= Pterygotus rani- sumably mentioned the Eusarcus problem, and also re- ceps Woodward, 1868], C. scoticus (Laurie, 1899), ?C. spiniferum ceived confirmation from Strand that Carcinosoma was Kjellesvig-Waering & Heubusch, 1962. not preoccupied and still available. For reasons best known to himself, Strand subsequently proposed – Eocarcinosoma Caster & Kjellesvig-Waering, 1964 without comment – a replacement name for Eusarcus; namely Eusarcana Strand, 1942. Why he felt the urge 1964 Eocarcinosoma Caster & Kjellesvig-Waering, p. 314. to replace a name which had already been dealt with Type and only species. Eocarcinosoma batrachophthalmus Caster & satisfactorily by Størmer’s nomenclatural act is hard to Kjellesvig-Waering, 1964. fathom, but contemporary critiques of Strand’s ap- proach to systematics (Mortensen 1929; Richter 1929) Eusarcana Strand, 1942 and his apparent desire to have as many taxon names associated with his own authorship as possible probably 1875 Eusarcus Grote & Pitt, p. 1 [preoccupied by Eusarcus Perty, goes some way towards explaining this. In this context, 1833 (Opiliones)] Strand also gets a mention in the review by Evenhuis 1942 Eusarcana Strand, p. 387 [replacement name] (2008) of the so-called ‘Mihi itch’, or its German var- 1964 Paracarcinosoma Caster & Kjellesvig-Waering, p. 312. iant ‘Mihisucht’ – a term Strand apparently coined Type species. Eusarcus scorpionis Grote & Pitt, 1875. himself in response to Mortensen (1929)! This ‘itch’ is Included species. E. acrocephala (Semper, 1898), E. obesa (Wood- essentially an insatiable desire to name taxa. ward, 1868). Strand’s name was completely overlooked. Eusarcana Remarks. Eusarcus Grote & Pitt, 1875 was proposed has not been mentioned in any subsequent eurypterid for a eurypterid from the Buffalo Waterlime of New publications, including summary works such the Trea- York State, USA with Eusarcus scorpionis Grote & Pitt, tise (Størmer 1955), the systematic overview of Toller- 1875 as the type species by original designation. The ton (1989), or the online summary of nomenclature by genus name is preoccupied. Eusarcus Perty, 1833 was Dunlop et al. (2011). As long as only Carcinosoma re- already in use for (living) laniatorid harvestmen (Ara- mained in use this was not really an issue and Strand’s chnida: Opiliones: Gonyleptidae); a genus recently re- name could effectively be ignored. However, Caster & vised by Hara & Pinto da Rocha (2010). The homonym Kjellesvig-Waering (1964), in their major revision of was not initially recognised and Eusarcus continued to Mixopteroidea, created a new genus Paracarcinosoma be used for eurypterids throughout the late 19th and Caster & Kjellesvig-Waering, 1964 with Eusarcus scor- early 20th centuries (e.g. Clarke & Rudemann 1912; pionis as the type species. Eusarcus had effectively Kindle 1913; Woodward 1913; Williams 1915; O’Con- been revalidated, but Strand’s Eusarcana was not men- nell 1916; Ruedemann 1919; Diener 1924). tioned by Caster & Kjellesvig-Waering and they were Another scorpion-like eurypterid was described as presumably unaware of its existence. Indeed, Caster & Eurysoma newlini Claypole, 1890a from Kokomo in In- Kjellesvig-Waering (1964: 312–313) simply introduced diana, USA. Eurysoma Claypole, 1890a is also preoc- the name Paracarcinosoma as a new genus and made cupied and the name has been introduced independently no mention of any previous nomenclatural discussions. on at least six occasions; the oldest of which was for a Paracarcinosoma has been used by all subsequent beetle Eurysoma Dejean, 1831 (Insecta: Coleoptera). authors working on carcinosomatid eurypterids (Stør- Claypole quickly recognised the homonym (Fig. 2) and mer 1974; Copeland & Bolton 1985; Tollerton 1989; proposed Carcinosoma Claypole, 1890b as a replace- Ciurca 1990; Braddy & Dunlop 2000; Braddy et al. ment name for his genus – again with Eurysoma new- 2002; Tetlie 2007; Tetlie et al. 2008; Lamsdell & Braddy lini as its type species. Clarke and Ruedemann (1912) 2010; Dunlop et al. 2011) – with the exception of compared Carcinosoma with Eusarcus and concluded Ritchie (1968) who still considered the Paracarcinoso- in their monograph that the two genera were synonyms. ma species to belong to Carcinosoma. Priority went to Grote & Pitt’s older name, Eusarcus, We are thus faced with the following nomenclatural whereby the fact that it was also a homonym had still problem for Paracarcinosoma scorpionis. Does the not been recognised. Principle of Priority force us to revert to the next oldest In his discussion of his new family Carcinosomati- available name and treat Eusarcana scorpionis (Grote dae, Størmer (1934) now realised that Eusarcus was & Pitt, 1875) as the correct name? Could the Interna- preoccupied by a harvestman and (correctly) proposed tional Commission on Zoological Nomenclature (ICZN) that the best course of action was to go to the next old- be asked to rule in favour of suppressing the barely est available and valid name: Claypole’s Carcinosoma. known Eusarcana and replacing it with the more wide- While preparing this 1934 paper Størmer consulted spread Paracarcinosoma? Or can we argue that Strand’s Embrik Strand (Fig. 3), who at the time was investing replacement name was unnecessary and in some way an inordinate amount of effort into nomenclatural is- did not constitute a valid nomenclatural act? sues; replacing homonyms and (less helpfully) applying Taking these in reverse order, Strand (1942: 387) ex- new species names to incomplete and/or poorly pre- plicitly wrote “ Eusarcus Grote & Pitt, Bull. Buffalo served fossils figured in the literature (cf. Dunlop & Soc. III, p. 1 (1875)y (nec Eusarcus Perty 1833) benen- Tetlie 2006 for two eurypterid examples). Størmer pre- ne ich Eusarcana m. (Gigantostraca).” Taking Arti- y

# 2012 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 22 Dunlop, J. A. & Lamsdell, J. C.: Nomenclatural notes on Carcinosomatidae

Figure 2. Timeline of the major events affecting the nomenclature of both Eusarcus scorpionis Grote & Pitt, 1875 and Eurysoma newlini Claypole, 1890a respectively. cle 13.3 of the ICZN, an available genus group name With respect to conservation of usage, we fear that published after 1930 must be accompanied by the fixa- while Eusarcana is a long-forgotten name, the ten or so tion of a type species – which Strand does not do – or citations using Paracarcinosoma (see above) – and the be expressly proposed as a new replacement name: fact that this only affects a minor aspect of eurypterid which he does do by saying “I name it Eusarcana systematics – are unlikely to be sufficient to warrant a m[ihi] (= me).” Although Strand did not formallyy as- successful application to the ICZN to suppress Eusar- sign a type species for Eusarcana, Article 67.8 states cana. Given that Eusarcana and Paracarcinosoma both that the type species of a replacement name must be appear to be available names, and given that both have fixed as the same type species for the original name; the same type species, we fear that that the Principle of i.e. we are obliged to accept Grote & Pitt’s scorpionis Priority (Article 23) forces us to adopt Strand’s genus as the type species of Eusarcana. While Strand’s name as the oldest valid name available (Fig. 2) and the (1942) replacement name was, at the time of writing, three species currently assigned to Paracarcinosoma not really necessary we see no provision in the ICZN must be transferred, all as comb. nov., to Eusarcana. It code which formally invalidates it. remains a further example of Embrik Strand’s less than

museum-zoosyst.evol.wiley-vch.de # 2012 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Zoosyst. Evol. 88 (1) 2012, 19–24 23

turkunde, Berlin) for the portrait of Embrik Strand and for informa- tion about his academic career. The reviewers, Mark Harvey and Erik Tetlie, are thanked for helpful comments.

References

Braddy, S. J. & Dunlop, J. A. 2000. Early Devonian eurypterids from the Northwest Territories of Arctic Canada. – Canadian Journal of Earth Science 37: 1167–1175. Braddy, S. J., Selden, P. A. & Truong, D. N. 2002. A new carcinosomatid eurypterid from the Upper Silurian of Northern Vietnam. – Palaeontology 45: 897–915. Caster K. E. & Kjellesvig-Waering, E. N. 1964. Upper Ordovician eurypterids of Ohio. – Palaeontographica Americana 4 (32): 297– 358. Ciurca Jr., S. J. 1990. Eurypterid biofacies of the Silurian-Devonian evaporite sequence: Niagra peninsula, Ontario, Canada and New York. – New York State Geological Association, 62nd Annual Meeting and Guidebook: pp. 1–30. Clarke, J. N. & Ruedemann, R. 1912. The Eurypterida of New York. – New York State Museum, Memoir 14: 1–439. Claypole, E. W. 1890a. Palaeontological notes from Indianapolis (A.A.A.S.) Pterichthys – Castoroides – Eurysoma g.n. – Ameri- can Geologist 6: 255–260. Claypole, E. W. 1890b. Carcinosoma newlini. – American Geolo- gist 6: 400. Copeland, M. J. & Bolton, T. E. 1985. Fossils of Ontario Part 3: The eurypterids and phyllocarids. – Life Sciences Miscellaneous Pub- lications, Royal Ontario Museum, University of Toronto Press, 48 pp. Dejean, P. F. M. A. 1831. Species general des Coleopteres, de la col- lection de M. le Comte Dejean. Tome cinquieme. Mequignon- Marvis, Paris, 883 pp. Diener, C. 1924. Eurypterida. In Diener, C. (ed.). Fossilium Catalogus I: Animalia, 25. W. Junk, Berlin: pp. 1–26. Figure 3. Portrait of Embrik Strand (1876–1947); Museum fçr Dunlop, J. A. & Tetlie, O. E. 2006. Embrik Strand’s eurypterids. – Naturkunde Berlin, historical image and document archives (of- Neues Jahrbuch fçr Geologie und Palåontologie, Monats- ficial seal: MfN, HBSB). Inventory: Zool. Mus. Signature: B I/ hefte 2006 (11): 696–704. 486. Strand worked as a scientific assistant in entomology and Dunlop, J. A., Penney, D. & Jekel, D. 2011. A summary list of fossil arachnology at the Berlin Museum from 1907–1912; from spiders and their relatives. In Platnick, N. I. (ed.). The world spi- 1923 onwards he was professor of zoology in Riga. der catalog, version 12.0 American Museum of Natural History, online at http://research.amnh.org/entomology/spiders/catalog/in- dex.html doi: 10.5531/db.iz.0001 Evenhuis, N. L. 2008. The “Mihi itch” – a brief history. – Zoo- helpful contributions to zoological nomenclature taxa 1890: 59–68. through creating whole swathes of (sometimes unneces- Grote, A. R. & Pitt, W. H. 1875. I. Description of a new Crustacean sary) names in his own journal ‘Folia Zoologica et Hy- from the Water Lime Group at Buffalo. – Bulletin of the Buffalo drobiologica’, which would later re-emerge to create Society of Natural Sciences 3: 1–2. avoidable systematic problems. Hara, M. R. & Pinto-da-Rocha, R. 2010. Systematic review and cla- distic analysis of the genus Eusarcus Perty, 1833 (Arachnida, Opi- liones, Gonyleptidae). – Zootaxa 2698: 1–136. Rhinocarcinosoma Novojilov, 1962 Kindle, E. M. 1913. The age of the eurypterids of Kokomo, Indiana. – American Journal of Science 36: 282–288. 1962 Rhinocarcinosoma Novojilov, p. 413. Lamsdell, J. C. & Braddy, S. J. 2010. Cope’s rule and Romer’s theory: Type species. Eusarcus vaningeni Clarke & Ruedemann, 1912. patterns of diversity and gigantism in eurypterids and Palaeozoic vertebrates. – Biology Letters 6: 265–269. Included species. R. cicerops (Clarke, 1907), R. dosonensis Braddy, Mortensen, T. 1929. Some remarks and proposals concerning zoologi- Selden & Doan Nhat, 2002. cal nomenclature. – Verhandlungen des X. Internationalen Zoolo- gen-Kongresses 1927: 1569–1574. Novojilov, N. J. 1962. Order Eurypterida. In Orlov, Y. A. (ed.). Funda- Acknowledgements mentals of Paleontology, Volume 9; Arthropoda, Tracheata, Cheli- cerata. Akademiya Nauk SSSR Publishers, Moscow: pp. 404–423. We are grateful to Joel Hallan (Austin, Texas) for bringing the Eusar- [in Russian] cus scorpionis problem to our attention, Steve Thurston (New York) O’Connell, M. 1916. The habitat of the Eurypterida. – Bulletin of the for photographing specimens and Sabine Hackethal (Museum fçr Na- Buffalo Society of Natural History 11: 1–277.

# 2012 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-zoosyst.evol.wiley-vch.de 24 Dunlop, J. A. & Lamsdell, J. C.: Nomenclatural notes on Carcinosomatidae

Perty, M. 1833. Delectus animalium articulorum, quae in itinere per Størmer, L. 1974. Arthropods from the Lower Devonian (Lower Em- Brasilia anni 1817–1820 peracta collegerunt J. B. Spix et de Mar- sian) of Alken an der Mosel, Germany. – Senckenbergiana tius. Monachii, 205 pp. lethaea 54: 359–451. Richter, R. 1929. Nomenklatur und Nomenklatur. – Senckenbergi- Strand, E. 1942. Miscellanea nomenclatorica zoological et palaeonto- ana 11: 380–386. logica. – Folia Zoologica et Hydrobiologica 11: 386–402. Ritchie, A. 1968. Lanarkopterus dolichoschelus (Størmer) gen. nov., a Tetlie, O. E. 2007. Distribution and dispersal history of Eurypterida mixopterid eurypterid from the Upper Silurian of the Lesmaha- (Chelicerata). – Palaeogeography, Palaeoclimatology, Palaeoecol- gow and Hagshaw Hills inliers, Scotland. – Scottish Journal of ogy 252: 557–574. Geology 4: 317–338. Tetlie, O. E., Brandt, D. S. & Briggs, D. E. G. 2008. Ecdysis in sea Ruedemann, R. 1919. Preservation of alimentary canal in an eurypter- scorpions (Chelicerata: Eurypterida). – Palaeogeography, Palaeo- id: Paleontologic contributions from the New York State Museum. climatology, Palaeoecology 265: 182–194. – New York State Museum Bulletin 227–228: 92–95. Tollerton Jr., V.P. 1989. Morphology, taxonomy, and classification of Størmer, L. 1934. Merostomata from the Downtonian Sandstone of the Order Eurypterida Burmeister, 1843. – Journal of Paleontol- Ringerike, Norway. – Skrifter utgitt av Det Norske Videnskaps- ogy 63: 642–657. Akademi i Oslo I. Matematisk-Naturvidenskapelig Klasse 10: 1– Williams, M. Y. 1915. An eurypterid horizon in the Niagara Forma- 125. tion of Ontario. – Geological Survey of Canada Museum Bulle- Størmer, L. 1955. Merostomata. In Moore, R. C. (ed.). Treatise on In- tin 20: 1–21. vertebrate Paleontology Part p. Arthropoda 2. Geological Society Woodward, H. 1913. III. The position of the Merostomata. – Geologi- of Amerrica and University of Kansas Press, Lawrence: pp. 4–41. cal Magazine 10: 293–300.

museum-zoosyst.evol.wiley-vch.de # 2012 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim