Journal of Paleontology, 84(6), 2010, p. 1206–1208 Copyright ’ 2010, The Paleontological Society 0022-3360/10/0084-1206$03.00

AN ISOLATED PTERYGOTID RAMUS (CHELICERATA: EURYPTERIDA) FROM THE DEVONIAN BEARTOOTH BUTTE FORMATION, WYOMING

JAMES C. LAMSDELL1 AND DAVID A. LEGG2 1Paleontological Institute, University of Kansas, 1475 Jayhawk Boulevard, Lawrence, Kansas 66045, USA, ,[email protected].; and 2Department of Earth Science and Engineering, Imperial College, London SW7 2AZ, UK

ABSTRACT—An isolated ramus of the pterygotid eurypterid Jaekelopterus cf. howelli from the Early Devonian (Pragian) Beartooth Butte Formation (Cottonwood Canyon, north-western Wyoming) is described. Pterygotid taxonomy and synonymy is briefly discussed with the genera Pterygotus, Acutiramus and Jaekelopterus shown to be potential synonyms. The use of cheliceral denticulation patterns as a generic-level character is discouraged in light of variations within genera and its unsuitability as a major characteristic in the other eurypterid families.

INTRODUCTION the type section of the Beartooth Butte Formation in TERYGOTIDS ARE a diverse group of predatory Palaeozoic Beartooth Butte, which is Early Devonian in age. For a more P eurypterids, famed for being amongst the largest arthro- comprehensive discussion of the formation, its stratigraphy pods ever to have lived (Braddy et al., 2008a). The and paleoenvironment, see Tetlie (2007b). Vertebrate biostra- Pterygotidae are characterized by the possession of enlarged tigraphy (Elliot and Ilyes, 1996) indicates that the Cotton- raptorial chelicerae, non-spiniferous appendages III–V, undi- wood Canyon section is Pragian whereas the type section at vided medial appendages, laterally expanded pretelson and a Beartooth Butte is Emsian. Stable oxygen and isotope data broad paddle-like telson with marginal ornamentation (Tetlie (Poulson in Fiorillo, 2000) indicate that the Beartooth Butte and Briggs, 2009). They originated in the late early Silurian Formation was deposited in an estuarine environment, with and went extinct during the Middle Devonian (Tetlie, 2007a), the Cottonwood Canyon section being slightly less saline than possibly due to increased competition from vertebrates the type section. (Lamsdell and Braddy, 2010). There are currently 46 The Beartooth Butte Formation is best known for its fish recognized species assigned to 5 genera within Pterygotidae, fauna; however, despite their lower numbers, the associated making them the most speciose clade of eurypterids. Pter- eurypterids are also of importance. The eurypterids of the ygotids are almost certainly taxonomically over-split however Beartooth Butte section were originally described by Ruede- (Braddy et al., 2008b): Erettopterus monroensis (Sarle, 1903) mann (1934, 1935a), Kjellesvig-Waering (1955) and Kjellesvig- and Pterygotus impacatus Kjellesvig-Waering, 1964 were Waering and Størmer (1952). Tetlie (2007b) recently rede- synonymized with Erettopterus osiliensis (Schmidt, 1883) by scribed the fauna, synonymizing some species and reducing the Ciurca and Tetlie (2007) while Pterygotus minor Woodward, eurypterids known from the site to Dorfopterus angusticollis 1864 and Pterygotus atlanticus Clarke and Ruedemann, 1912 Kjellesvig-Waering, 1955, Jaekelopterus howelli (Kjellesvig- have both been suggested to be synonyms of Pterygotus Waering and Størmer, 1952) and Strobilopterus princetonii anglicus Agassiz, 1844 (Braddy, 2000; Miller, 2007). (Ruedemann, 1934). He further suggested that Dorfopterus Herein is described an isolated free ramus of a pterygotid and Strobilopterus may be synonyms. The eurypterid fauna chelicera, FMNH 6177.2, from the Early Devonian Beartooth from the Cottonwood Canyon section is as yet undescribed, Butte Formation of Cottonwood Canyon, north-western although Tetlie (2007b) noted that the most common Wyoming. This is the same specimen as referred to by eurypterid there closely resembles S. princetonii. Three Kjellesvig-Waering (1986) in his posthumous scorpion mono- scorpion specimens are also known from Cottonwood graph as Pterygotus mcgrewi Kjellesvig-Waering and Richard- Canyon, each assigned its own monospecific genus; Hydro- son, apparently an unpublished manuscript name that made it scorpius denisoni Kjellesvig-Waering, 1986, Acanthoscorpio into the literature. mucronatus Kjellesvig-Waering, 1986 and Branchioscorpio richardsoni Kjellesvig-Waering, 1986. However, A. mucrona- MATERIALS AND METHODS tus, preserved on the same slab as the ramus, may in fact be a The material described herein is deposited in the Field juvenile eurypterid (Legg et al., 2009). Museum of Natural History, Chicago (FMNH). Morpholog- ical terminology follows Tollerton (1989) with denticle SYSTEMATIC PALEONTOLOGY terminology as used by Miller (2007). Specimens were Order EURYPTERIDA Burmeister, 1843 examined using a Nikon binocular microscope and photo- Suborder EURYPTERINA Burmeister, 1843 graphed immersed in alcohol using a Nikon D80 camera. Superfamily PTERYGOTOIDEA Clarke and Ruedemann, Camera lucida drawings were prepared using Adobe Illustra- 1912 tor 10. Family PTERYGOTIDAE Clarke and Ruedemann, 1912 GEOLOGICAL SETTING JAEKELOPTERUS Waterston, 1964 JAEKELOPTERUS cf. HOWELLI (Kjellesvig-Waering and The ramus was recovered from the Cottonwood Canyon Størmer, 1952) section of the Beartooth Butte Formation by Robert H. Figure 1 Denison and E. S. Richardson, Jr. in 1962. The Cottonwood Canyon section is situated in the Bighorn Mountains of Pterygotus mcgrewi (nomen nudum)KJELLESVIG-WAERING northern Wyoming (Sandberg, 1961), roughly 100 km east of AND RICHARDSON IN KJELLESVIG-WAERING, 1986, p. 73 1206 LAMSDELL AND LEGG—A PTERYGOTID RAMUS FROM THE DEVONIAN OF WYOMING 1207

reconstructed by Størmer (1936); three large principal denticles with a curved terminal denticle oriented at a right angle to the ramus. The intermediate denticle pattern of two before the anterior principal denticle and two between the anterior and primary principal denticles is identical to that described for Jaekelopterus by Waterston (1964) and seen in the large chelicera of Braddy et al. (2008a). The free ramus of Pterygotus Agassiz, 1839 is also similar to that of Jaekelopterus, to the extent that Braddy et al. (2008b) considered the two genera indistinguishable by the chelicerae alone. The chelicerae of Pterygotus are highly variable; however, the free ramus does differ from Jaekelopterus in possessing seven principal denticles (Miller, 2007) and having the terminal denticle angled slightly away from the ramus (Waterston, 1964). Although a number of pterygotid species have been erected based on isolated chelicera it is felt that, given the high probability of species-level oversplitting, a single chela (or ramus thereof) is not sufficient material to warrant the description of a new species. The presence of Jaekelopterus howelli in the same formation (albeit in slightly younger strata) suggests that the ramus should be assigned to this species, however the temporal difference means such an interpretation must be treated with caution, especially given that the Strobilopterus-like species is very similar to Strobi- lopterus princetonii but not identical (Tetlie, 2007b). The potential for similar eurypterids between the localities to be chronospecies, as is the case in the Eurypterus Dekay, 1825 of the Bertie Group in New York and Ontario (Tetlie et al., 2007), cannot currently be discounted. The chelicerae of the Beartooth Butte section Jaekelopterus are unknown and so no direct comparison between it and the Cottonwood Canyon Jaekelopterus can be made beyond the fact that they are cogeneric. FIGURE 1—Jaekelopterus cf. howelli (Kjellesvig-Waering and Størmer, 1952), photographs and interpretive drawings. Scale bars represent 10 mm. Several lines of evidence point to the Jaekelopterus being 1, Photograph of FMNH PE 6177.2a, comprising the free ramus of the conspecific: they both have a similar projected body length chelicera; 2,photographofcounterpartFMNHPE6177.2b;3, (80 cm for the Beartooth Butte specimens, 100 cm for the interpretive drawing of FMNH PE 6177.2a; 4, interpretive drawing of Cottonwood Canyon specimen [Lamsdell, pers. obs.]), they counterpart FMNH PE 6177.2b. Abbreviations: td9—terminal denticle on free ramus, d19–d39—principal denticles on free ramus, d19 indicates both come from geographically close localities in the same primary denticle, i19–i59—intermediate denticles on free ramus. formation within a relatively short geological period, the two localities have extremely similar faunal components, and there Description.—FMNH PE 6177.2—Fig. 1.1 and 1.3 (part), is no evidence from the Beartooth Butte section of there being Fig. 1.2 and 1.4 (counterpart): Free ramus of chelicera more than one pterygotid species present. Nevertheless, the preserving terminal, three principal and five intermediate authors prefer to assign the specimen described herein to denticles. Total length 91 mm, width at base 18 mm, indicating Jaekelopterus cf. howelli until post-cheliceral specimens are an animal with a total body length of 100 cm (for dimensions found from Cottonwood Canyon or the chelicerae of of individual denticles see Table 1). Terminal denticle oriented Jaekelopterus howelli are known from Beartooth Butte. almost at a right angle to ramus, primary denticle partially PTERYGOTID TAXONOMIC DIVERSITY collapsed into ramus. Anterior principal denticle curves Despite the apparent differences in the chelicerae of posteriorly along anterior edge, while its posterior equivalent Jaekelopterus and Pterygotus, there may be an argument for has broken off along its length. Paired intermediate denticles synonymizing Jaekelopterus, Pterygotus and Acutiramus Rue- are located before and after the anterior principal denticle, demann, 1935b based on similarities in the genital appendage, with a single intermediate denticle at the posterior of the reducing the recognized pterygotid genera to Pterygotus, ramus. Erettopterus Salter (in Huxley and Salter), 1859 and Ciurcop- Discussion.—The specimen bears close comparison with the terus Tetlie and Briggs, 2009. It appears that Waterston (1964) free ramus of Jaekelopterus rhenaniae (Jaekel, 1914) as had similar views on pterygotid taxonomy, considering the morphology of the chelicerae to be too dependent on mode of TABLE 1—Denticle dimensions for FMNH PE 6177.2a+b, free ramus of life and variable through ontogeny to be a suitable basis for a Jaekelopterus cf. howelli. All measurements in millimetres. Column headings correspond to the abbreviations listed in figure 1. Asterisk (*) generic-level taxonomic framework. indicates an incomplete measurement. Furthermore, the criteria applied to pterygotids when organizing their taxonomy differs somewhat to that applied td9 d19 d29 d39 i19 i29 i39 i49 i59 to other eurypterid families. Of the primary characteristics Length/width at base 10/6 7/5 7/4 -/3* 2/2 4/3 2/3 4/4 2/2 defining Jaekelopterus, the marginal cuticular ornamentation Distance from terminal of the telson is not a suitable genus-level character and while denticle — 35 14 53 6 9 25 32 69 the telson shape can be used to distinguish genera, the more 1208 JOURNAL OF PALEONTOLOGY, V. 84, NO. 6, 2010 triangular telson still falls within the morphological range of FIORILLO, A. R. 2000. The ancient environment of the Beartooth Butte the paddle-shaped telson as defined by Tollerton (1989) that is Formation (Devonian) in Wyoming and Montana: Combining paleon- tological inquiry with federal management needs. USDA Forest Service diagnostic of both Acutiramus and Pterygotus. The chelicera Proceedings RMRS-P-15, 3:160–167. of pterygotids can be highly variable, as shown within HALL, J. 1859. 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