TRANSACTIONS OF THE KANSAS ACADEMY OF SCIENCE 103(1-2), 2000, pp. 76-94 Horseshoe Crabs (Arthropoda: Xiphosurida) from the Pennsylvanian of Kansas and Elsewhere

LOREN E. BABCOCK Department of Geological Sciences, The Ohio State University, Columbus, Ohio 43210

DANIEL F. MERRIAM Kansas Geological Survey, The University of Kansas, Lawrence, Kansas 66047

Pennsylvanian strata of Kansas have yielded two species of xiphosurids (horseshoe crabs). The exceptionally preserved fossils are from inferred coastal (estuarine or tidal flat) paleoenvironments. One species, the euproop- id Euproops danae (Meek and Worthen), occurs in the Lawrence Formation (Upper Pennsylvanian: Virgilian) in Douglas County, Kansas. Siderite con­ cretions in the Lawrence Formation preserve organisms similar to those from Braidwood-type assemblages of the Mazon Creek biota of Illinois. This oc­ currence of E. danae in the Virgilian Stage (North American usage) is one of the few stratigraphically above the Desmoinesian Stage of North America or the Westphalian D Series of Europe. Another xiphosurid, the limuline Paleolimulus signatus (Beecher), occurs in the Pony Creek Shale Member of the Wood Siding Formation (Upper Pennsylvanian: Virgilian) in Wabaunsee County, Kansas. This occurrence extends the known stratigraphic range of P. signatus downward from the Permian into the Pennsylvanian. Another specimen that is here referred to P. signatus, and which was collected from the Mazon Creek biota of Illinois, shows that the species ranges downward to the Desmoinesian Stage (of North American usage).

INTRODUCTION Xiphosurids or horseshoe crabs are an important, although generally un­ common, component of Pennsylvanian and Permian strata of Kansas. As a measure of their importance among xiphosurids generally, approximately 45 genera of Phanerozoic horseshoe crabs have been described (not all of which are valid), of which six genera were described using specimens from the Lower Permian of Kansas. This figure is mitigated, however, by the fact that five genera of putative xiphosurids that were described from the insect beds in the Lower Permian Wellington Formation of Elmo or Annelly, Kansas VOLUME 103, NUMBERS 1-2 77

(Tasch, 1961, 1963, 1964), have been reassigned subsequently to other ar­ thropod groups (Bergstrom, 1975). Another three genera have been reported from Pennsylvanian or Permian rocks of the state (Dunbar, 1923; Tasch, 1961, 1963, 1964; Bennett, 1984; Babcock, Merriam, and West, in press) after having been described using specimens collected elsewhere. It is note­ worthy that the genus Paleolimulus, which was described first from the Permian of Kansas (Dunbar, 1923), is a valid xiphosurid and represents one of the oldest known limulines, or so-called modern horseshoe crabs (Selden and Siveter, 1987; Anderson and Selden, 1997). The purpose of this paper is to document the Pennsylvanian horseshoe crabs that occur in Kansas. This work provides further information on the biogeographic and paleoenvironmental distribution of the taxa now known from the state, provides new information on their temporal distribution, and provides data relevant for a deeper understanding of the exceptional pres­ ervation of nonmineralized arthropod cuticle. Two xiphosurid species pres­ ently are known from the Pennsylvanian of Kansas: Euproops danae (Meek and Worthen) and Paleolimulus signatus (Beecher). Paleolimulus signatus, which also occurs in the Permian, is described more fully, primarily from Permian remains, by Babcock, Merriam, and West (in press). Previously, this species was documented from Kansas alone, but evidence presented here shows that the species distribution extended to Illinois. Euproops danae has not been documented from Kansas or adjoining states until now, although it has a wide distribution in Pennsylvanian strata of North America and Upper Carboniferous strata of Europe. This work shows that both E. danae and P. signatus range upward from the Desmoinesian Stage of North Amer­ ica; E danae extends into the Virgilian, and P. signatus extends into the Lower Permian.

REPOSITORIES Specimens cited here are deposited in the following collections: Orton Geological Museum, The Ohio State University, Columbus, Ohio (OSU); Peabody Museum of Natural History, Yale University, New Haven, Con­ necticut (YPM); University of Kansas Natural History Museum (Division of Invertebrate Paleontology), Lawrence, Kansas (KUMIP); and U.S. Na­ tional Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM).

LOCALITY AND STRATIGRAPHIC INFORMATION Pennsylvanian horseshoe crabs are reported here from two localities in Kansas (Fig. 1). Locality 1 has yielded only one specimen that we have examined, whereas locality 2 has yielded tens of specimens. A generalized stratigraphic section, showing locality intervals, is given in Figure 2. Ad­ ditional stratigraphic information concerning the units in which specimens 78 TRANSACTIONS OF THE KANSAS ACADEMY OF SCIENCE

102°W 40°N"

KANSAS Locality 2 V Locality 1

100 km

Figure 1. Map of Kansas showing localities yielding Pennsylvanian xiphosurans. Locality 1, near Lawrence, Douglas County (Lawrence Formation Konservat-Lagerstatte). Locality 2. near Maple Hill, Wabaunsee County (Pony Creek Shale Konservat-Lagerstatte). occur was provided by Jewett, O'Connor, and Zeller (1968). Productive beds for nonmineralized fossils such as those described in this paper are often referred to by the German term "Konservat-Lagerstatten" (following Sei­ lacher, 1970), which translates into English as "fossil conservation deposits" (Shields, 1998). Locality 1. Lawrence Formation Konservat-Lagerstatte. Lawrence Forma­ tion (Douglas Group; Upper Pennsylvanian: Virgilian), near Lawrence, Douglas County, Kansas. We have examined a single specimen of Euproops danae (YPM 35153) from the Lawrence Formation Lagerstatte. According to a specimen label written by W.H. Twenhofel, who probably acquired the specimen for the Yale Peabody Museum, the illustrated specimen (Fig. 3) is from "Train[?] mounds," "Lawrence? Shales," "near Lawrence, Kansas." The specimen is preserved as part and counterpart in a siderite concretion and, on the back of the counterpart, someone (presumably the collector) carved the words "Oct. 12th 1884 I dug this f[rom] the coal." Based on our experience with upper Paleozoic strata of eastern Kansas, the specimen probably was col­ lected from siderite concretions that occur in the shale immediately above the Williamsburg coal bed (see Moore and others, 1951; Jewett, O'Connor, and Zeller, 1968; Feldman and others, 1993; Fig. 4), near the top of the Lawrence Formation. The available locality data are insufficient to judge precisely where the specimen was collected although today there are only two reasonably good exposures of the Williamsburg coal bed in the vicinity of Lawrence. At both VOLUME 103. NUMBERS 1-2 79

UJ Q_ a: D JANESVILLE SHALE I o FALLS CITY LIMESTONE 5 DC ONAGA SHALE BROWNVILLE LS. < o MEMBER WOOD SIDING PONY CREEK FORMATION Locality 2 SHALE MEMBER D. ROOT SHALE \ => GRAYHORSE LS. O STOTLER LIMESTONE MEMBER or PILLSBURY SHALE PLUMB SHALE o MEMBER UJ ZEANDALE LIMESTONE LU WILLARD SHALE NEBRASKA CITY CO EMPORIA LIMESTONE LS. MEMBER z AUBURN SHALE < BERN LIMESTONE z C < _J r- SEVERY SHALE >- 0) CO z TOPEKA LIMESTONE z < z _l D_ UJ CALHOUN SHALE oZ) a. o rr DEER CREEK CL O LIMESTONE uj > UJ TECUMSEH SHALE Q. UJ D. 3 z LECOMPTON < LIMESTONE I KANWAKA SHALE CO

OREAD LIMESTONE -Locality 1

AMAZONIA LS. MEMBER 00 LAWRENCE < 0- FORMATION IRELAND SS. _J Z) MEMBER o o ROBBINS SHALE 3 rr MEMBER O es STRANGER HASKELL LS. o FORMATION MEMBER

Figure 2. Generalized stratigraphic relationships of Upper Pennsylvanian (Virgilian) strata in Kansas and horizons yielding xiphosurans. Approximate positions ol" fossil conservation deposits discussed in text, keyed to Figure I, arc indicated. Upper part of Lawrence Formation, which contains locality I, is not divided into members (following the usage of Zeller. I968). Abbreviations: Ls., Limestone: Ss., Sandstone. 80 TRANSACTIONS OF THE KANSAS ACADEMY OF SCIENCE

1CT5«Kx

.t, fmmk

Figure 3. Euproops danae (Meek and Worthen, 1865), large specimen preserved in siderite concretion from the Lawrence Formation (Upper Pennsylvanian), near Lawrence, Kansas (lo­ cality 1 in Figs. 1, 2); YPM 35153, X 1. A, Counterpart, dorsal view, showing proximal part of telson. B, Part of same specimen, dorsal view, showing strong wrinkling and slight forward inclination of prosoma.

siderite concretions Williamsburg coal bed

Figure 4. Generalized stratigraphic section of the Lawrence Formation and parts of adjacent formations in Pennsylvanian Douglas Group of northeastern Kansas (modified from Zeller, 1968). Positions of Williamsburg coal bed and overlying siderite concretion-bearing shales of Lawrence Formation are indicated to right of stratigraphic column. Lithologic symbols are: dots, sandstone; dashed horizontal lines, shale: continuous horizontal lines, black shale; blocks, limestone; solid black, coal; cross hatches, red color. Thickness of Lawrence Formation in northeastern Kansas ranges upward to about 45 meters. VOLUME 103, NUMBERS 1-2

QUADRANGLE LOCATION

Contour interval 50 feet. Figure 5. Map of localities in Douglas County, Kansas, showing exposures of Lawrence Formation from which specimen of Euproops danae (Fig. 3) may have been collected. Base map is from Lawrence, Kansas, 30' quadrangle topographic map (U.S. Geological Survey, 1889). Features such as Union Pacific Railroad tracks and Clinton Station no longer exist. Superimposed on map for reference to present-day geographic features are locations of dams along stream channels and lakes (Clinton Lake and Lone Star Lake) that they have produced. Position number I is in bluffs of Wakarusa River Valley, near Union Pacific Railroad tracks; this area is covered now by Clinton Lake. Position number 2 is in vicinity of present-day dam at Lone Star Lake. Contour interval of map (50 feet) equals 15.2 meters. localities, fossil-bearing siderite concretions are abundant in the shales im­ mediately above the coal. Judging from a contemporary map, surveyed in 1885 (Lawrence, Kansas 30' topographic quadrangle; U.S. Geological Sur­ vey, 1889; Fig. 5), the same exposures were as well exposed, or better exposed, in 1884, when the specimen was collected. The most likely locality (Fig. 5, number 1) is near the confluence of the Wakarusa River and Rock Creek. Today, most exposures of the Lawrence Formation in that general 82 TRANSACTIONS OF THE KANSAS ACADEMY OF SCIENCE vicinity are in sees. 8 and 17, T. 13 S., R. 19 E., Lawrence West, Kansas, 7.5' topographic quadrangle (U.S. Geological Survey, 1950, photorevised 1967, 1978), approximately 7 km southwest of the campus of The University of Kansas. According to the 1889 map (Fig. 5), tracks of the Union Pacific Railroad passed just north of the Wakarusa River along a west-southwest line from southern Lawrence to Clinton Station, Kansas. A likely collecting spot would have been in the bluffs of the Wakarusa River Valley, in an area now covered by Clinton Lake. The railroad tracks and Clinton Station are no longer in existence. The second, and less likely, locality (Fig. 5, number 2) is along the banks of Washington Creek, in the vicinity of what is now Lone Star Lake (see Feldman and others, 1993). Today, exposures of the Lawrence Formation at this locality are in sees. 11 and 14, T. 14 S., R. 18 E., Globe, Kansas 7.5' topographic quadrangle (U.S. Geological Survey, 1955), approximately 17 km southwest of The University of Kansas. Locality 2. Pony Creek Shale Konservat-Lagerstatte. Pony Creek Shale Member of the Wood Siding Formation (Wabaunsee Group; Upper Penn­ sylvanian: Virgilian); 3.2 km southwest and 3.2 km south of Maple Hill, Wabaunsee County; along the north section line of sec. 35, T. 11 S., R. 12 E., Maple Hill, Kansas 7.5' topographic quadrangle (U.S. Geological Sur­ vey, 1953, photorevised 1978). More than 20 specimens of Paleolimulus signatus have been collected from this locality. Most are moldically preserved on the outsides of gray to tan limestone concretions ranging in size from two to eight centimeters in maximum dimension. Some specimens occur within or on the outsides of thin, fine-grained, lenticular limestone beds. Specimens usually are articu­ lated, and many retain appendages or book gills.

DEPOSITIONAL SETTINGS AND RELATIONSHIP TO EXCEPTIONAL PRESERVATION Relatively few arthropod clades are characterized by having cuticles re­ inforced by minerals such as calcite. The principal groups that have min­ eralized exoskeletons are trilobites, ostracodes, and some crustaceans. Other forms, which lacked mineralized exoskeletons, tend to have a much poorer fossil record, and instances of their preservation can be regarded as excep­ tional. Xiphosurids, and the related synxiphosurines (see Anderson and Sel­ den, 1997), are examples of arthropods lacking mineralized exoskeletons. Despite the unusual occurrence of exceptional preservation (see Allison and Briggs, 1991 for summary information), such preservation can occur in some predictable circumstances (see Kluessendorf, 1994; Liddell, Wright, and Brett, 1997), usually at the interface of strongly contrasting oxygen, salinity, or other conditions in the environment (Babcock, 1998). Pennsylvanian xiphosurids from Kansas have been preserved in inferred coastal settings where a variety of factors may have contributed to the ex­ ceptional preservation of nonmineralized organisms. In coastal settings, sed- VOLUME 103, NUMBERS 1-2 83

imentation rates, related to tidal deposition (Feldman and others, 1992, 1993) may have been relatively high, and this would have aided in burial of or- ganismic remains. Additionally, taphonomic experiments have demonstrated that the disarticulation of xiphosurid remains is relatively slow (Babcock and Chang, 1997; Babcock, Merriam, and West, in press); this would provide ample time for burial of remains, assuming that the remains were not af­ fected severely by scavengers. Once buried, remains could be expected to remain intact unless sediment bioturbators disturbed them. Conditions that may have limited both scavengers and sediment bioturbators in Pennsylva­ nian coastal settings would have included fluctuating salinity (Babcock, 1998; Babcock, Merriam, and West, in press), and perhaps desiccation. Euproops danae from the Lawrence Formation occurs in a layer of siderite concretions just above the Williamsburg coal. The concretions contain, in addition to the xiphosurid, such plants as Pecopteris, Neuropteris, Cordaites, and Annularia; such insects as cockroaches and dragonflies; bivalves; and coprolites (see Feldman and others, 1993). This biota is similar to the Braid- wood biota present in siderite concretions of the Mazon Creek Konservat- Lagerstatte (Francis Creek Shale Member of the Carbondale Formation) in northeastern Illinois (see Nitecki, 1979; Baird and others, 1985a, 1985b, 1986; Baird, 1997a; Shabica and Hay, 1997). The Lawrence Formation biota is less diverse than the Braidwood biota. The lower diversity of the Lawrence Formation reflects in part a considerably lesser degree of exposure and collecting. The Lawrence Formation in Douglas County, Kansas, is interpreted to have been deposited in an estuarine to coal-forming swamp and floodplain setting along a coastal delta plain (see Baird and Maples, 1997). Deposition was tidally influenced evidently (Feldman and others, 1993; Joeckel, 1994), as indicated by rhythmic, light-dark couplets of mm-scale sediment. The depositional interpretation is based on outcrop study of stratigraphic and sedimentational patterns (compare Baird, 1997c), and on the strong similar­ ity among its preserved biota with the Braidwood biota of Illinois (see Pfef- ferkorn, 1979; Shabica, 1979; Schram, 1979; Baird, 1979; Baird and others, 1985a, 1985b, 1986). The formation of siderite concretions was probably early diagenetic, just as in Mazon Creek-type assemblages of Illinois (Wood­ land and Stenstrom, 1979; Schopf, 1979; Baird and others, 1985a, 1986; Baird, 1997b). Conditions for siderite precipitation are restricted (Woodland and Stenstrom, 1979; Berner, 1981; Baird, 1997b), and imply a deltaic en­ vironment where there was a rapid transition from freshwater to marine water conditions. Assuming that pore waters in recently deposited sediment contained iron and fresh to brackish water, bacterial decomposition could have sufficiently increased the pH in sediment in the vicinity of organic debris to produce chemical microenvironments that mediated the precipita­ tion of iron carbonate (see Woodland and Stenstrom, 1979; Baird and others, 84 TRANSACTIONS OF THE KANSAS ACADEMY OF SCIENCE

1985a, 1986; Baird, 1997b). The Williamsburg coal bed likely represents a coastal peat swamp analogous to the Colchester Coal Member of the Car- bondale Formation (Baird and others, 1985a; Baird, 1997c), which imme­ diately underlies the Francis Creek Shale Member in Illinois. Paleolimulus signatus from the Pony Creek Shale Member of the Wood Siding Formation (locality 2) occurs in a succession of rocks representing a marine-to-nonmarine transition (West and Matsumoto, 1989). Rhythmic, mm-scale, light-dark laminations at the site, together with a low diversity of body fossils, and a trace fossil assemblage that includes trails attributed to horseshoe crabs (Babcock, Merriam, and West, in press), suggest that exceptional preservation of Paleolimulus remains in this deposit occurred mostly in a tidal flat to estuarine setting. Other taxa from the site (principally bivalves) do not show preservation of nonmineralized body parts. Preser­ vation of nonmineralized cuticle is inferred to have been related at least partly to fluctuating salinity (Babcock, 1998), which would have had the secondary effect of limiting scavenging and sediment reworking by ma- croorganisms, in a marginal-marine environment. Desiccation on a tidal flat is to be expected, and it too may have contributed to the exceptional pres­ ervation of xiphosurids in the Pony Creek Shale Member.

SYSTEMATIC PALEONTOLOGY Order Xiphosurida Latreille, 1802 Suborder Bellinurina Eastman, 1913 Family Euproopidae Eller, 1938 Genus Euproops Meek, 1867b Type species. Bellinurus danae (Meek and Worthen, 1865), by inferred original designation (Meek, 1867b, p. 394, 395). Although the binomen B. danae was not mentioned by Meek (1867b) at the time that he proposed the name Euproops, it is clear from Meeks's (1867b, p. 395) reference to ". . . our Illinois fossil . . ." and ". . . remarks accompanying the description of the Illinois species as being probably of more than specific importance ..." that Euproops was erected to embrace B. danae. Remarks. Anderson (1994) provided an emended diagnosis of the genus and discussed synonyms. Later, Anderson and Selden (1997) discussed syn- apomorphies of Euproops and the phylogenetic relationships of xiphosurid taxa including ones assigned to this genus. Euproops danae (Meek and Worthen, 1865) Figure 3

Bellinurus danae Meek and Worthen, 1865, p. 44; Meek and Worthen, 1866, p. 395-398, pi. 32, figs. 2, 2a. Prestwichia danae (Meek and Worthen). Meek, 1867a, p. 257, 258; Packard, VOLUME 103, NUMBERS 1-2 85

1886, p. 146, 147, pi. 5, figs. 3, 3a, pi. 6, figs. 1, la, 2, 2a; Weller, 1898, p. 479; Eastman, 1913, fig. 1499. Euproops danae (Meek and Worthen). Meek, 1867c, p. 320, 321; Meek and Worthen, 1868, p. 547-549, text fig. A; Woodward, 1868, p. 2; White, 1884, p. 170-172; pi. 39, fig. 1; Packard, 1885, p. 291; Packard, 1886, fig. 9; Lesley, 1889, p. 234, fig.; Kobayashi, 1933, p. 181, pi. 16, fig. 3; Raymond, 1944, p. 484-486; Raymond, 1945, p. 4-7, pi. 1, figs. 1, 2, pi. 2, figs. 1, 2; Shimer and Shrock, 1944, p. 707, pi. 300, fig. 5; St0rmer, 1955, p. P20, fig. 15B; Murphy, 1970, p. 281, 282, pi. 1, fig. 1; Fisher, 1977, p. 175-194, figs. 1-8; Fisher, 1979, p. 379-440, figs. 1, 2, 4C-F, 5-7; Fisher, 1981, p. 51-58, figs. 2B, 4A, 4B; Fisher, 1984, fig. 2; Baird, Shabica, Anderson, and Richardson, 1985a, fig. 5.7; Selden and Siveter, 1987, p. 388, fig. 1H; Eldredge, 1991, pi. 89; Anderson, 1994, p. 270- 273, figs. 1-3; Baird, 1997a, p. 21, fig. 4A.3; Mikulic, 1997, p. 135-137, figs. 10.1, 10.2. Euproops colletti White, 1884, p. 172, pi. 39, fig. 2; Lesley, 1889, p. 233, fig- Euproops longispina Packard, 1885, p. 292; Raymond, 1944, p. 491, 492. Prestwichia longispina (Packard). Packard, 1886, p. 147, 148, pi. 5, fig. 4, pi. 6, fig. 3. Belinurus [sic] lacoei Packard, 1885, p. 292; Packard, 1886, p. 149, pi. 5, fig. 5. Prestwichia colletti (White). Weller, 1898, p. 479. Prestwichia (Euproops) scheeleana Ebert, 1892, p. 215—220, figs. 1-3; Dix and Pringle, 1929, p. Ill; Dix and Pringle, 1930, table 1. Euproops amiae Woodward, 1918, p. 465-467, figs. 2-4; Raymond, 1944, p. 493; Copeland, 1957, p. 49, pi. 17, figs. 1-8. Prestwichianella nitida Dix and Pringle, 1929, p. 101, 102, fig. 9; Dix and Pringle, 1930, table 1. Euproops graigolae Dix and Pringle, 1929, p. 104, 105, 114, figs. 10, 16; Dix and Pringle, 1930, table 1. Euproops gwenti Dix and Pringle, 1929, p. 105—107, figs. 11, 16; Dix and Pringle, 1930, table 1. Euproops islwyni Dix and Pringle, 1929, p. 107, 108, figs. 12, 16; Dix and Pringle, 1930, table 1. Euproops meeki Dix and Pringle, 1929, p. 108, 109, figs. 13, 16; Dix and Pringle, 1930, table 1. ? Euproops cambrensis Dix and Pringle, 1929, p. 110, fig. 14; Dix and Pringle, 1930, table 1. Euproops packardi Willard and Jones, 1935, p. 127—131, figs. 1, 2; Ray­ mond, 1944, p. 493. Euproops darrahi Raymond, 1944, p. 489, 490, pi. 2, fig. 4. 86 TRANSACTIONS OF THE KANSAS ACADEMY OF SCIENCE

Euproops laevicula Raymond, 1944, p. 490, fig. 3; Raymond, 1945, p. 7; Murphy, 1970, p. 282-284, pi. 1, fig. 2. Euproops laticephalus Raymond, 1944, p. 491, fig. 4. Euproops thompsoni Raymond, 1944, p. 486-489, fig. 1; Raymond, 1945, p. 7; St0rmer, 1955, p. P20, figs. 13,4a, 13,4b. Euproops spec. ind. Raymond, 1944, p. 492. Euproops lacoei (Packard). Raymond, 1944, p. 494, 495. Euproops cf. danae (Meek and Worthen). Copeland, 1957, pi. 17, fig. 9, pi. 21, fig. 1; Fisher, 1979, p. 431. Euproops kilmersdonensis Ambrose and Romano, 1972, p. 570-576, text- figs. 1, 2, pi. 112, figs. 1-3; pi. 113, fig. 1, table 1; Brauckmann, 1982; p. 22; Jarzembowski, 1989, p. 223; Pollard and Hardy, 1991, p. 170, 176. Euproops [sp.]. Jarzembowski, 1989, fig. 5.

Material. One specimen, YPM 35153, preserved as part and counterpart, from locality 1. Remarks. This species has been well described and illustrated from re­ mains collected in Illinois (e.g., Meek and Worthen, 1865; Packard, 1886; Raymond, 1944; Fisher, 1977, 1979), Indiana (e.g., White, 1884; Raymond, 1944), Pennsylvania (Raymond, 1944), Ohio (Murphy, 1970), Nova Scotia (Woodward, 1918; Raymond, 1944; Copeland, 1957), Great Britain (e.g., Dix and Pringle, 1929, 1930; Ambrose and Romano, 1972; Anderson, 1994), and Germany (Ebert, 1892; Brauckmann, 1982). The specimen from north­ eastern Kansas (Fig. 3) agrees in all respects with Euproops danae except that the opisthosoma in the illustrated specimen (Fig. 3) is wider than in many illustrated specimens. This minor difference, however, is interpreted to be taphonomic in origin and attributed to the flattening of a pliable exo­ skeleton sometime after death or molting (compare Babcock and Chang, 1997). As discussed by Anderson (1994), considerable variability in the appearance of the exoskeleton within this species has resulted from tapho­ nomic distortion of the relatively pliable exoskeleton. Most of the species listed here as synonyms can be attributed to original misinterpretation of the effects of such postmortem distortion (see Anderson, 1994). Eup>roop>s danae is no longer referred to Bellinurus, as it was originally (Meek and Worthen, 1865). Currently, the rationale for retaining E. danae in a separate genus (of which it is the type species) is that it possesses such distinguishing characters as ophthalmic ridges that have a concave course to the cardiac lobe, and opisthosomal flanges that are composed of the fused bases of the fixed lateral spines (Anderson and Selden, 1997). Previously, E. danae was not documented from the Pennsylvania of Kan­ sas. A rather stylized reconstruction of Prestwichianella (= Euproops), how­ ever, appeared in a general book on Kansas geology (Bennett, 1984, fig. 72). The material basis for this citation is not known. VOLUME 103, NUMBERS 1-2 87

Stratigraphically, E. danae is confined to the Pennsylvanian (or Upper Carboniferous). Previous reports of this species were mostly in rocks as­ signed to the Desmoinesian Stage of North America or equivalents (West­ phalian D) of Europe (see Raymond, 1944; Anderson, 1994). By interna­ tional standards, these occurrences are in the Myachkovskian Stage of the Moscovian Series. A few specimens, however, have been reported (Ray­ mond, 1944) from the Missourian and Virgilian Stages (North American usage). The specimen from northeastern Kansas discussed here seems to have been collected from a stratigraphic position at or close to the upper limit of the species. It is from the middle part of the Virgilian Stage (North American usage), which is equivalent to the Klazminskian Stage of the Gze- lian Series (international usage). Suborder Limulina Richter and Richter, 1929 Superfamily Paleolimuloidea Anderson and Selden, 1997 Family Paleolimulidae Raymond, 1944 Genus Paleolimulus Dunbar 1923 Type species. Paleolimulus avitus Dunbar, 1923, by original designation (Dunbar, 1923, p. 444). Babcock, Merriam, and West (in press) proposed suppression of P. avitus as a junior subjective synonym of Prestwichia sig- nata Beecher, 1904. Remarks. Prestwichia, to which P. signata was referred originally (Bee­ cher, 1904), is no longer considered to be a valid genus, as it was synony­ mized previously with Euproops (see Anderson, 1994 for discussion). Bee- cher's (1904) species P. signata is here referred to Paleolimulus Dunbar, 1923, which is characterized by, among other things, longitudinal opistho­ somal ridges and a pyramidal cheek node (Anderson and Selden, 1997). A change in gender of the trivial name (signata, feminine, to signatus, mas­ culine) has accompanied transfer of the species from Prestwichia (feminine) to Paleolimulus (masculine). Included species. At present, only one species, Prestwichia signata Bee­ cher, 1904 (= Paleolimulus signatus) is included in Paleolimulus. Paleoli­ mulus signatus incorporates as a junior synonym P. avitus Dunbar, 1923. Most other named species previously assigned to Paleolimulus are referable to other genera. For example, Paleolimulus? randalli (Beecher, 1902) from the Devonian of Pennsylvania and New York has been reassigned to Kasi­ belinurus (Babcock and others, 1995); P. longispinus Schram, 1979, from the Mississippian of Montana, probably should be assigned to Rolfeia (Wa- terson, 1985; Anderson and Selden, 1997); and P. fuchsbergensis Hauschke and Wilde, 1987, from the Triassic of Germany, is not a paleolimulid (An­ derson and Selden, 1997), although its generic assignment at present is un­ certain. The correct generic assignment for P. juresanensis Chernyshev, 1933, from the Upper Carboniferous of Russia, is also uncertain at present. 88 TRANSACTIONS OF THE KANSAS ACADEMY OF SCIENCE

Figure 6. Paleolimulus signatus (Beecher. 1904); specimens viewed from dorsal side, pre­ served in small limestone concretions from Pony Creek Shale Member of Wood Siding For­ mation (Upper Pennsylvanian). near Maple Hill. Kansas (locality 2 in Figs. I. 2). A. Prosoma showing strong wrinkles; USNM 484410: X I. B. Internal mold of specimen retaining most of appendages but lacking all except proximal part of telson: USNM 465528; X1.25. C. Dorsal exoskeleton lacking telson: USNM 484408; XI. D. Internal mold of specimen retaining ap­ pendages and book gills; posterior opisthosoma and telson evidently extended beyond limits of concretion and have been lost: USNM 484407: XI.25.

Paleolimulus signatus (Beecher, 1904) Figure 6

Prestwichia signata Beecher, 1904, p. 24, fig. 1. Paleolimulus avitus Dunbar, 1923, p. 444-451, figs. 1-6; Raymond, 1944, VOLUME 103, NUMBERS 1-2 89

p. 505-507; Shimer and Shrock, 1944, p. 707, pi. 300, figs. 1, 2; St0rmer, 1955, p. P21, P22, fig. 15C, 16,1; Fisher, 1981, p. 51-58, figs. 3B, 4A- 4C; Selden and Siveter, 1987, p. 387, fig. 1A; Novozhilov, 1991, fig. 1183. Paleolimulus signatus (Beecher). Dunbar, 1923, p. 450, fig. 8; Babcock, 1996, fig. 1A; Babcock, Merriam, and West, in press, figs. 3, 4A-H. Paleolimulus [sic] avitus Dunbar. Kobayashi, 1933, p. 181; Raymond, 1944, pi. 1, pi. 2, figs. 1-3; St0rmer, 1952, fig. lj. Paleolimulus avitas [sic] Dunbar. Tasch, 1963, p. 1247, pi. 173, fig. 1. Paleolimulus [sp.]. Robison, 1987, fig. 13.46.A; Eldredge, 1991, pi. 90. Paleolimulus [sic] sp. Mikulic, 1997, p. 138, figs. 10.6, 10.7.

Types. Holotype of Prestwichia signata Beecher, 1904, broken prosoma, USNM 48305, from the Fort Riley Limestone Member of the Barneston Limestone (Chase Group; Lower Permian: Wolfcampian Stage), about 4.8 km west of Stockdale, Riley County, Kansas; plastoholotypes, YPM 26319, 26319A. Syntypes of the junior subjective synonym Paleolimulus avitus Dunbar, 1923 are YPM 26324, 26317, 26325, from the Insect Hill deposit, Elmo Limestone Member of the Wellington Formation (Summer Group, Lower Permian: Wolfcampian Stage), Insect Hill, about 4 km south-south­ east of Elmo, Kansas. Material. More than 20 specimens from locality 2, including OSU 46358-46363; YPM 170019-170024; KUMIP 286276-286281; and USNM 465528, 484407-484411. Remarks. Babcock, Merriam, and West (in press) discussed reasons for considering Paleolimulus avitus to be a junior subjective synonym of P. signatus, and discussed the taphonomy of the species. The morphology of this species is known in detail from exceptionally preserved specimens re­ corded from the Permian Elmo Limestone Member of the Wellington For­ mation (Dunbar, 1923; Raymond, 1944; Babcock, 1996; Babcock, Merriam, and West, in press) and the Pennsylvanian Pony Creek Shale Shale Member of the Wood Siding Formation (Babcock, Merriam, and West, in press; here­ in, Fig. 6). Some specimens from these occurrences retain appendages and book gills. Previously, the known stratigraphic range of P. signatus was restricted to the Wolfcampian and Leonardian Stages (North American usage) of the Permian (Beecher, 1904; Dunbar, 1923; Raymond, 1944). By interna­ tional standards, that range extends from the Asselian Stage to the Artinskian Stage of the Lower Permian Series. New material from the Pony Creek Shale Member of the Wood Siding Formation extends the known stratigraphic range of this species downward into the upper part of the Virgilian Stage (North American usage) of the Upper Pennsylvanian, which is equivalent to the Ko- ginskian Stage of the Gzelian Series (international usage). The range is ex­ tended into the Desmoinesian Stage (North American usage), which is equiv­ alent to the Myachkovskian Stage of the Moscovian Series (international us- 90 TRANSACTIONS OF THE KANSAS ACADEMY OF SCIENCE age) by inclusion of a specimen from the Mazon Creek biota of northeastern Illinois (Robison, 1987, fig. 13.46.A; Eldredge, 1991, pi. 90; Mikulic, 1997, fig. 10.6) that had not been previously assigned to species.

ACKNOWLEDGMENTS R. D. White (Peabody Museum of Natural History, Yale University) called attention to the specimen of Euproops danae illustrated herein, and arranged for a loan of it. F. A. Abegg (Chevron Company U.S.A.), L. I. Anderson (University of Aberdeen), A. W. Archer (Kansas State University), S. M. Bergstrom (The Ohio State University), W. G. Coyle (Chevron Company U.S.A.), C. R. Easterday (The Ohio State University), K. R. Evans (formerly University of Kansas), H. R. Feldman (Shell Oil Company), R. L. Kaesler (University of Kansas), the late A. Kamb (University of Kansas Natural History Museum, Division of Invertebrate Paleontology), J. Keim (Univer­ sity of Kansas), C. G. Maples (Indiana University), R. A. Robison (Univer­ sity of Kansas), W. L. Watney (Kansas Geological Survey), and R. R. West (Kansas State University) provided field assistance, locality information, or assistance in locating reference materials. This paper has benefitted from the helpful reviews of R. L. Kaesler and B. S. Lieberman (both of the University of Kansas). This work was partly supported by a grant from the Kansas Geological Survey to Babcock.

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-;,-.>;.•:-..:-: .,',< •: .•*.; . ••. > r, ':•>-:' :•• '. ' " "!r •'; .:>, ' .•(•. •' -,\''',••. :- - •-'• •'.' •.• ••*• " ; . .; Tjq-.M' •••-• -in: ,t 1 )f..-i,'.c:

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