Cretaceous Confluence in the Coon

CRETACEOUS CONFLUENCE IN THE COON CREEK FORMATION (MAASTRICHTIAN) OF MISSISSIPPI AND TENNESSEE, USA: TAPHONOMY AND SYSTEMATIC PALEONTOLOGY OF A DECAPOD KONSENTRAT- LAGERSTÄTTE

A thesis submitted to Kent State University

In partial fulfillment of the requirements

For the degree of Masters of Science

Krystyna M. Kornecki

December 2014

Thesis written by

Krystyna M. Kornecki

B.S., St. Lawrence University, 2012

M.S., Kent State University, 2014

Approved by

______Dr. Rodney Feldmann, Advisor

______Dr. Daniel Holm, Chair, Department of Geology

______James Blank, Dean, College of Arts and Sciences

TABLE OF CONTENTS

List of Figures ...... 4

List of Tables ...... 6

Acknowledgments ...... 7

Abstract……………………………….………………………………………………8

Localities……………………………….…………………………………………….10

Institutional abbreviations..…………….…………………………………………….12

Introduction…………………………….……………………………………………13

i. Tectonic setting…………………………………………………………14

ii. Stratigraphy……………………………………………………………..14

Methods……………………………………………………………………………17

Results……………………………………………………………………….18

i. Sedimentology……………………………………………………18

ii. Systematic paleontology……………………………………………20

Discussion……………………………..…………………………………………….97

i. Taphonomy…………………………………………………………97

ii. Paleoecology………………………………………………………..100

iii. Paleobiogeographic comparisons…………………………………103

Conclusions……..…………………………………………………………………106

References…………………………………………………………………………108

LIST OF FIGURES

Figure 1. – Maastrichtian outcrops of North America, black. Intercontinental seaway

outlined, gray (Schuchert, 1955)……………………………………………160

Figure 2. – Late Cretaceous facies along the Mississippi Embayment, with dominantly

siliciclastic sediments in the Selmer – Tupelo area (Mancini et al., 1995)…161

Figure 3. – Regional Upper Cretaceous stratigraphy (C.C. Smith, 1989)…………162

Figure 4. – Interpretation of stratigraphic context of the Coon Creek beds (Mancini et

al., 1995) …………………………………………………………………….163

Figure 5. – Blue Springs Locality of the Coon Creek Formation near New Albany,

Mississippi (Google Earth)………………………..……………………………164

Figure 6. – Thin section images, magnification 10 X 0.4. ………………………….165

Figure 7. – Thin section images, magnification 10 X 20.………………………….166

Figure 8. – Thin section images, magnification 10 X 40.……………………………167

Figure 9. – Illustrations of Hoploparia tennesseensis and Hoploparia georgeana…168

Figure 10. – Illustrations of Mesostylus mortoni, New genus new species, Bournelyridus

n. sp., Prehepatus harrisi, and majid fragments………………………..169

Figure 11. – Illustrations of Linuparus n sp.……………………………………………170

Figure 12. – Illustrations of Tetracarcinus subquadratus and Dakoticancer australis..171

Figure 13. – Ratios of Dakoticancer australis and Tetracarcinus subquadratus

measurements………………………………………….………………………..172

Figure 14. – Number of male and female specimens of Dakoticancer australis………173

Figure 15. – Male vs. female ratios of Dakoticancer australis width to frontal width...174

Figure 16. – Male vs. female ratios of Dakoticancer australis frontal width to fronto-

orbital width…………………………………………………………………….175

Figure 17. – vs. female ratios of Dakoticancer australis width to frontal orbital

width………………………………………………………………………………...……176

Figure 18. – Dakoticancer australis claw height vs. granule size………….………….177

Figure 19. – Tetracarcinus subquadratus measurements of width against ontogenetic ratios………………..……………………………………….…………………………..178

Figure 20. – Illustrations of Cretacoranina testacea, Cristipluma mississippiensis,

Letheticocarcinus atlanticus……………………………………………………179

Figure 21. – Elemental maps of cuticle thin section of Tetracarcinus subquadratus….180

Figure 22. – Elemental maps of cuticle thin section of Linuparus sp..……………..…..181

Figure 23. – Elemental maps of cuticle thin section of Bournelyridus n. sp.……….…182

Figure 24. – Elemental maps of cuticle in thin section of Hoploparia tennesseensis….183

Figure 25. – Elemental maps of cuticle in thin section of Linuparus n. sp……………184

Figure 26. – Thin section of Tetracarcinus subquadratus carapace cuticle…………..185

Figure 27. – Thin section of Dakoticancer australis carapace cuticle…………………186

Figure 28. – SEM images of Dakoticancer australis claw………………….………….187

Figure 29. – Number of species in common with Coon Creek fauna………………..…188

Figure 30. – Distribution of species in common with the Coon Creek fauna in North and

Central America……………………………………………………………..….189

LIST OF TABLES

Table 1. – Percent composition of lithology…………………………...……………….190

Table 2. – Table of Linuparus species characters of the most similar species to Linuparus

n. sp. …………………………………………………………………………191

Table 3. – Dakoticancer australis measurements…………………..…………………..192

Table 4. – Tests of normality…………………………………….……………………..199

Table 5. – Tests of similarity between males and females of Dakoticancer australis…200

Table 6. – Tetracarcinus subquadratus measurements……………..………………….201

Table 7. – Cretoranina testacea measurements………………………………………..202

ACKNOWLEDGMENTS

For providing access to the Bishop Collection of the Blue Springs Locality of the

Coon Creek Formation, I thank Sally Shelton of the South Dakota School of Mines and

Technology Museum and likewise the Memphis Pink Palace Museum, Memphis,

Tennessee, and the Mississippi Museum of Natural Science, Jackson, Mississippi for providing material through loan for study. I am in great gratitude to Richard Keyes for his generosity, help in the field, and loan of an excellent specimen as well as Kendal, Nancy, and other members of the North Mississippi Gem and Mineral Society. Robert Langford also aided in the field and provided innovative field collecting tools. I could not have completed this research without the time and generosity of George Phillips and his correspondence and help in the field. I thank Amanda Millhouse, Daniel Levin, John

Pojeta, and others at the National Museum of History in Washington, D.C. for access to collections for comparison and Merida Keatts for her help using the XRF, SEM, EDX, and all other technical troubleshooting.

I also greatly appreciate Drs. Ovidiu and Adina Franţescu for help and instruction in the lab and helpful discussion as well as Dr. J. Mark Erickson, Dr. Joseph Ortiz, Nidal

Atallah, AnnMarie Jones, Jessica Tashman, Wade Jones, Samantha Yost, Evin Maguire,

Andrew Gerwitz, Kevin Engle, and Ashleigh Stepp for helpful conversation and for sharing their enthusiasm. I also thank Eric Sload for help using the SEM, carbon coating, and gold coating specimens. I sincerely appreciate Dr. Feldmann, Dr. Schweitzer, and Dr.

Wells for their help as my committee members, their support, their patience, and their passion. I also could not have pursued my joy of paleontology without my family and their love and support.

ABSTRACT

The Late Cretaceous (Maastrichtian) Coon Creek Formation of Mississippi and

Tennessee possesses a diverse and abundant assemblage of decapods including lobsters, ghost shrimp, and crabs. The formation lies in a temporally and paleogeographically significant location, situated between the Atlantic Coastal Plain and the Western Interior

Seaway, shortly before the closing of the seaway and the K-Pg mass extinction.

Coon Creek decapods have been little studied since the fauna was first described in the 1920’s. A large collection of specimens, ranging in preservation from poor to excellent, has recently become available for study. Because of the visible variation in preservation, the abundant material, and the paucity of cuticular data of this type of preservation, an investigative study of elemental composition of the sediment and cuticle of six species of decapod from six families (Palinuridae, Nephropidae, Callianassidae,

Dakoticancridae, Raninidae, and Retroplumidae) is conducted using material collected at the Blue Springs Locality in Mississippi.

Cuticle, concretions, decapod burrow, and sediment from the site are analyzed with X-Ray Florescence and Elemental Reflectance for preliminary elemental composition and subsequent mineral composition. Concretions and the burrow were observed in thin section and were mapped for elemental distribution using Energy-

8 dispersive X-Ray spectroscopy and dot mapping. The six species of decapod were analyzed using the same techniques. Taphonomic data supports preservation ranging from well preserved phosphatization to secondary alteration to silica-rich exterior and weathering clay minerals. Because the silica is not replacing the phosphatized exocuticle and microscopic structure of cuticle in cross section is not preserved in the silica layer, the cause of this alteration is uncertain. This preservation is un-like the decapod cuticle preservation of the concretions of the Bearpaw Shale Formation (Late Cretaceous) of

Montana.

Species of the Coon Creek Formation were re-assessed and assigned to modern taxonomic schemes. Sixteen species are identified and two new species are described including Hoploparia tennesseensis, Hoploparia mcnairyensis, Linuparus new species,

Linuparus sp., Palaeopetrochirus enigmus, Seorsus wadei, Bournelyreidus new species,

Cristipluma mississippiensis, Hoploparia georgeana, Mesostylus mortoni, Tetracarcinus subquadratus, Avitelmessus grapsoideus, Cretacoranina testacea, New genus new species (Carcineretidae), Dakoticancer australis, Prehepatus harrisi, and

Latheticocarcinus atlanticus. This decapod assemblage shares common species with correlative units of the Western Interior Seaway, the Gulf Coastal Plain, and the Atlantic

Coastal Plain, supporting the hypothesis that the Mississippi Embayment is an ecotone for North American decapods.

LOCALITIES

“Ft. Pierre Locality,” Pierre Shale Formation.- north section line of sec. 21 to the south section line of sec. 26, T. 6 N., R. 29 E., Stanley County, South Dakota) in the

Upper Cretaceous Pierre Shale, from the zone of Baculites grandis Hall and Meek

(Bishop, 1972, p. 3823).

Pierre Shale Formation (holotype local for Plagiophthalmus izetti) - NE ¼, NE

¼, Sec. 25, T.3 N., R. 17 W., Grand Co., Colorado (USGS Locality D8090); Baculites scotti zone (index fossil to upper Campanian rocks) (Bishop, 1988, p. 377).

Pierre Shale Formation (paratype location for Plagiophthalmus izetti) - SE ¼, SE

¼, Sec. 29, T.2 S., R. 70 W., Jefferson County, Colorado (Bishop, 1988, p. 377).

Prairie Bluff Formation (collection site by Mr. Ralph Harris, 1982 of Prehepatus harrisi) - sec. 3, T8S, R3E, Union County, Mississippi (Bishop, 1985, p. 1030).

Fox Hills Formation (holotype location for Latheticocarcinus shapiroi) - Grand

River, Corson County, South Dakota; two paratypes collected from either Red Bird,

Wyoming, or from the Fox Hills of Central South Dakota; paratypes are either

Campanian or Maastrichtian (lacking any other more specific record) (Bishop, 1988, p.

378).

Bearpaw Shale Formation - specimens present at the U.S. National Museum of

Natural History and the Carter County Museum (Montana) but locality has not be successfully found by Bishop; known to be near Baker, Montana (Bishop, 1976-78, pg.

199).

Coon Creek type section - on the Coon Creek, 1.6 km south of Enville, Coon

Creek Formation, lower lithofacies. Dark greenish gray, glauconitic sandy silt. McNairy

County, TN. Index fossils: Nanno-assemblage 23a Nostoceras hyatti, B. undatus, B. claviformis (Ebersole, 2009, p. 31).

Blue Springs Locality - Lower Maastrichtian Coon Creek Formation (Fig. 2)

(34°24’30.14” N, 88°53’08.36” W). Unvegetated roadcut at the exchange of new U.S.

Highway 78 and Mississippi Highway 9, approximately 22 km northwest of Tupelo,

Mississippi and 2.5 km southwest of the village of Blue Springs (locality GAB 37; E ½,

NW ¼, Sec. 16, T8S, R4E, Union County, Mississippi. 6 meters of sandy, glauconitic and phosphatic light olive gray mudstone. Also includes an interval of phosphatic nodules (Bishop, 1983, p. 417).

Coon Creek Formation (collection site by Mr. Ralph Harris, 1982 of Prehepatus harrisi) - sec. 9, T8S, R4E, Union County, Mississippi (Bishop, 1985, p. 1030).

Coon Creek fauna locality - Known as the “Dave Weeks Place” on Coon Creek.

Located in the Northeastern part of McNairy County, Tennessee, 3 ½ miles south of

Enville, 7 ½ miles north of Adamsville, and 1/8 of a mile east of the main Henderson-

Adamsville Road. Best exposed fossil beds are in the valley 250 yards east of Dave

Weeks’ house, along the headwaters of Coon Creek. Coon Creek is a small stream flowing northward into White Oak Creek, a tributary of Tennessee River (Wade, 1926, page. 9).

“Thompson Farm Fossil Bed” – Coon Creek Formation, along Melton Creek near

Enville, Tennessee (Conover, 1990, in Dunagan and Gibson, 1993).

Merchantville/Marshaltown Formation Spoils or Railroad Bridge Spoils – located on the north side of the canal near a railroad bridge, 39°32’49”N, 75°42’57”W, New

Castle County, Delaware (Feldmann et al., 2013).

Deep Cut Locality– located about one kilometer east of Delaware Route 896 on the C and D Canal, New Castle County, Delaware, 39°32’44”N, 75°42’57”W. Contains in situ material from the Merchantville, Marshalltown, and Englishtown Formations

(Feldmann et al., 2013).

INSTITUTIONAL ABREVIATIONS

USNM - United States National Museum of Natural History, Smithsonian

Institute, Washington, District of Columbia; MMNS – Mississippi Museum of Natural

Science, Jackson, Mississippi; MPPM – Memphis Pink Palace Museum, Memphis,

Tennessee; GAB – Gale A. Bishop Collection, loaned from South Dakota School of

Mines; B – Gale A. Bishop Collection, loaned to Kent State University without specimen numbers, material collected at the Blue Springs Locality; KSU – Kent State University,

Kent, Ohio; SDSM – South Dakota School of Mines, Rapid City, South Dakota; GSUM

– Georgia Southern University Museum, Statesboro, Georgia 30460-8064 (referenced as

GSCM or Georgia Southern College Museum in Bishop, 1988); MGUH – Museum

Geologicum Universitatis Hauniensis, Copenhagen, Denmark; MHN LM – Musée d’Histoire naturelle (“Musée Vert”), Le Mans (Sarthe), France.

INTRODUCTION

The paleontology of the Coon Creek Formation (Maastrichtian, of Tennessee and

Mississippi) has been a popular area of study since the 1920’s, beginning with a thorough assessment by Bruce Wade (1926). These descriptions of marine invertebrate taxa have been the starting point for all subsequent paleontological research of the fossil invertebrates of the formation. The Coon Creek Formation is fossiliferous, with excellent preservation of a diverse assemblage of decapod taxa, first described by Rathbun (in

Wade, 1926). An assemblage of so many fossil decapods is a rarity, both in quantity of specimens as well as diversity. The vertebrate taxa (Carpenter et al., 2003; Gibson et al.,

2008; Gallagher et al., 2012; Stringer, 2003; Whetstone, 1977), ichnofossils (Hall and

Savrda, 2008), and associated sediments have been addressed from the formation or correlative units within the associated seas of North America. However, little has been done to qualify or quantify the decapods of the formation as a whole since Wade’s original work, except for those works by Bishop (1983; 1985; 1991), which focused heavily on the Dakoticanceridae, nor to compare the decapods of correlative units in the region.

This work aims to compile all known decapod taxa of the Coon Creek Formation and to present the taxa according to recent systematic schemes. The second aim of this work is to compare these taxa to correlative units in the Western Interior Seaway, the

Atlantic Coastal Plain, and the Gulf Coastal Plain in order to address the hypothesis that the Mississippi Embayment represents an ecotone for North American decapod fauna. A secondary objective for this work incorporates a taphonomic and paleoecological description of the unit.

i. Tectonic setting

The Cretaceous-Paleogene gulf, known as the Mississippi Embayment, was located on the southern margin of North America during the Maastrichtian. It comprises a southwestward plunging trough that has been filled with about 1.5 kilometers of

Cretaceous and Cenozoic sediments. The embayment is underlain by an early Paleozoic graben and basin fault complex (Cox and Van Arsdale, 2002). The subsidence of the embayment is attributed to forming the Gulf of Mexico, but Cox and Van Asdale (2002) proposed that because sea floor spreading ceased in the Gulf of Mexico 60 million years before the formation of the embayment that the subsidence was caused by the passage of the faulted crust over the Bermuda hotspot during the mid-Cretaceous. The associated volcanism began around 115 million years ago and ended around 65 million years ago in central Mississippi (Cox and Van Arsdale, 2002).

ii. Stratigraphy

The Coon Creek fauna was dated as early Maastrichtian by Russell and Parks

(1975), but Bishop (1986, p. 119) re-dated it as latest Campanian to earliest

Maastrichtian, based on using the Baculites reesidei zone (Obradovich and Cobban,

1975) as the base of the Maastrichtian boundary.

The beds containing the Coon Creek fauna are located in northern Mississippi and southern Tennessee, near what was then the northern end of the Mississippi embayment at around 30 - 35° N (Mancini et al., 1995; Fig. 1 and 2: see also Schuchert, 1955, and

Blakey, 2011). Mancini et al. describe the Late Cretaceous section through the region as a succession of coastal regression and transgression cycles, with the Coon Creek beds

14 comprising part of a progradational to regressive cycle (Fig. 3 and 4). In simplified terms

(temporarily setting aside some nomenclatural complications), the section enclosing the

Coon Creek beds comprises some basal chalks and marls (the Demopolis Formation) that become more argillaceous up-section (the ‘Coon Creek Formation’), which in turn pass up into non-calcareous and non-glauconitic, mainly fluvial sands and clays (the ‘McNairy

Sand’). Wade (1917) described the Coon Creek beds as consisting specifically of a lower clay zone (the “Transitional Clay” of Conant, 1941) that grades down into the Demopolis marls below) and an upper “ferruginous clay horizon” that presages the overlying sands.

In its type area of Coon Creek, in the McNairy County of Tennessee, the Coon Creek

Formation consists of about 58 meters of marine sandstone and shale. Being at the end of a comparatively long and narrow embayment, the section is predominantly (but not entirely) siliciclastic, particularly compared to same-age beds that are farther from the coast: it passes southward into marine sands, clays and limestones of the Ripley

Formation, while it grades and pinches northward into the McNairy Sands (Smith, 1989).

(The Coon Creek beds have at various times and places been regarded as a clay-rich tongue of the Ripley Formation.) Mancini et al. (1995, 1996) identify the chalk as the maximum flooding surface, the shales as progradation during highstand, and the sands as progradation and regression prior to the next transgression (Fig. 3).

The stratigraphic nomenclature applied to this sequence has some complications, especially regarding the question of whether the Coon Creek beds rank as a formation

(the Coon Creek Formation), or just as a member (the Coon Creek tongue of the Ripley

Formation). Russell and Parks (1975) provide a detailed summary of the early literature:

“Conrad (1858, p. 324) informally proposed the name Ripley, but Hilgard

(1860, p. 83-95) first used the term in a formal sense to describe the uppermost marine sediments of Late Cretaceous age near Ripley, Mississippi. He included units now known as the Ripley Formation, the Owl Creek Formation, and also limestones and sands presently known to be of Tertiary age (Harris, 1896, p. 18-

25). Safford (1869, p.417-419) later used the term Ripley Group in a provisional sense and also included the basal Clayton limestones in the Ripley.

“Stephenson (1914, p. 17-18, 22) named a nonmarine sequence of beds in the Ripley Formation in Tennessee and northern Mississippi the McNairy Sand

Member from exposures along railroad cuts in southern McNairy County. Later

Wade (1917a) used the term Ripley Formation to include the Coon Creek

Member, the McNairy Sand Member, and the Owl Creek Member. Subsequently,

Stephenson and Monroe (1937, p. 808-809) excluded the Owl Creek from the

Ripley Formation on recognition of a regional unconformity at the base of that formation. In Tennessee, Sohl (1960, p. 9-22) formally recognized the

Transitional Clay, the Coon Creek as a tongue, and the McNairy Sand as members of the Ripley Formation.”

In 1960, Sohl (1960, p. 9-22) formally recognized the section as comprising the “Transitional Clay”, the “Coon Creek tongue” and the McNairy

Sand”, all as members of the Ripley Formation. The name Coon Creek Formation was raised from the rank of member to its own formation on the 1966 geologic map of Tennessee (Hardeman et al., 1966). In 1975, Russell and Parks Proposed:

“The Coon Creek Formation is composed of well-sorted fine-grained glauconitic, marine sands and clays that are characteristically dark gray. They weather initially to greenish gray and finally to pale reddish brown. There are, typically, two facies, the lower calcareous sand and clay that grades downward through a transitional clay sequence into the Demopolis marls; and an upper facies of sand and clay with large, flattened concretions. The combined thickness of both is approximately 140 feet,” (Russell and Parks, 1975, pg. A35).

Since the Russell and Parks paper, usage of Coon Creek Formation and

Coon Creek tongue has been inconsistent. For simplicity, this paper will refer throughout to “the Coon Creek Formation” and localities will be noted as to whether they are in Tennessee or Mississippi.

METHODS

This study primarily examined Bishop’s Collection, on loan from the South

Dakota School of Mines, and newly collected material of roughly 500 specimens from the Blue Springs Locality, New Albany, Mississippi (Fig. 2), collected over the course of four days during December, 2012 and May, 2013. These specimens were compared to previously published studies and to other collections of fossil decapods, including material at Kent State University. Comparisons were made during two visits to the U.S.

Museum of Natural History in Washington, D.C. Collections from both the Pink Palace

Natural History Museum in Memphis, Tennessee, and the Mississippi Museum of

Natural History in Jackson, Mississippi were obtained through loan for further study at

Kent State University and have been described and illustrated where appropriate. All specimens were prepared (when necessary), whitened, and photographed with a Nikon

D3100 digital camera.

All Coon Creek species were compared to age equivalent species from the Gulf

Coastal Plain, the Western Interior Seaway, and the Gulf of Mexico, using the taxonomic systematic scheme as revised by Schweitzer et. al.(2010).

Taphonomic and sedimentologic studies were conducted through petrographic analysis, scanning electron microscopy, and energy dispersive x-ray spectrometry. For use in the SEM and EDX, specimens were sectioned and polished to 1200 grit scale after impregnation in epoxy when necessary. Specimens were mounted on aluminum stubs using double sided carbon tape tabs. They were then carbon coated twice to ensure even and thick coating. One sample was gold coated to permit more detailed observations of

Dakoticancer australis cuticle morphology using scanning electron microscopy.

RESULTS

i. Sedimentology

The lithology was studied in detail at an outcrop in New Albany, Union County,

Mississippi, just off exit 73B on Route 78. The unit is massively bedded. In hand sample, the rock is a very micaceous, glauconitic, clay-rich sandy siltstone, with common burrows and small concretions. It is grey-brown to grey-green (Munsell colors). It is weakly cemented with calcite. Some layering is visible in hand sample that is not seen in the outcrop, which may be highlighted or even caused by weathering and re-cementation

(Fig. 6D, 6E). Thin section study shows that the rock consists of approximately 10% pore space, and (recalculating the remaining components to 100%), 49.38% muscovite (ca. ¼ mm), 48.15% clay (<1/16 mm), 12.35% glauconite (ca. ¼ mm), and 1.23% organics (1/8 to ¼ mm).

The glauconite occurs as large and well rounded cryptocrystalline grains that give the sediment bimodal grain-size distribution (Fig. 6D, 7A, 7F). Some glauconite grains are fractured. In some cases, glauconite has clearly replaced fecal pellets, and possibly also foraminiferal tests, given poor resemblance of some grains to tests (Fig. 8A).

Foraminiferal tests are white with infilling opaque brown material, possibly phosphate

(Fig. 8C). Fragments of dark brown, opaque organic material are ubiquitous (Fig. 7E).

Many concretions are black to green on the outside, although some are beige.

Freshly broken surfaces react vigorously to HCl, and cracks in the concretions have infillings of calcite cement. Many show round pits and longer thin burrow-like features on their surfaces. Compositions are given in Table 1. In the example of the dark concretions that was studied in thin section, the external color zone was different from the inside primarily in its color, although the outer zone had large micas and more pores, with a central hollow with secondary calcite infilling (Fig. 6A, 6C, 6H). In the example of the lighter concretions that was studied in thin section, the concretion also had a dark, dense, external rind. It contained glauconite, although the grains of glauconite are sparser and less rounded than those in the host bedrock (Fig. 6G). Its nucleus consisted of large fragments of crab cuticle (Fig. 6B). The fragments are brown, and are layered with considerable pore space between them and around them. Some concretions contain claws and associated fragments of Mesostylus.

Some other concretions are even lighter grey-brown in color: these are weakly cemented, micaceous, and silty to sandy, and do not contain fossils (Fig. 6I, Fig. 7B, 7C,

Fig. 7E). A flat concretion was observed in thin section: its outer rim is dark and fine, brownish-black in color. Some of the rock sample is lightly cemented to the concretion, giving it an outer zone of glauconite, clay, and mica, while internally it consists of very fine clay with some large muscovite grains (Fig. 8B, 8D, 8E), with a central hollow that has been filled with some clay, mica, and organic fragments. This was likely a burrow

(Fig. 6A). No glauconite is present in the concretion: it consists of 60% clay and fine muscovite and 20% large muscovite (1/4 – 1/8 mm). The concretion has subtle layering perpendicular, with bands of coarse micas similarly oriented perpendicular to flattening, presumable therefore representing original bedding.

Some annelid worm burrows are present. These are tube-shaped with pointed ends and wrinkle-like features on their surfaces. Crescentic microstructure is visible in thin section. They share the same colors as the concretions, some being black, hard, and dense, and others being light beige with weakly cemented exteriors. In thin section the burrows have dense outer rinds as seen in the concretions, so they presumably all represent an early phase of diagenesis. Cross-sections of burrows show layers of different lithologies, alternating between coarse grains and fine grains, arranged in crescents parallel to the convex margin of the specimen (Fig. 6F).

Fossil preservation will be covered in detail in the Discussion.

ii. Systematic paleontology

See Rathbun (1918) for schematic of brachyuran morphology.

Order DECAPODA Latreille, 1802

Infraorder ASTACIDEA Latreille, 1802

Superfamily NEPHROPOIDEA Dana, 1852

Family NEPHROPIDAE Dana, 1852

Subfamily NEPHROPINAE Dana, 1852

Genus HOPLOPARIA McCoy, 1849

Type species.—Astacus longimanus Sowerby, 1826, by subsequent designation of Rathbun (1926, p. 129).

Included species.— H. albertaensis Tshudy et al., 2005; H. alpines (Van

Straelen, 1936c) as Homarus alpinus; H. antarctica Wilckens, 1907; H. arbei

Aguirre-Urreta, 1989; H. aspera Harbort, 1905; H. bearpawensis Feldmann, Bishop and Kammer, 1977; H. belli McCoy, 1849; H. benedeni Pelseneer, 1886; H. bennetti

Woodward, 1900; H. beyrichi Schlüter, 1862; H. biserialis Fritsch and Kafka, 1887;

H. blossomana Rathbun, 1935; H. buntingi (Feldmann and Holand, 1971) as

Nephrops bunting; H. catalunica Garassino, Artal and Pasini, 2009; H. calcarifera

Schlüter, 1879; H. collignoni (Van Straelen, 1949) as Palaeohomarus Collignoni; H. columbiana Beurlen, 1938; H. corneti Van Straelen, 1921; H. dentata (Roemer, 1841)

(= H. prismatica McCoy, 1849; Palaeno roemeri Robineau-Desvoidy, 1849); H. dentonensis Rathbun, 1935a; H. edwardsi (Robineau-Desvoidy, 1849) as Nephrops samviensis; H. eocenica Lőrenthey in Lőrenthey and Beurlen, 1929; H. falcifer

Fritsch and Kafka, 1887; H. fraasi (Böhm, 1891) as Homarus fraasi; H. gabbi

Pilsbry, 1901; H. gadzicki Feldmann and Crame, 1998; H. gammaroides McCoy,

1849 (= H. belli McCoy, 1849; H. victoriae Quayle, 1987); H. georgeana Rathbun,

1935; H. gladiator Pilsbry, 1901; H. groenlandica Ravn, 1903; H. hakelensis (Fraas,

1878) as Pseudostacus hakelensis; H. hemprichi (Mertin, 1941) as Paleohomarus hemprichi; H. heterodon (Bosquet, 1854) as Oncopareia Bredaï; H. horrida

Schweitzer, Feldmann et al., 2003; H. intermedia Secretan, 1964; H. johnsoni

Rathbun, 1935; H. kamimurai Kato and Karasawa, 2006; H. kamuy Karasawa and

Hayakawa, 2000; H. klebsi Noetling, 1885 (=H. knetschii Zimmermann, 1944

[imprint 1942]); H. lehmanni Haas, 1889; H. longimana (Sowerby, 1826) (type) (=H. granulosa Bell, 1863; H. punctulata Bell, 1863; H. sulcirostris Bell, 1863); H. mcnairyensis Rathbun, 1926a; H. mesembria Etheridge, Jr., 1917; H. minima

Tribolet, 1876; H. miyamotoi Karasawa, 1998; H. muncki Pelseneer, 1886; H. natsumiae Karasawa, Ohara and Kato, 2008; H. nephropiformis Schlüter, 1879; H. pelseneeri (Van Straelen, 1936a) as Homarus pelseneeri; H. percyi Beneden, 1872;

H. pusilla Secretan, 1964; H. riddlensis Feldmann, 1974; H. saxbyi McCoy, 1849; H. scabra Bell, 1863; H. schluteri Tribolet, 1876b; H. sculpta Secretan, 1964; H. senonensis Forir, 1887; H. seucica Schlüter, 1879; H. tennesseensis Rathbun, 1926a;

H. triboleti Borrisiak, 1904; H. trigeri (A. Milne-Edwards, 1886) as Ptychodus

Trigeri; H. tshudyi Schweitzer and Feldmann, 2001a; H. victoriae Quayle, 1987; H. wardi Quayle, 1987.

Generic diagnosis.— Cephalic ridges and spines developed to varying degrees; well-developed rostrum with or without spines; cervical groove extends about half the distance to the dorsal surface; postcervical groove that extends from the dorsal surface to curve around and merge with the cervical groove; well developed hepatic groove; branchial regions with granules, scabrous ornamentation, or keels; exopod of

22 the uropod with a diaresis; chelae clearly differentiated into crushers and cutters,

(Feldman et al., 2007, p. 703).

Discussion.— The genus has been referred to as a “wastebasket” (Tshudy and

Sorhannus, 2003). However, the species within the genus are strikingly similar, with many characters in common. One might hope that a pattern would emerge to further differentiate among nearly fifty species, but no such pattern has been recognized

(Feldmann, et al., 2007). Hoploparia ranges widely geographically and ranges from the Early Cretaceous to the Miocene (Feldmann, et al., 2007).

HOPLOPARIA TENNESSEENSIS Rathbun, 1926

Figures 9.1-9.6

Diagnosis.— Postorbital keel with large spine at anterior margin. Lacking ornamentation on thorax. Cephalon with post antennal spines and postorbital spines.

Cuticle granular on cephalothorax. Pleura with sulcus on posterior margin. Chelae long and slender with large denticles.

Description.— Carapace narrow and laterally compressed. Cuticle granular.

Rostrum short with small, spinose keels on either side of midline. Postrostral carina with small spines decreasing in size posteriorly. Two well defined antennal spines; one suborbital spine and one, well defined postorbital spine at anterior end of postorbital keel. Inferior groove, cervical groove, intercervical groove, and postcervical groove deep and well defined with faint continuation in a concave upward arc between the cervical and postcervical groove. Terga weakly granular or punctate with a small node on anterior margin at the transition point between the first tergum and pleuron; posterior margin of pleura with arcuate sulcus. Pleural ventral

23 margin sigmoidal, terminating in an acute point at posterior edge. Chelae with robust and pustulose palm becoming wider distally; fingers slightly arcuate to straight and elongate with very low, flat denticles proximally, becoming pointed distally.

Material examined.— Specimen B-001; B-005; MPPM 72.46.445; B-005;

MPPM-007; B-008; MPPM-003; MPPM-004; MPPM-005; MPPM-007; MPPM

1972.46.446; MPPM 1972.46.447. Measurements, in millimeters, are given in the

Appendix: Specimen catalogue.

Occurrence.— GAB Blue Springs locality 37 (34°23’31.49”N,

88°53’06.94”W), Mississippi, Coon Creek Formation (Late Cretaceous); Type locality at “Dave Weeks Place” (35°07’23.25”N, 88°33’26.53”W), on Coon Creek in northeastern McNairy County, Tennessee. Other localities include the state line cut on the Southern Railway in McNairy County, Tennessee, one mile northwest of

Wenasoga, Mississippi; on the line between Selma and Ripley Formations as well as a few hundred yards west of the station near Wenasoga, Mississippi (Rathbun in

Wade, 1926 p. 187) as well as in Union County at Lee’s old mill, (34°32’29.32”N,

88°54’43.13”W), located 2 miles northeast of Keownville on the road to Molino

(Rathbun, 1935, p. 25).

Discussion.— In Rathbun’s initial description of Hoploparia tennesseensis (in

Wade, 1926, p. 187), few carapaces are preserved and described. The lengths of specimens in the present study are congruent to the species, the type measuring a total of 48.6 mm and study material being of similar size. Like Rathbun’s material, the

Blue Springs Locality specimens are cracked along the midline and possess a cervical and inverted, Y-shaped hepatic groove that are also well defined.

The presence of more material aids in verification of the species and illustrates variation between individuals. Most notably, specimens from the Blue Springs

Locality vary in size. Most carapaces are about the same size as Rathbun illustrated and described, but two carapaces are preserved in a different orientation, are more inflated, and are generally larger than those found preserved lying on their side. A number of claws were also found, though they are not articulated with any carapaces.

Claws also show a range in size and form. Robust claws possess a long, large palm that increases in size distally. Dactylus and propodus of these claws are broken either at the base or shortly distally. Claw segments of the dactylus and/or propodus, on the other hand, possess long, slender, weakly arcuate fingers with pronounced denticles that become very pointed distally. One claw, preserved on specimen MMNH

1972.46.446 illustrates this form, fully articulated. Another, impressive specimen

MMNH 1972.46.447, does not possess the palm of the claw but is extremely large.

The type specimen of Hoploparia tennesseensis is in fact a claw and was not associated with an carapace specimens. Most notably, this claw possesses broad, flat tubercles that become oblong punctae on the finger. Rathbun also described carapaces that match the form of Hoploparia tennesseensis from the Coon Creek Formation in a later publication, to which she referred to as H. tennesseensis (Rathbun, 1935).

Rathbun also described a new species, Hoploparia mcnairyensis (in Wade,

1927). One specimen, loaned from Vanderbilt University, possessed a longer rostrum and five distinct carinae on the carapace. The degree and size of granulation was also recognized as being larger and more prominent than that of Hoploparia tennesseensis.

Rathbun expressed uncertainty that her specimen and that of the loan from Vanderbilt

25 were in fact the same species. Further observation of the specimen USNM 73118 of

Hoploparia mcnairyensis at the National Museum of Natural History aided in clarification that all specimens illustrated in this study are in fact Hoploparia tennesseensis.

HOPLOPARIA MCNAIRYENSIS Rathbun, 1926

Diagnosis.— Rathbun (in Wade, 1926) did not provide a diagnosis and present material observed did not yield a definitive diagnosis of the species.

Description.— “Carapace behind the cervical groove is granulated.

Granulation is scarce immediately in front of the cervical groove, becoming more crowded and flattened toward the front and middle of the carapace. There is also a spine near the orbit. A second specimen also has granulation and a long rostrum. The carapace has a median carina with a lateral carina on either side. The lateral carina are slightly more prominent and irregular, possessing spines and tubercles where as the median carina do not. There is also a blunt, uneven ridge that extends from the orbital angle to the cervical suture with two interruptions,” (Rathbun, in Wade, 1927, p.

187).

Material examined.— No. 10272 (Specimen from Vanderbilt University?, described by Rathbun, 1926, only); USNM No. 73118.

Occurrence.— Half a mile northwest of Gravel Hill, McNairy County,

Tennessee, on line between Selma and Ripley formations; State Line cut, in McNairy

County, Tennessee, near Wenasoga, Mississippi; Coon Creek Formation,

Maastrichtian.

Discussion.— Rathbun (1927) stated her uncertainty that the two described specimens of Hoploparia mcnairyensis were the same species, citing a need for more material. Having observed specimen USNM No. 73118, and without access to the

Vanderbilt University specimen, I am unable to conclude that any material from this study, including all material from the Blue Springs Locality, is representative of H. mcnairyensis.

HOPLOPARIA GEORGEANA Rathbun, 1935

Plate 1; Figures 7, 8, 9

1935. Hoploparia georgiana. Rathbun, Geological Society of America Special Papers, 3: p. 25, pl.

9, figs. 9-12.

2010. Hoploparia georgiana. Schweitzer et al., Crustaceana Monographs, 10: p. 29.

Diagnosis.— Left and right claws, carpus (?). Cuticle granular. Upper surface of claw possessing anteriorly directed spines; first two spines form a pair parallel to the point of articulation with the carpus, the outermost of which is directed laterally, followed by a row of about three spines that form the upper keel of the claw, beginning at outer margin of upper surface continuing to end at the inner margin, close to the articulation of the dactylus. Lower margin becoming increasingly narrow from the upper margin, becoming keeled toward proximal margin. Articulated joint of merus and carpus (?) possessing widely spaced spines (direction and placement of fragment indiscernible).

Material examined.— GAB 37-686, length: 25.92 mm, height: 15.3 mm;

GAB 37-888, length: 21.44 mm, height: 13.72 mm, GAB 37-1147, length: 40.74 mm, height: 16.98 mm; GAB 37-964, length: 27.02 mm, height: 14 mm, width: 10 mm;

GAB 37-51, length: 25.7 mm, height: 13.7 mm.

Occurrence.— Rathbun (1935) described the holotype from Maryland at an erosional exposure at Brightseat, Prince Georges County from the Monmouth

Formation, Upper Cretaceous (p. 26). Specimens described herein occurred at the

Blue Springs locality, Coon Creek Formation, Union County, Mississippi,

Maastrichtian.

Discussion.— The identification of these claw fragments extends the geographic range of the species from the Atlantic Coastal Plain to the Mississippi

Embayment and is the third species of Hoploparia found at the Blue Springs Locality.

These specimens can clearly be differentiated from Hoploparia tennesseensis in that they possess spines that are not observed on chelae of H. tennesseensis. Because there are no articulated claws with H. mcnairyensis, it is not impossible that the claws, H. georgeana are a junior synonym of H. mcnairyensis, but ornamentation of H. mcnairyensis does not possess such dramatic spines. More specimens are needed to confirm that H. mcnairyensis and H. georgeana are not the same species.

Infraorder AXIIDEA de Saint Laurent, 1979

Superfamily CALLIANASSOIDEA Dana, 1852

Family CALLIANASSIDAE Dana, 1852

Genus MESOSTYLUS Bronn and Roemer, 1852

Type species. – Pagurus faujasi Desmarest, 1822.

Included species.— Mesostylus faujasi (Desmarest, 1822); Mesostylus mortoni (Pilsbry, 1901) as Callianassa mortoni.

Generic diagnosis.— “Merus of major cheliped longer than high, upper margin convex, lined with large granules; large knob on distal margin articulating

28 with carpus. Carpus longer than high, distal margin concave, serrate, lower distal margin with flange extending onto outer surface in weak rim separated from distal margin by prominent sulcus, distal margin of flange serrate. Proximal margin of manus at 100–110º angle to lower margin; upper and lower margins finely serrate; fingers with stout teeth on occlusal surfaces. Major chela exhibiting notable dimorphism; chelae stouter and more rectangular in presumed males, chelae more slender and higher proximally in presumed females. Minor chela longer than high, highest proximally; fingers with parallel ridges. Pleonal somites apparently smooth.”

(Schweitzer and Feldmann, 2012, p. 17).

MESOSTYLUS MORTONI (Pilsbry, 1901)

Fig. 10.1, 10.6

1901. Callianassa mortoni . Pilsbry. Proceedings of the Academy of Natural Sciences of Philadelphia,

p. 112, pl. 1, figs. 1–7.

1926. Callianassa mortoni. Rathbun, In B. Wade (ed.), The fauna of the Ripley Formation of Coon

Creek, Tennessee. U.S. Geological Survey, Professional Paper, 137: p. 188, pl. 67, figs. 1, 2, 4–9.

1935. Callianassa mortoni. Rathbun, Geological Society of America, (Special Paper), 2, i–viii,: p. 29.

1941. Protocallianassa mortoni (Pilsbry, 1901).Mertin, Nova Acta Leopoldina, 10 (68): p. 208.

1962. Protocallianassa mortoni (Pilsbry, 1901). Roberts, New Jersey Geological Survey Bulletin, 61:

p. 169, pl. 81, fig. 8, pl. 83, figs. 1–6.

1991. Protocallianassa mortoni (Pilsbry, 1901). Bishop, Mississippi Geology, 12 (1): 12-13.

2012. Mesostylus mortoni (Pilsbry, 1901). Schweitzer and Feldmann, Bulletin of the Mizunami Fossil

Museum, no. 38, p. 19.

2013. Mesostylus mortoni (Pilsbry, 1901). Feldmann et al., Bulletin of the Mizunami Fossil Museum,

no. 39: 12-15.

Description.— “Merus longer than high, upper surface convex; proximal margin at about 60º angle to lower margin, with projection near lower margin

29 articulating with ischium; lower margin nearly straight; distal margin concave, articulating with long projection of carpus; outer margin convex, with row of large granules, very large swelling ornamented with tubercles distally, knob on distal margin articulating with carpus. Carpus longer than high, with sinuous proximal margin, with projection at upper margin to articulate with merus, marked concavity below it, then becoming convex at lower margin; lower margin sloping downward distally so that entire carpus becomes higher distally; upper margin weakly convex, serrate; distal margin concave, serrate where articulating with manus; at lower distal corner, flange projecting anteriorly, extending onto outer surface as weak ridge separated from distal margin by prominent sulcus, distal margin of flange serrate; strongly vaulted on outer surface, slightly depressed on inner surface. Manus longer than high, strongly vaulted on outer surface; proximal margin sinuous, with weak projection centrally for articulation with carpus, lined with setal pits, entire margin oriented at about 100º angle to lower margin; lower and upper margins serrate, rimmed, nearly straight, manus becoming slightly less high distally; distal margin straight where intersecting upper margin, then directed slightly obliquely to intersection with fixed finger, serrate; inner surface of manus flattened, with row of setal pits along upper margin. Fixed finger with triangular central spine; movable finger with basal blunt spine and long, blunt tooth centrally; outer surface of movable finger with two rows of setal pits; row of setal pits along upper margin of inner surface of movable finger. Movable finger of minor chelae with two or three granular keels with rows of setal pits or granules between them. Ischia of other pereiopods

30 longer than high, smooth. Abdominal somites poorly known, smooth,” (Schweitzer and Feldmann, 2012, p. 19).

Description of material.— Specimens of chelae and carpus, some of which bear both major and minor claws. Merus and carpus longer than high; narrow. Cuticle smooth. Dactyl and fixed finger edentulous; proximal margin of carpus greater than

90°, top and bottom margin of carpus and merus finely serrate, distal outer margin of carpus with finely serrate keel or flange extending from lower margin upward and proximally from the distal margin of the carpus ¼ of total height; row of tubercles on the lower margin of the merus.

Material examined.— B-014, no measurements; GAB 37-103, length of merus (l): 15.58 mm, height of merus (h): 13.92 mm; MPPM 1974.35.16, l: 17.48 mm, h: 15.28 mm; MPPM 72.46.456, l: 15.93 mm, h: 12.38 mm; MPPM 016, no measurements; MPPM 022, no measurements; MPPM 020, h: 17.22 mm; MPPM

019, l: 20.28, h: 15.4 mm; MPPM 013, no measurements; MPPM 023, l: 21.52 mm, h: 17 mm; MPPM 021, (major claw) l: 19.34, h:18.26, (minor claw) l: 16.04, h: 14.45 mm; MPPM 018, no measurements; GAB 37-237, l: 11.66 mm, h: 10.14 mm; MPPM

017, no measurements; MPPM 006, no measurements; MPPM 024, l: 15.64 mm,

13.94 mm; GAB 37-208, (major claw) l: 13.64 mm, h: 12.69 mm, (minor claw) l: 9.4 mm, h: 8.78 mm; GAB 37-177, h: 16 mm; GAB 37-296, no measurements.

Occurrence.— Early Campanian to Maastrichtian throughout the Atlantic and

Gulf Coastal Plains and the Mississippi Embayment (Schweitzer and Feldmann,

2012; Feldmann et al., 2013).

Discussion.— Schweitzer and Feldmann (2012) proposed to restrict

Protocallianassa to Protocallianassa archiaci, the type species, and to refer

Protocallianassa mortoni to Mesostylus because it possesses the well developed flange on the lower distal margin of the carpus, the proximal margin of the merus is at an angle greater than 100°, and it has a ridge of large tubercles on the merus (p. 17).

The tubercles on the merus of the Mississippi specimens are not well preserved and generally are observed as pits, but are verified as tubercles on specimen MPPM 019.

Modern studies such as Tsang et al. (2008) and Sakai (2011) have reevaluated the systematics of extant Axiidea, but necessary morphology for classification is not preserved in the fossil record. Because of this, the placement of Mesostylus mortoni has been maintained in this study and reflects that of Schweitzer and Feldmann

(2012) and Feldmann et al. (2013). This species of mud shrimp is relatively wide spread geographically and ranges from New Jersey to Tennessee and Mississippi

(Feldmann et al., 2013).

Infraorder ACHELATA Scholtz, 1995

Superfamily PALINUROIDEA Latreille, 1803

Family PALINURIDAE Latrielle, 1802

Genus LINUPARUS White, 1847

1847. Linuparus. White, List of Specimens of Crustacea in the Collection of the British

Museum, London, p. 40.

1849. Podocratus. Becks MS., in Geinitz, Das Quadersand-Steingebirge oder Kreidegebirge in

Deutschland, Freidberg, p. 96.

1862. Podocrates. Schlüter, Deutsche geol. Gesell. Zeitschr., vol. 14, p. 710.

1897. Linuparus (part). Ortmann, Am. Jour. Sci., 4th ser., vol. 4, p. 296.

Type species.— Palinurus trigonus von Siebold, 1824.

Included species.— L. adkinsi Rathbun, 1935a; L. africanus Glaessner, 1932a;

L. bererensis Secretan, 1964; L. bigranulatus Glaessner, 1930; L. canadensis

(Whiteaves, 1885) as Hoploparia? canadensis = L. atavus Ortmann, 1897; L. carteri

(Reed, 1911) as Thenops carteri; L. dentatus Van Straelen, 1936a; L. dulmenensis

(Geinitz, 1849-1850) (Schlüter, 1899) as Podocrates dulmenensi; L. dzheirantuiensis

Feldmann, Schweitzer et al., 2007;L. eocenicus Woods, 1925; L. euthymei (Roman and Mazeran, 1920) as Podocrates euthymei; L. grimmeri Stenzel, 1945; L. hantscheli

Mertin, 1941; L. japonicas Nagao, 1931; L. kleinfelderi Rathbun, 1931a; L. korura

Feldmann and Bearlin, 1988; L. laevicephalus Mertin, 1941; L. macellarii Feldmann and Tshudy, 1988; L. petkovici Bachmayer and Marković, 1955; L. pustulosus

Feldmann in Feldmann et al, 1977; L. richardsi (Roberts, 1962) as L. (Podocratus) richardsi; L. schluteri (Tribolet, 1874) as Podocrates schlüteri; L. scyllariformis

(Bell, 1858) as Thenops scyllariformis; L. spinosus Collins and Rasmussen, 1992; L. stolleyi (Haas, 1889) as Podocrates stolleyi; L. straili (Forir, 1887) as Thenops straili;

L. somniosus Berry and George, 1972; L. sordidus Bruce, 1965; L. tarrantensis

Davidson, 1963; L. texanus Rathbun, 1935a; L. trigonus Siebold, 1824 as Palinurus trigonus; L. vancouverensis (Whiteaves, 1885) as Hoploparia (?) canadensis; L. watkinsi Stenzel, 1945; L. wilcoxensis Rathbun, 1935a.

Generic diagnosis.— “Carapace sub-rectangular with three longitudinal keels; rostrum absent; supraorbital spines close to median line, fused to form plate or separated by indentation; well marked cervical groove; longitudinal median carina extends from posterior margin to cervical groove; prominent ridges form swelling of

33 flank just posterior to cervical groove. Pleon with variously spinose margins on pleurae and keeled terga. Pereiopod 1 stout; pereiopods 2-5 long, slender; telson sub- rectangular,” (Feldmann et al., 2013, p. 15).

Discussion.— Specimens from the Coon Creek possess all diagnostic characters of the genus. Important features not preserved on Rathbun’s (1926) specimen are observed including keeled terga, inflated flanks (stridulating region or appareil stridulant of Secretan, 1964), and spines on the margins of the pleura.

Podocratus is herein treated as a junior synonym of Linuparus. Subgeneric diagnoses are not included in the following discussion and are not utilized in classification because they lack utility as suggested by Feldmann et al. (2007). Features used to determine subgeneric placement are not mutually exclusive. Linuparus (Linuparus) as well as Linuparus (Thenops), Linuparus (Podocrates), and Linuparus (Eolinuparus) are therefore not recognized.

LINUPARUS new species

Figure 11.1 – 11.7

1926. Podocratus canadensis. Rathbun, in Bruce Wade, U. S. Geol. Surv., Prof. Pap. 137, p. 185, pl. 63,

figs. 12, 16; U. S. Nat. Mus., Bull. 138, p. 134, pl. 35, fig. 2, pl. 36.

1983a. Linuparus canadensis. Bishop, G.A., Journal of Crustacean Biology, 3(3): 417-430.

1986. Linuparus canadensis. Bishop, Journal of Crustacean Biology, 6(3): 345-346.

1991. Linuparus canadensis. Bishop, G. A., Mississippi Geology, 12(1,2): 8-17.

Diagnosis.— Large to medium sized carapace for the genus. Gastric region aristate. Pleurae with three spines on lateral keel and ventral margin, one or two pustules at center of pleura.

Description.— Carapace sub-rectangular. Rostrum distinct, bifid; supraorbital spines close to midline and separated, single pair of post-supraorbital spines. Cephalic region with marginal keel possessing three or more forward directed spines and antennal keel possessing 4 or more forward direct spines. Cephalic region slightly inflated with aristate gastric region outlined by five large, forward directed spines and a variable range of small spines in between the larger ones. Cephalic region sparsely granular. Cervical groove parabolic, deep and well defined. Thoracic region with three longitudinal keels with numerous, anteriorly directed spines of nearly equal size. Axial keel possesses seven to nine spines, lateral keels with 10 to 15 spines.

Slightly inflated flank posterior to cervical groove indicating appareil stridulant (of

Secretan, 1964). Nearly vertical ventral walls of carapace. Flank with prominent, finely granular marginal rim. Thoracic region uniformly granular. Pleon sub- rectangular and coarsely punctate. Axial keel of pleon with pair of anteriorly directed spines with smooth, posterior-laterally sloping sulcus rising between the two spines on the first through fourth somite, spines on fifth and sixth somite posteriorly directed. Angular demarcation between terga and pleura with three outward projecting spines, declining posteriorly. Pleura with 3 spines at ventral margin and two pustules at the center of the second and third pleura, with only one on fourth pleuron. Basal antennal segment spinose on margin and on central keel of dorsal surface. Right mandible rectangular with small, arcuate, anterior process.

Material examined.— Specimen loaned from Richard Keyes’ personal collection, MMNH 2000.1.11; 1972.2.444; MPPM-001; MPPM-002; KSU-009-1; B-

006. Other specimens: SDSM 37-1115, SDSM 37-1156, SDSM 37-30, SDSM 37-29,

SDSM 37-1141, SDSM 37-23, and SDSM 37-31. Measurements, in millimeters, are given in the Appendix under Specimen Catalogue.

Occurrence.— Blue Springs Locality (GAB 37; 34°23’31.49”N,

88°53’06.94”W), Mississippi, Coon Creek Formation; Maastrichtian, Late Cretaceous.

Other specimen (No. 10272) collected ½ mile northwest of Gravel Hill, McNairy County,

Tennessee, on line between Selma and Ripley Formations near 35°44’51.08”N,

87°03’14.88”W (Rathbun, in Wade, 1927, p. 186).

Discussion.— Rathbun (in Wade, 1927; p. 185; pl. 63, fig. 12, 16) described the specimen as Podocratus canadensis. Her description of the available material conforms to the morphology of the new specimens from the Coon Creek Formation.

That being said, Rathbun’s material consisted of the thoracic region of the carapace only. The fragmented carapace is tricarinate behind the cervical groove and the keels possess irregularly-sized spines. The median keel has nine spines with the third to last being double (Rathbun in Wade, 1927 p. 185). Rathbun also noted that the Coon

Creek specimen possessed a longitudinal sulcus or furrow around the median keel as was described from the specimens of Linuparus canadensis from Alberta, Canada.

The longitudinal furrow was not observed in the studied material, but if the cause of this furrow is in fact post-mortem deformation which it almost certainly is, then the character is not of taxonomic significance.

When compared to descriptions of more complete specimens of Linuparus canadensis, the Blue Springs Linuparus new species is different in size, form of gastric region, and nature of the pleura. There are only a handful of species of

Linuparus from North America. Linuparus canadensis is possibly the most widely

36 distributed in the Cretaceous (Feldmann and McPherson, 1980, p. 12). Feldmann and

McPherson (1980) suggested that L. canadensis occurs from Alberta, Canada to

Louisiana, USA; however, observations from this study lead to the conclusion that this distribution should exclude the material from the Coon Creek Formation of

Mississippi.

Because of the fragmental nature of decapod preservation, only portions of the whole animal are sometimes used to describe new species. Linuparus species infrequently have the pleon available for comparison, yet some species are described solely on the pleon, such as Linuparus texanus Rathbun, 1935. Because the Coon Creek

Formation specimens possess both the carapace and the pleon, the specimens can be compared to the morphology of most previously described species (Table 2).

Stenzel (1945) noted the close similarity and relationship of Linuparus canadensis with Linuparus watkinsi. Linuparus watkinsi also possesses three keels on the thorax, but the median keel has seven spines, though Stenzel (1945) noted that these are the major spines and that a few accessory spines also exist. Lateral keels possess twelve major spines. Both Linuparus canadensis and Linuparus watkinsi, therefore, have about 7-8 prominent spines on median keel (illustrated in Stenzel, 1945, plate 34 and described in

Table 2 of Feldmann et al., 2007, p. 709), similar in character to the nine median spines on the Coon Creek specimens. The lateral keels had twelve spines on Rathbun’s Coon

Creek specimen as well, just as in Linuparus watkinsi.

Whiteaves (1985) initially described Hoploparia? canadensis as having large conical tubercles on three keels on the cephalon with five tubercles forming an isosceles triangle of the gastric region. Rathbun (1935) also described the gastric region of L.

37 canadensis as having five spines that form an isosceles triangle. Stenzel (1945) described the cephalon of Linuparus watkinsi in great detail. The cephalon of Linuparus watkinsi has nine keels, three of which make the oval shaped gastric region, much like the Coon

Creek specimens. The antennal keels (beginning at the fronto-lateral margin), the supraorbital keels, and the suborbital keels (gently curving and spreading posteriorly) are also spinose.

The pleon of Linuparus watkinsi also has three spinose keels, like the thorax. The lateral keels have only two spines on each lateral somite, except for the first segment, which has none. The median keel has one central spine on the first and second segment, two laterally compressed spines on the third segment, four spines on the forth segment and two simple spines on the fifth segment. The telson has one spine and the uropods are narrow with sigmoidal edges. The upper surface of each pleonal segment of Linuparus canadensis has a tubercle in the center, on its anterior edge, and another one on the margin of each of the sides at the tergal-pleural margin.

Linuparus texanus is a specimen of the pleon only (Rathbun, 1935, p. 73). Each segment has two median spines as well, the pleura are rounded, and the edge of the pleura is separated from the tergum by a smooth, blunt ridge that terminates in a stout and conical spine, pointing anteriorly.

Linuparus wilcoxensis has lateral keels with “crowded granules” and a median ridge with granules that becomes spinose with age (Rathbun, 1935, p. 74). The pleon is coarsely punctate, but lacks carinae sloping toward the pleura and lacks pleural spines characteristic of the Coon Creek specimens. Linuparus wilcoxensis also possesses one to two spines rather than three on the ventral margin of the pleura. Specimens included in

No. 371498 (103 fragmented specimens) at the National Museum of Natural History of

Linuparus wilcoxensis from the Eocene of the Prairie Creek Formation interestingly possess a rectangular mandible with an anterior process and spines on the pleonal segments pointing both anteriorly and becoming posteriorly, as Linuparus new species does.

Linuparus grimmeri has a median keel bearing about nine spines and lateral keels with about twenty small spines (Stenzel, 1945). The lateral keels on the terga have two spines per somite, except for the first, which has none. Linuparus grimmeri is generally narrow and more finely ornamented species and is very small for the genus.

Linuparus richardsi Roberts, 1962 (Feldmann et al., 2013, p. 15), was collected from the Graham Brick Company pit, Maple Shade, Burlington County, New Jersey from the Merchantville Formation. Another specimen had been collected from the

Marshalltown/Merchantville spoils along the C and D Canal, Delaware. It is diagnosed as having supraorbital spines that are close and fused, but lacking post-supraorbital spines.

The gastric region is triangular and outlined by five large spines. The lateral keels on the thoracic region possess spines that decrease in size posteriorly. The cephalic region is smooth rather than granular (Feldmann et al., 2013, p. 15).

The carapace of one specimen of Linuparus new species, MPPM 002, has a more triangular gastric region and is generally large in overall size, but it does not possess irregularly granular ornamentation, and it is not apparent that the spines on the keels decrease in size posteriorly. The poor nature of preservation of the specimen makes comparison difficult. Both MPPM 001 and MMNS 2000.1.11 are both slightly smaller than MPPM 002 as is the specimen from New Jersey, Linuparus richardsi (Feldmann et

39 al., 2013, p. 15, fig. 5). One cannot help but find MPPM 002 to be very reminiscent of

Linuparus richardsi and could easily be mistaken for Linuparus canadensis due to its preservation. Furthermore, Linuparus richardsi is undoubtedly closely related to

Linuparus canadensis.

Though the material from the Coon Creek Formation was determined to be

Linuparus canadensis by Rathbun (in Wade, 1926) and supported by Bishop (1991), new specimens available have more morphological characters preserved. They cannot be assigned to Linuparus canadensis because of the gastric region and ornamentation on the pleura. The species noted to be the most similar to Linuparus canadensis by Stenzel

(1945), Linuparus watkinsi, has a distinct rostrum that is nearly identical to those preserved on small specimens of Linuparus new species. Linuparus watkinsi is also a smaller species, but lacks the same number of spines on the lateral keel of the terga, and the pleura do not have pustules at the center as the Linuparus new species do. Linuparus grimmeri possesses many similarities of the carapace, but the species is much smaller with finer ornamentation and generally less ornamentation on the pleura as well.

Stenzel (1945, p. 410) contrasted Linuparus watkinsi and Linuparus canadensis, citing the shape of the gastric region, the number of accessory spines in the gastric region, and the medial keel of the thorax as being the main differences. Because of this, the characters of the Coon Creek specimens document a new species, sharing characters with Linuparus watkinsi and Linuparus canadensis, as well as possessing a character unique from either species. Figure 6 illustrates key differences between closely related species to those from the Coon Creek. The variable nature of preservation of the carapaces affects approximations of thoracic spines. If pleonal segments did not illustrate

40 consistent morphology and the specimens were not generally large, I do not believe that the thoracic keel characters would be sufficient to describe this species as they vary by specimen, possibly due to age or the individual. For example, specimen SDSM 37-1141 may be an example of a juvenile of Linuparus new species. It has acutely pointed, well pronounced spines and is a great deal smaller than other specimens with nearly identical morphology other than having a slightly larger number of thoracic spines.

LINUPARUS sp.

Diagnosis.— Cuticle punctate. Cephalic spines rounded. Thorax with three longitudinal keels. Spines on lateral keels reduced; spines on median keel not present.

Description.— Carapace medium sized, sub-rectangular, and narrow. Pair of supraorbital spines and pair of protogastric spines, slightly granular and rounded in form.

Two or three spines on cephalic margin but generally smooth. Gastric region elliptic to ovate, slightly inflated with rounded, blunt spines outlining the narrow region and connecting to the bluntly granular median cephalic carina. Anterior-most spine is reduced and located between the protogastric spines. Gastric region slopes steeply toward the cervical groove. Thoracic region with three longitudinal keels; axial keel poorly preserved and not obviously spinose; lateral keel finely nodose with three larger nodes at posterior margin. Somites coarsely punctate with two anterior directed median spines on second and third somite. Sulcus connecting anterior margin of second spine to the tergal- pleural margin.

Material examined.— SDSM 37-1115.

Occurrence.— Blue Springs Locality (GAB 37; 34°23’31.49”N,

88°53’06.94”W), Mississippi, Coon Creek Formation; Maastrichtian, Late Cretaceous.

Discussion.— As illustrated in Table 1, this specimen of Linuparus also lacks key features to be assigned to any of the most closely related species. Because the preservation of the specimen is incomplete – gastric region is compromised, pleurae are not visible or present, specimen is crushed and anterior of cephalon is distorted – I do not believe it is appropriate to use the available characters to assign it to a new species or to an existing one. The most similar species is Linuparus kleinfelderi Rathbun (1931a), but the wide carina are smooth, rather than spinose. Specimen USNM 2009820, Linuparus sp., of the Eagleford Formation from the University of Dallas roadcut in Irving, TX, also has wide keels but keels are granular. The gastric region of the specimen from the

Eagleford Formation is also similar in nature to Linuparus sp. of the Coon Creek

Formation in that it is aristate, but other similar morphology cannot be confirmed.

Infraorder ANOMURA Mac Leay, 1838

Superfamily PAGUROIDEA Latreille, 1803

Family DIOGENIDAE Ortmann, 1892a

Genus PALAEOPETROCHIRUS Bishop, 1991

Type species. – Palaeopetrochirus enigmus Bishop, 1991

Included species.— Palaeopetrochirus enigmus Bishop, 1991

Generic diagnosis.— As for species.

PALAEOPETROCHIRUS ENIGMUS Bishop, 1991

Diagnosis.— “Left chelipeds robust; merus long, triangular in cross section, flattened on bottom and front; posterior edge with en echelon crinkles; carpus short;

42 claw longer than high, outer face convex, lower margin level but slightly sinuous, upper margin strongly arched; fixed finger relatively short, stout, and triangular; dactylus relatively long, narrow, and curved,” (Bishop, 1991, p. 11).

Description.—Left claw; lower margin straight, upper margin forming continuous arc with dactylus when fingers are closed; weakly convex outer surface.

Outer surface with distally directed nodes. Nodes coarsest at lowermost and uppermost margin of propodus.

Material examined.— The holotype, 1700 MGS is illustrated in Bishop (1991, p. 15) and is the sole specimen of the species.

Occurrence.— Bishop (1991) stated that the specimen did not occur with any written information on locality, etc. by the collector, but specimens associated in the box of material left by the collector, Ralph Harris, and described by Bishop, were likely from the Blue Springs Locality, GAB 37, New Albany, Mississippi according to Bishop (1991) and would therefore be Maastrichtian.

Discussion.— The specimen described by Bishop (1991) has not been referenced since the publication and the subfamily (Diogeninae Ortmann, 1892), genus, and species are not recognized by Schweitzer et al. (2010). The species was described in a less widely circulated journal, Mississippi Geology. Because of this, the species has gone unreported since its initial description. The singular specimen is likely from the Coon Creek Formation, but there is no significant evidence to give the author justification to include this potential hermit crab claw, especially without the presence of any other material available. Bishop (1991, p. 11) diagnosed the new genus as having “equal or subequal” claws with the possession of a single specimen

43 of left claw, of which one can only observe the outer surface and part of the lower margin (Bishop, 1991, p. 15, fig. 5, A, B, and C). Bishop also illustrated the “lower edge of merus” (Bishop, 1991, p. 15, fig. D), but did not reference it in his description.

There are many specimens of Paguristes Dana, 1851a in the fossil record, and one species of Parapaguristes Bishop, 1986a, P. tuberculatus. Palaeopetrochirus enigmus is tentatively retained in the Diogenidae, though there is no way of knowing that this species is left-handed.

Infraorder BRACHYURA Latreille, 1802

Section DAKOTICANCROIDA Rathbun, 1917

Family DAKOTICANCRIDAE Rathbun, 1917

Included genera.—Dakoticancer Rathbun, 1917; Tetracarcinus Weller, 1905;

Avitelmessus Rathbun, 1923.

Included species.— Dakoticancer overanus Rathbun, 1917. Dakoticancer australis Rathbun, 1935. Tetracarcinus subquadratus Weller, 1905. Avitelmessus grapsoideus Rathbun, 1923.

Family diagnosis.— “Carapace quadrate, as wide as long or longer than wide; rostrum narrow, bilobed; orbits well developed, rimmed; eyes sheltered by orbits when retracted; anterolateral margins entire; posterior margin nearly straight; medial part of cervical groove weakly developed; gastric regions poorly separated from cardiac and intestinal regions; branchiocardiac groove well developed; pleural sutures located on sides of carapace; fifth pereiopods very reduced, subdorsal; sternum broad, sternites visible to posterior of carapace, sternite 4 with ridge parallel to anterior end,

44 sternites 5, 6, and 7 with granular transverse ridges; sternum of female without longitudinal grooves; lateral portion of posterior part of sternites visible; male pleon with all somites free, lateral terminations on pleonites rectangular, telson rounded triangular; female pleon wide, with long epimeres, all pleonites free; coxae of pereiopods at same level as sternum; first pereiopods isochelous,” (Karasawa et al.,

2011, p. 556).

Genus DAKOTICANCER Rathbun, 1917

Type species. –Dakoticancer overanus Rathbun, 1917.

Included species.— Dakoticancer overanus Rathbun, 1917; Dakoticancer australis Rathbun, 1935.

Generic diagnosis.— “Carapace rectangular to transversely ovoid, length and width about equal, front narrow, rostrum bilobed; orbits well developed; median part of cardiac groove weak, gastric regions hardly separated from cardiac-intestinal region, branchiocardiac groove well developed, pleural sutures on carapace sides; genital openings on coxae, female on third leg and male on the fifth; fifth legs much reduced. Chelae equal,” (Bishop et al., 1998).

Discussion.— Dakoticancer is the type genus of the Dakoticancridae. Bishop,

Feldmann, and Vega (1998) addressed the reproductive structures, intersexuality, functional morphology, feeding structures, and life habit of species of the

Dakoticancridae where material was available. These topics will be expanded upon by species, where applicable, below.

DAKOTICANCER OVERANUS Rathbun, 1917

1917. Dakoticancer overana. Rathbun, Proc. U.S. nat. Mus., 52: 385-391.

1929. Dacoticancer overanus. Glaessner, Fossilium Catalogus I:Animalia, pars 41, p. 134.

1930b. Dakoticancer overana. Rathbun, Proceedings of the United States National Museum, 78: 1-10,

pls. 1-6.

1935. Dakoticancer overana. Rathbun, Geol. Soc. Of America, Spec. Pap. 2, 160p.

1969. Dakoticancer overanus. Glaessner in Moore (ed.), Treatise on Invertebrate Paleontology, R (4)

(2): R400-R533, R626-R628.

1972b. Dakoticancer overanus. Bishop, Journal of Paleontology, 15(3): 631-636.

1974. Dakoticancer overanus. Bishop, Crustaceana, 26(2): 212-218.

1977. Dakoticancer overanus. Bishop, Journal of Sedimentary Petrology, 47(1): 129-136.

1977. Dakoticancer overanus. Feldmann, Bishop, and Kammer, Journal of Paleontology, 51(6):1161-

1180.

1981. Dakoticancer overanus. Feldmann, Géobios, 14 (4): 449-468.

1981b. Dakoticancer overanus. Bishop, The GSC Museum Contributions to Natural History: 383-414.

1983a. Dakoticancer overanus. Bishop, Journal of Crustacean Biology, 3(3): 417-430.

1983c. Dakoticancer overanus. Bishop, Crustaceana 44(1): 23-26.

1984. Dakoticancer overanus. Bishop, Journal of crustacean biology 4.3 (1984): 514-517.

1985c. Dakoticancer overanus. Bishop, Journal of Paleontology, 59(3): 605-624.

1986c. Dakoticancer overanus. Bishop, Published in: Gore, R.H., and K.L. Heck: Crustacean

Biogeography. Crustacean Issues 4. A.A. Balkena, Boston: 111-141.

1986. Dakoticancer overanus. Bishop, Journal of Crustacean Biology, 6(3): 345-346.

1987. Dakoticancer overanus. Tucker, Feldmann, Holland, and Brinster, Annals of Carnegie Museum,

56(7): 275-288.

1991. Dakoticancer overanus. Bishop, Mississippi Geology, 12(1,2):8-17.

1993b. Dakoticancer overanus. Dunagan and Gibson, Tennessee Academy of Science Journal,

68(3):87-93.

1999. Dakoticancer overanus. Bishop, Journal of Crustacean Biology, (Wrong version)***

1991. Dakoticancer overanus. Bishop, Mississippi Geology, 12 (1/2): pg. 14.

1998. Dakoticancer overanus. Bishop, Feldmann, and Vega, Contributions to Zoology, 67(4):237-255.

2003b. Dakoticancer overanus. Bishop, In Gibson, M.A., and Dunagan, S.P., eds., The Coon Creek

Formation of Tennessee: Tennessee Division of Geology Bulletin 86: ??

2011. Dakoticancer overanus. Karasawa, Schweitzer, and Feldmann, Journal of Crustacean Biology,

31: 523-565.

2013. Dakoticancer overanus. Jones, Thesis, Kent State University, December, 2013.

Diagnosis.—“Carapace of moderate size, rectangular, slightly wider than long, widest across epibranchial regions, sides sinuous, slightly convergent posteriorly; high, moderately arched; grooves deep, cervical groove prominent, relatively continuous except across continuous sagittal ridge; pits present at junction of cervical and antennal grooves. Tumid regions granulate, hind margin nearly width of carapace, not developed into a shelf, raised and granulate. Chelae equal, twice as long as high,” (Bishop et al., 1998).

Occurrence.— Pierre Shale in South Dakota, North Dakota, and Montana;

Maastrichtian.

Discussion.—Dakoticancer overanus does not occur in the Mississippi

Embayment in Tennessee or Mississippi, but is very common in correlative units of the Pierre Shale of South Dakota, North Dakota, and Montana within the Western

Interior Seaway (Bishop, et al., 1998). Dakoticancer overanus also illustrates a number of intersex forms that are not observed in any other species of the family

(Jones, 2013).

DAKOTICANCER AUSTRALIS Rathbun, 1935

Figures 12.3 – 12.9

1926. Dakoticancer overana. Rathbun, in Bruce Wade, U. S. Geol. Surv., Prof. Pap. 137, p. 189, plate

LXVII, fig. 3.

1935a. Dakoticancer overana australis. Rathbun, Geological Society of America Special Papers, 2: 1-

160.

1969. Dakoticancer overanus australis. Glaessner in Moore (ed.), Treatise on Invertebrate

Paleontology, R (4) (2): R400-R533, R626-R628.

1983. Dakoticancer australis. Bishop, Journal of Crustacean Biology, 3(3):417-430.

1983. Paguristes whitteni. Bishop, Journal of Crustacean Biology, 3(3): 420.

1985a. Dakoticancer australis. Bishop, Journal of Paleontology, 59(5): 1028-1032.

1986. Dakoticancer australis. Bishop in Gore and Heck (eds.), Crustacean Biogeography, Crustacean

Issues 4, A. A. Balkema, Rotterdam, The Netherlands, p. 292.

1986. Dakoticancer australis. Bishop, Journal of Crustacean Biology, 6(3): 345-346.

1991. Dakoticancer australis. Feldmann and Vega, Annals of Carnegie Museum, 60 (2): p. 165.

1991. Dakoticancer australis. Bishop, Mississippi Geology, 12(1,2):8-17.

1991. Parapaguristes whitteni (Bishop, 1983). Bishop, Mississippi Geology, 12(1-2): 13.

1995. Dakoticancer australis. Vega, Feldmann, and Sour-Tovar, J. of Paleontology, 69(2): fig. 4.

1998. Dakoticancer australis. Bishop, Feldmann, and Vega, Contributions to Zoology, 67(4):237-255.

2010. Paguristes whitteni. Schweitzer et al., Crustaceana Monographs, 10: 54.

Diagnosis.— “Carapace large, slightly longer than wide, widest across branchial and hepatic regions (sides nearly parallel), well differentiated by grooves, ornamented by granules over entire surface. Posterior produced into a fairly discrete shelf, not quite a maximum width of carapace, hind margin slightly raised. Chelae equal, short, stout, and crested, carpal articulation very oblique; fingers short, downturned,” (Bishop et al., 1998).

Addendum to chelae diagnosis: Propodus longest at lower margin, fixed finger long, slender, slightly downturned with 4 denticles. Dactyl long, slender down turned with denticles and tuberculate on topmost surface. Articulation of dactyl to

48 palm of propodus rimmed and triangular. Palm nearly straight on lower margin, slightly convex on upper margin. Horizontal keel at midline of the palm, extending from upper margin of fixed finger to proximal edge of propodus. Keel with distinct row of tubercles, tubercles above keel forming two loosely defined rows. Propodus generally evenly tuberculate on outer margin, slight sulcus below uppermost margin smooth. Carpus oval, longest from proximal to distal margins of articulation with well defined, smooth sulcus at midline; evenly tuberculate on outer surface. Inner surface becoming smooth at margin of articulation.

Material examined.— See Table 3.

Occurrence.— Coon Creek Formation in Union and Pontotoc Counties,

Mississippi and McNairy County, Tennessee; Navarro Formation near Castroville,

Bexar County, Texas; Difunta Group in Coahuila State, Mexico; Portrerillos

Formation in San Luis Potosi State, Mexico. Late Campanian to early Maastrichtian

(Bishop et al., 1998).

Discussion.— Material forming the basis for interpretations include specimens collected by Krystyna Kornecki, Dr. Rodney Feldmann, Dr. Carrie

Schweitzer, AnnMarie Jones, Dr. Ovidiu Frantescu, Dr. Adina Frantescu, Dr. George

Phillips, and members of the Mississippi Gem and Mineral Society on March 24th and

25th, 2012 and December 17th and 18th, 2013, as well as those received on loan from the Pink Palace Museum, Memphis, Tennessee; the South Dakota School of Mines and Technology Museum, Rapid City, South Dakota; and the Mississippi Museum of

Natural Science, Jackson, Mississippi. All available specimens of Dakoticancridae were measured and plotted where data permitted (Fig. 13). Ninety specimens of males

49 and females were identified using pleon or sternum morphology. Of these, three specimens have gonopores preserved (specimens 37-686; 37-379; 51-82) none of which possessed intersex characters.

The proportion of females to males is not a 1:1 ratio but rather 28 females and

47 males- a 0.59:1 ratio (Fig. 14). Perhaps this is a relic of preservational bias or sampling bias, but that is unlikely as all material was collected that was found at the outcrop and there are multiple collecting events by many different people over a long time span. Furthermore, a preservational bias is unlikely between males and females due to their apparent similar preservation potential. Decapods tend to have less well calcified cuticle if their lifestyle is to swim, such as shrimp (Amato et al., 2008). This difference in cuticle density is not observed between sexes of the same species without adaptations for swimming. Therefore, the presence of a greater abundance of males than females could indicate that the community preserved is a life assemblage, but that the females had different behavioral or distributional patterns than the males, such as migrating further onshore while brooding their young (Hooper, 1986).

Though some specimens of Dakoticancer australis comprised of sterna only or carapace and sternum could be identified as male or female, the majority of them cannot offer data on other measurements of their morphology due to damage or incomplete preservation. Only six specimens of female Dakoticancer australis could be quantified for certain comparisons. Measurements of male and female D. australis are quantified in Figures 15, 16, and 17. Measurements of width vs. width/frontal orbital width and width vs. frontal width to frontal orbital width were of normal,

Gaussian distribution, but measurements of width vs. width to frontal orbital width

50 are Non-Gaussian (Table 4). When tested for similarity between males and females, all p-values indicated a rejection of the null hypothesis, concluding that sexual dimorphism is not present in frontal ratios of D. australis (Table 5).

In 1983, Bishop described a new species of hermit crab, Paguristes whitteni, based on “major” and “minor” claws of “male” and “female” specimens as well as carpus and manus specimens that possessed a distinct, tuberculate ornamentation. His description (Bishop, 1983, p. 422-423) embraces both articulated specimens of

Dakoticancer australis as well as disarticulated Paguristes whitteni. His description is as follows: “Merus triangular in cross section; 1.2 times longer than carpus, 1.8 times longer than wide with broad distal articulators separated from rest of well-developed distal ridge. Inner surface in plane of carpal articulators somewhat convex, tuberculate over entire surface; tubercles largest on upper angle. Upper face of merus more convex, coarsely tuberculate on angles with tubercles decreasing in size away from edges. Lower surface smooth to slightly undulatory except for tuberculate angles.

Carpus 1.4 times longer than wide, very convex above, concave below; bordered at both ends by articulating ridges; deep furrow running from upper meral articulator to midway between propodal articulators. Outer surface densely tuberculate; inner surface smooth on concave portion, tuberculate on convex portion.

Claws subequal, left usually larger, heterochelous. Major propodus ~ 1.5 times as long as high, upper and lower margins convex, carpal articulation strongly oblique, propodal articulation nearly vertical. Merus 1.4 times as long as high; fixed finger short and turned downward. Outer face strongly convex with shallow trough

51 running from just below upper carpal articulator to outer dactyl articulator; surface tuberculate. Tubercles largest and densest on lower part, randomly located except for those arranged in 3 poorly defined rows below shallow trough, then in dense random pattern below, onto lower surface and lower part of inner surface. Shallow trough mostly smooth, giving way above to crested upper surface overhanging inner face to a degree, especially distally. Below tuberculate crest inner face mostly smooth and flared into convexity by ridge running from middle of proximal edge upward to inner dactyl articulator. Articulating ridge present on inner and outer distal (propodal) edges. Fixed finger short, downturned, slightly grooved, and finely tuberculate with three teeth increasing in size distally. Only stub of dactylus preserved, but apparently stout and presumably short.

Minor claw relatively higher than major, outer face more uniformly convex and tuberculate with only shallow depression paralleling upper edge, forming straight ridge surmounted by coarser tubercles. Inner surface dominated by oblique ridge running from midpoint of proximal margin to dactylus articulator. Claw concave above this ridge and convex below it except for depression at base of finger.

Tubercles becoming coarser on lower part of outer face. Lower margin sharp, almost keel-like as upper margin,” (Bishop, 1983, p. 422-423).

Some noteworthy aspects of this description include the reiteration of the short, stout, and downturned dactyl, which is later clarified as being “only [the] stub of dactylus preserved, but apparently stout and presumably short,” (Bishop, 1983, p.

423). The diagnosis of Dakoticancer australis claws by Bishop et al. (1998) stated,

“chelae equal, short, stout, and crested, carpal articulation very oblique; fingers short,

52 downturned,” (p. 242 after Bishop, 1983b). Without illustration of the morphological differences that Bishop indicates, it is not possible to confirm the morphological differences suggested, utilizing available material of approximately 80 claws.

Bishop stated (1983, p. 420-421) that “the specimens representing this taxon

[Paguristes whitteni] are all claw elements” and that the “preservational bias is consistent with the pagurids because of the heavy mineralization of the claws,” referring to the absence of a carapace. He goes on to state that the claws “seemed to resemble claws of Dakoticancer australis; a similarity so striking that I concluded that the large “hermit crab claws” might be nothing more than claws of large specimens of D. australis. A search for D. australis carapaces with claws attached yielded only four specimens (out of 528) all amongst the largest of the carapaces.” In the numerous collections of Blue Springs Locality and Coon Creek Formation material, observed 14 articulated carapaces of D. australis (KSU 002, GAB 37-1461,

GAB 37-615, B-013, GAB 37-no number, GAB 37-740, GAB 37-381, GAB 37-532,

GAB 37-460, GAB 37-715, GAB 37-372, GAB 37-508, GAB 37-631, GAB 37-525), ranging within the intermediate size range; carapaces much smaller as well as much larger are also present in the collection, but lack articulated first pereiopods. The data that Bishop (1983, p. 424, fig. 5) illustrated suggests that he had 26 specimens of

Dakoticancer australis claws to compare to Paguristes whitteni claws, of which he documented measurements for 12 specimens. With only four articulated specimens, the author finds the assessment presented in this graph lacking. Specimens illustrated in fig. 3 (Bishop, 1983, p. 421) and fig. 4 (Bishop, 1983, p. 422) do not seem to illustrate morphological differences, but rather preservational bias in mode of

53 preservation and ornamentation. Furthermore, the data does not indicate which data points are of a left or right claw, though the plot of Paguristes whitteni does group into two distinct sizes and could be interpreted as heterochelous without more data.

Of course, these specimens cannot be associated with a single individual because they are disarticulated and the assessment of heterochelous claws may be a relic of preservational bias.

Bishop’s (1983) observation that the claws on articulated specimens of

Dakoticancer australis are quite small, and that the claws attributed to Paguristes whitteni are larger and disarticulated is indeed a preservational bias. To suggest that these morphologically identical forms of different sizes belong to two different species within two different infraorders, the Anomura and the Brachyura, yet occurring in the same locality, would indeed be a dramatic case of convergent evolution. Preservational bias caused by disarticulation is more prominent in specimens with larger claws, perhaps due to decomposition, predation, or transport.

The majority of carapaces lack articulation of the first pereiopods altogether. Thus,

Paguristes whitteni is herein considered to be a junior synonym of Dakoticancer australis.

The range of sizes in these claws is noteworthy; the increase in the size of the propodus correlates with the increase in size of ornamentation and may indicate allometric growth, though the small forms are identical to large forms in all other instances (Figure 18).

Also of note, Bishop illustrated paratype specimens of Dakoticancer australis

USNM no. 73840 (1983, p. 425, fig. 6A; Bishop et al., 1998, p. 243, fig. 1-6, 1-8), a

54 male specimen, and specimen USNM no. 18628 (1983, p. 425, fig. 6B; Bishop et al.,

1998, p. 243, fig. 1-7). These specimens possess noticeably different morphology.

Specimen USNM. No. 73840 is widest at about 50% of the carapace length, has a larger posterior shelf, is less tumid in the branchial regions and has a less well defined cardiac region. Specimen USNM. No. 73840 is narrowest at 50% the carapace length, has very tumid branchial regions, and a well defined cardiac region ending in a sharp point about 75% the carapace length. The specimens of the Coon Creek Formation and Blue Springs Locality conform morphologically to the type specimen illustrated by Rathbun (1935, p. 134-135, pl. 10, fig. 20) but do not resemble the paratypes illustrated by Bishop. The paratypes of Dakoticancer australis should be re-examined to determine if they represent a different species.

Genus TETRACARCINUS Weller, 1905

Type species. –Tetracarcinus subquadratus Weller, 1905

Included species.— Tetracarcinus subquadratus Weller, 1905.

Generic diagnosis.— “Carapace generally small, subquadrate, length nearly equal to width, widest at position of epibranchial regions; orbits rimmed; lateral margins sinuous; posterior margin rimmed; cervical groove shallow medially and poorly developed distally; regions flattened, weakly inflated; epibranchial and metabranchial regions separated by broud depression enclosing narrow mesobranchial region; epibranchial regions transversely weakly inflated; cardiac region with posterior tubercle; first pereiopods isochelous,” Feldmann et al., 2013, p. 28).

TETRACARCINUS SUBQUADRATUS Weller, 1905

Figures 12.1, 12.2

1905. Tetracarcinus subquadratus, Weller, Journal of Geology, 13: 324-337.

1907. Tetracarcinus subquadratus, Weller, Geol. Survey of New Jersey, Paleontological Survey IV:

846-853.

1935. Tetracarcinus subquadratus, Rathbun, Geological Society of America Special Papers, 2: p. 41, pl. 10, fig. 16-17.

1958. Tetracarcinus subquadratus. Holland, Jr., and Cvancara, Journal of Paleontology, 32:3: pg. 496.

1991. Tetracarcinus subquadratus. Mississippi Geology, 12 (1/2): pg. 13.

1998. Tetracarcinus subquadratus, Bishop et al., Contributions to Zoology, 67 (4): p. 244, fig. 2.1.

2010. Tetracarcinus subquadratus, Schweitzer et al., Crustaceana Monographs, 10: p. 58.

2013. Tetracarcinus subquadratus, Feldmann et al., Bulletin of the Mizunami Fossil Museum, 39: 28.

Diagnosis.— As for the genus.

Material examined.— See Table 6.

Occurrence.— Tetracarcinus subquadratus occurs at the Blue Springs

Locality (GAB 37), Coon Creek Formation, New Albany, Mississippi. The species is also known from the Magothy Formation at Cliffwood, New Jersey; Woodbury

Formation at Lorillard; Merchantville Formation on the Delmarva Peninsula; as well as from one occurrence in the Lewis Shale of Wyoming. It ranges from the late

Santonian to the Maastrichtian (Bishop et al., 1998).

Discussion.— The diagnosis of Tetracarcinus subquadratus by Bishop et al.

(1998) does not include any sternal morphology. It is also of note that Bishop et al.

(1998) illustrated in Figure 2.2 (p. 245) a claw of T. subquadratus, unarticulated.

Catalogue numbers are not given, thus making it impossible to determine if these specimens were in fact closely associated, though both are recognized as being from the Coon Creek Formation, Mississippi. The diagnosis described this claw as “small; palm longer than high, smooth except for granulate upper surface,” (Bishop et al.

1998, pg. 244). I have not observed any articulated specimens of T. subquadratus, but

56 the specimen of claw illustrated by Bishop et al. (1998) does not seem to fit the description, though abbreviated, as “small” as it is nearly the same width as the typical carapace width of the species. Weller (1905a, 1905b, 1907) illustrated carapaces only of the species, and Rathbun (1935) illustrated carapaces of the species and one, disarticulated claw specimen that she assigned to the species as well.

Roberts (in Richards, 1962) also illustrated T. subquadratus without chelae, articulated or disarticulated. He also noted the specimens reported as T. subquadratus by Rathbun (1935) of a palm and carapace were not definitively assignable to the species. The specimen of claw illustrated by Rathbun (1935, p. 134-135, plate 10, fig.

18) does not have any portion of the dactyl preserved and the palm is much shorter than the claw illustrated by Bishop et al. (1998). The claw specimen (Rathbun, 1935, p. 134-135, plate 10, fig. 18) has no catalogue number but is referenced to be in the

New Jersey State Museum and occurred in the Tinton beds, Beers Hill, New Jersey.

Her other illustrated specimens are from Wyoming and the Cliffwood Clay of

Cliffwood Point, New Jersey. This claw is not considered at this time to be that of

Tetracarcinus subquadratus. There is insufficient evidence to assign it to a new species or any other species at present. The diagnosis by Feldmann et al. (2013) also included a reference to chelae, “first pereiopods isochelous,” (p. 28). The specimen illustrated in figure 13.2, NJSM 23337, of a “crushed dorsal carapace” may have these pereiopods preserved, but there is also no specific reference to a specimen with preserved pereiopods or chelae.

Specimens of Tetracarcinus subquadratus do not have sterna articulated with carapaces frequently and, therefore, do not provide evidence to correlate carapace

57 morphology with sex. Fewer specimens of T. subquadratus are available for study than are of Dakoticancer australis. Because the specimens are smaller, this proportion could be related to collecting bias.

Measurements of Tetracarcinus subquadratus yielded fairly well constrained data with one clear outlier. The ratio of the frontal width to the fronto-orbital width of

Tetracarcinus subquadratus is negatively correlated (Figure 19). In all specimens of

Dakoticancer australis, these data barely trend with a negative slope and are poorly constrained. The relationship of total width to frontal width in Tetracarcinus subquadratus is more positively trending with size than that observed in

Dakoticancer australis. The ratio of body width to fronto-orbital width in

Tetracarcinus subquadratus also trends positively, as is illustrated in Dakoticancer australis.

Genus AVITELMESSUS Rathbun, 1923

Type species. –Avitelmessus grapsoideus Rathbun, 1923.

Included species.— Avitelmessus grapsoideus Rathbun, 1923.

Generic diagnosis.— “Carapace very large, circular with concave front, widest at midpoint; arched longitudinally, fairly level transversely. Rostrum small, narrow, medially grooved. Orbits with two fossae forming two concavities in plan view. Carapace margin nearly forms circle from lateral orbital spines, spinose, carapace grooves poorly developed except for gastro-cardiac, proximal cervical

(except obsolete gastric portion), and epimesobranchial [sic]. Sagittal ridge well delineated; cephalic arch poorly differentiated except for mesogastric; scapular arch with well-delineated cardiac and branchial regions, branchial regions differentiated

58 into epibranchial and mesobranchial by epibranchial groove, posterior of branchial separated into mesobranchial and metabranchial by oblique ridges running outward and backward. Epibranchial posterior and anterior of mesobranchial with subparallel ridges. Subtle ridges parallel gastrocardiac groove along outer edges from epibranchials to orbits. Chelae equal, large, triangular in cross section, spiny; palm slightly longer than high, oblique; fingers long, slightly downturned, toothed,”

(Bishop et al., 1998, p. 244).

AVITELMESSUS GRAPSOIDEUS Rathbun, 1923

1923. Avitelmessus grapsoideus. Rathbun, North Carolina geol. Survey, 5, 403-407.

1927. Avitelmessus grapsoideus. Rathbun in Wade, Geological Survey of Tennessee, Professional

Paper 137: p. 190, pl. LXIX figs. 1-7, LXX figs. 1-12.

1957. Avitelmessus grapsoideus. Kesling and Reimann, Contributions from the Museum of

Paleontology, XIV (1): 1-15.

1991. Avitelmessus grapsoideus. Mississippi Geology, 12 (1/2): pg. 13.

1998. Avitelmessus grapsoideus. Bishop et al., Contributions to Zoology, 67(4): p. 249 figs. 2.1-2.5.

Diagnosis.— As for the genus.

Material examined.— Avitelmessus grapsoideus is not available from collected material at the Blue Springs locality, but specimens of the Coon Creek

Formation werre observed at the U.S. National Museum of Natural History.

Occurrence.— Peedee Formation of North Carolina as well as from Georgia and Alabama and the Mississippi Embayment into Texas, ranging from Campanian to early Maastrichtian (Bishop et al., 1998).

Discussion.— Rathbun’s description of Avitelmessus grapsoideus (1923, p.

403-404) and that by Kesling and Reimann (1957, p. 4-11) should be referenced for a complete assessment of the species.

Family IBERICANCRIDAE Artal et al., 2008

Included genera.—Ibericancer Artal et al., 2008; Seorsus Bishop, 1988;

Sodakus Bishop, 1978.

Family diagnosis.—“Carapace subrectangular, about as long as wide, generally widest just under half the distance posteriorly but may be at position two- thirds the distance; rostrum narrow, downturned, bilobed or quadrilobed; orbits square, directed forward, fronto-orbital width ranging from about 40-70% maximum width but usually about half; branchiocardiac groove deep, cervical groove discontinuous; axial regions well defined and distinct; sternum narrow, deep sterno- pleonal cavity, sternite five with pleonal locking mechanism, sterna sutures 4/5 through 7/8 interupted; female gonopore on coxa of pereiopod 3, male gonopore on coxa of pereiopod 5, spermatheca of female at sterna suture 7/8; male pleon very narrow, all somites free, female pleon wider, all somites free; pereiopods 4 and 5 apparently subdorsal, 5 reduced in size,” (Feldmann et al., 2013, p. 29).

Genus SEORSUS Bishop, 1988

Type species. –Seorsus wadei Bishop, 1988.

Included species.— Seorsus wadei Bishop, 1988; Seorsus kauffmani

Feldmann et al., 2013; Seorsus millerae (Bishop, 1992a).

Generic diagnosis.— “Carapace slightly longer than wide, width about

93% maximum carapace width, width at position of single anterolateral spine about

40-50% the distance posteriorly on carapace; rostrum long, with four blunt spines including inner-orbital spines; orbits square, rimmed, with intraorbital spine, fronto- orbital width ranging from half to 70% maximum carapace width; well-defined branchiocardiac groove and moderately defined cervical groove, and well-defined axial regions; sternum narrow, sterno-pleonal cavity narrow, sternites 1-3 fused, sternite 4 long, with concave lateral margins, sterna suture 4/5 incomplete; male pleon with all somites free, subdorsal pereiopods 4 and 5; major chela granular; propodus bulbous, two nodes at articulation with carpus; carpus granulated with X-shaped groove; fixed finger and dactylus thin, delicate,” (Feldmann et al., 2013, p. 29).

SEORSUS WADEI Bishop, 1988

1988. Seorsus wadei, Bishop. Proc. Biol. Soc. Wash., 101(1): p. 72, fig.1A-F.

1991. Seorsus wadei, Bishop. Mississippi Geology, 12 (1/2): p.13.

1998. Seorsus wadei. Bishop et al., Contributions to Zoology, 67(4): p. 249 figs. 2.1-2.5.

2008. Seorsus wadei. Artal, Guinot, Van Bakel, and Castillo, Zootaxa, 1907:1-27.

Diagnosis.— “Carapace of moderate size, longer than wide (L/W=1.10), widest at anterior 1/3; lateral margins distinctively convergent posteriorly; grooves broad, moderately defined; aerolations very tumid, especially epibranchial lobes; cardiac region with small central tubercle; metabranchial region with subtle transverse and submarginal ridges. Claws and legs unknown,” (Bishop et al., 1998, p.

245).

Material examined.— Seorsus wadei was not available for observation in the study material. Type species Seorsus wadei holotype GSUM 1693, and paratype

GSUM 1694 (Bishop, 1988), were collected from the Coon Creek Formation in

Union County, Mississippi from the NE1/4, NW1/4, SE1/4, section 9, T. 8S., R. 4E

(Blue Springs Locality).

Occurrence.— GAB 37, Coon Creek Formation, Blue Springs Locality, New

Albany, Mississippi. Maastrichtian (Bishop et al., 1998).

Discussion.— Bishop et al. (1998) cited two known specimens of Seorsus wadei. With the amount of available material, it is of interest that S. wadei has not occurred more frequently. This species was not numerically abundant within the

Mississippi Embayment fauna. Seorsus kauffmani occurs in the Turonian Mancos

Shale, Semilla Sandstone Member, Sandoval County, New Mexico, and Seorsus millerae from spoils piles of the early Campanian Merchantville Formation in

Newcastle County, Delaware (Feldmann et al., 2013).

Section HOMOLOIDA de Haan, 1839

Superfamily HOMOLOIDEA De Haan, 1839

Family HOMOLIDAE De Hann, 1839

Genus LATHETICOCARCINUS Bishop, 1988b

Type species.—Latheticocarcinus shapiroi Bishop, 1988b, by original designation.

Included species.—Latheticocarcinus adelphinus (Collins and Rasmussen, 1992);

Latheticocarcinus affinis (Jakobsen and Collins, 1997); Latheticocarcinus atlanticus

(Roberts, 1962); Latheticocarcinus brevis (Collins, Kanie, and Karasawa, 1993);

Latheticocarcinus brightoni (Wright and Collins, 1972); Latheticocarcinus centurialis

(Bishop, 1992b); Latheticocarcinus declinatus (Collins,Fraaye, and Jagt, 1995);

Latheticocarcinus dispar (Roberts, 1962); Latheticocarcinus ludvigseni Schweitzer,

Nyborg, Feldmann, and Ross, 2004; Latheticocarcinus pikeae (Bishop and Brannen,

1992); Latheticocarcinus punctatus (Rathbun, 1917a); Latheticocarcinus schlueteri

(Beurlen, 1928b); Latheticocarcinus shapiroi Bishop, 1988b; Latheticocarcinus spinigus

(Jakobsen and Collins, 1997); Latheticocarcinus transiens (Segerberg, 1900).

Diagnosis.—“Carapace longer than wide (width measured between lineae homolicae), typically widest just posterior to intersection of cervical groove and linea homolicae but relatively uniformly wide through entire length, surface granular, ornamentd with discrete, large tubercles; rostrum bifid or singular, sulcate; often with small pseudorostral spines; usually with supraorbital spine; protogastric, hepatic, mesogastric and cardiac regions ornamented with large tubercles; grooves defining lateral margins of mesogastric region deeply incised; cervical groove very deeply incised, arcuate, U-shaped, not typically sinuous, separating the carapace into distinctive anterior and posterior portions; branchiocardiac groove very deep anteriorly, beginning about midway between linea homolicae and axis, extending axially, curving around and extending laterally to intersect linea homolicae; lineae homolicae very well-developed, sub-hepatic and sub-branchial regions rarely preserved; cardiac region with two swellings positioned one beside the other, sometimes with lateral ridges extending onto cardiac regions; cardiac region not well differentiated from branchial regions by deep grooves,”

(Feldmann et al., 2013, p. 17).

? LATHETICOCARCINUS ATLANTICUS (Roberts, 1962)

Figure 20.5

1962. Homolopsis atlantica. Roberts, Bulletin of the New Jersey Division of Geology, 61: p. 179, pl. 89, fig. 4.

2004. Latheticocarcinus s atlanticus (Roberts, 1962). Schweitzer et al., Journal of Paleontology, 78: p. 136

2010. Lateticocarcinus atlanticus. Schweitzer et al., Crustaceana Monographs, 10: p. 68.

Diagnosis.—“Carapace including extralineal flanks; mesobranchial region with one large tubercle posteriorly and two obliquely ovate ones; cardiac region well-defined, with three large tubercles; tubercles ornamenting all regions very large; subepibranchial spine very large and long,” (Feldmann et al., 2013, p. 17).

Description.—Carapace fragmented; anterior margin obscured, posterior and posterior-lateral margin absent. Rostrum sulcate, appearing singular. Mesogastric region weakly defined by grooves with axial node anteriormost with slightly larger nodes defining protogastric region; pair of nodes lateral to protogastric nodes form hepatic region. Cervical groove well defined; mesogastric region tumid with deepening margin toward the gastric region. Gastric region most tumid with tubercles ending at a small spine. Lateral flanks vertical.

Material examined.— GAB 4-1352, length: 13.18 mm, width: 17.86 mm.

Occurrence.— ? Coon Creek Formation, Blue Springs Locality, Union County,

Mississippi, Maastrichtian. Marshalltown/Merchantville spoil pile on north side of the C and D Canal in Delaware (Feldmann et al., 2013); Roberts (1962) reported the species from the Merchantville Formation in New Jersey.

Discussion.— The specimen does not possess the Coon Creek Locality number assigned by Bishop and is therefore tentatively described in the assessment of the

64 decapod fauna. The preservation is also fragmentary. Because of this, the shape of the carapace is unknown as is the anterior and posterior margin morphology. This specimen was likely a molt: the subhepatic and subbranchial regions are not preserved, suggesting that the specimen has broken along the linea homolica (sensu Collins, 1997). The presence of this feature is also a defining character of the Homolidae. Though the material is quite fragmentary, the cardiac swellings situated beside each other and the distinct separation of the anterior and posterior of the carapace by the cervical groove suggests placement within Latheticocarcinus. Latheticocarcinus atlanticus also occurs on the Atlantic Coast. Thus, if the identification is confirmed as Latheticocarcinus atlanticus in the Coon Creek Formation, the range of this species can be extended to the Mississippi

Embayment.

Section RANINOIDA Ahyong et al., 2007

Superfamily PALAEOCORYSTOIDEA Lőrenthey in Lőrenthey and Beurlen, 1929

Included families.— Palaeocorystidae Lőrenthey in Lőrenthey and Beurlen, 1929.

Diagnosis.— As for family.

Discussion.— Karasawa et al. (2014) conducted a phylogenetic analysis of the

Raninoida. This classification scheme is followed herein.

Family PALAEOCORYSTIDAE Lőrenthey in Lőrenthey and Beurlen, 1929

Included genera.— Alessandranina Karasawa et al., 2014; Cenocorystes

Collins and Breton, 2009; Cretacoranina Mertin, 1941; Eucorystes Bell, 1863;

Ferroranina van Bakel et al., 2012; Joeranina van Bakel, 2012; Notopocorystes

M’Coy, 1849.

Diagnosis.— “Carapace obovate, usually longer than wide, widest at position of third or fourth anterolateral spine; frontal margin wide; anterolateral margin with 2 to 4 spines; carapace surface ornamented with ridges, straps, tubercles or unornamented; epibranchial region with weak tooth; fronto-orbital margin and orbits wide, orbital margin with two fissures; rostrum generally long, with two spines at tip; orbits with inner, intra- and outer orbital spines, some of which may be bifid; gymnopleuran condition absent; sternites 1-3 fused, 1 and 2 directed downward; sternite 4 long, pereiopod 1 articulating near posterior corner, moderately wide, lateral margins concave; sterna suture 4/5 sinuous laterally, then turning abruptly anteriorly parallel to axis; episternite 4 usually laterally directed but may be posteriorly directed (Notopocorystes); sternites 4/5 usually in broad contact (except

Notopocorystes); sternite 5 long, moderately wide, with double peg structure on episternal projection for attachment of pleon, episternites directed laterally or posterolaterally (Notopocorystes); sterna suture 5/6 complete; all female pleonites free, pleonite 6 long, pleonites 2-5 with central spine, entire pleon reaching to level of base of coxae of first pereiopods; male pleon narrower, telson triangular, somite 6 long, reaching to level of base of coxae of pereiopods 2; pleonal holding mechanism consisting of a double-peg structure; longitudinal furrow on merus of maxilliped 3; chelae with long fingers; female gonopore coxal, small, round; spermatheca at end of suture 7/8, separated from one another; pereiopod 5 reduced in size,” (Karasawa et al., 2014, p. 239).

Discussion.— The Palaeocorystidae range from the Early to Late Cretaceous.

The family was revised by van Bakel et al. (2012) and later by Karasawa et al.

(2014).

Genus CRETACORANINA Mertin, 1941

Type species. – Raninella? schloenbachi Schlüter, 1879.

Included species.— Cretacoranina denisae (Secretan, 1964); C. fritschi

(Glaessner, 1929); C. tesactea (Rathbun, 1926a); C. trechmanni (Withers, 1927).

Generic diagnosis.— “Carapace obovate, wide in anterolateral third; posterolateral margin narrowing considerably; rostrum extending beyond orbital margin, tip bifid, with spine just posterior to each spine at tip for a total of four, short spines on either side of rostrum forming inner orbital spines, rostrum sometimes rimmed; intra-orbital spine bifid, bounded by fissures; outer orbital spine bifid; anterolateral margin with 3 or 4 spines; spines becoming smaller posteriorly, last one nearly obsolete; cervical groove absent; branchiocardiac groove developed as arcs on either side of axis; dorsal carapace ornamentation developed as fungiform pillars overall but ending in indistinct scalloped termination to base of orbital margin or just barely onto orbital margin and anterolateral spines; sternite 3 appearing triangular; sternite 4 wide, episternite 4 wide, in broad contact with sternite 5, directed laterally, suture 4/5 incomplete, straight; sternite 5 narrower than sternite 4,” (Karasawa et al.,

2014, p. 239).

Discussion.— Van Bakel et al. (2012) limited the genus, though still quite morphologically diverse, to those species with hexagonal ornamentation overall but terminating at the base of the anterolateral spines and those lacking a cervical groove

(Feldmann et al. 2014, p. 240). It is not clear that the small pillars forming the texture of the carapace are hexagonal in the Mississippi specimens. The pillar structures do appear to have individual setal pits and to become smaller and closer together at the margins of the carapace. Specimen GAB 37-1143 illustrates exocuticle on the right lateral margin of the dorsal carapace, indicating that these pillars are less distinct when the external layer of cuticle is present, though exocuticle is not commonly observed.

CRETACORANINA TESTACEA (Rathbun, 1926a)

Figures 20.3 – 20.6

1926a. Raninella testacea. Rathbun, in Wade, ed., U.S. Geological Survey Professional Paper, 137: p.

190.

1929. Raninella testacea Rathbun, 1926. Glaessner, in F.J. Pompeckj, ed., Fossilium catalogus, 1,

Animalium, 41: p. 370.

1935a. Raninella testacea. Rathbun, Geological Society of America, Special Paper, 2: p. 50.

1983. Notopocorystes testacea (Rathbun, 1926). Journal of Crustacean Biology, 3(3): 419.

1991. Raninella testacea. Bishop, Mississippi Geology, 12(1-2): p. 12.

1998. Cretacoranina testacea (Rathbun, 1926). Tucker, Proceedings of the Biological Society of

Washington, 111: fig. 5, table 4.

2009. Cretocoranina testacea. Waugh et al., Bulletin of the Mizunami Fossil Museum, 35: p. 20.

2010. Raninella testacea. Schweitzer et al., Neues Jahrbuch fur Geologie und Palaontologie,

Abhandlungen, 256(3): p. 75.

2012. Cretocoranina testacea. van Bakel et al., Zootaxa, 3215: p. 19.

2013. Cretocoranina testacea. Feldmann et al., Bulletin of the Mizunami Fossil Museum, 39: p. 25.

Diagnosis.—“Carapace longer than wide, widest about one third the distance posteriorly at position of last anterolateral spine; moderately vaulted transversely and

68 longitudinally. Anterolateral margins convex, with four spines excluding bifid outerorbital spine, shorter than concave posterolateral margins. Outerorbital spines not extending as far anteriorly as bifid intra-orbital spines, intra-orbital spines separated from outerorbital spines and rostrum by open fissures; rostrum triangular, base composed of inner orbital spines, axial spines with axial groove. Surface covered with subhexagonal granules, visible to naked eye. Crescentic furrows at middle of carapace define inner limit of branchial region,” (Feldmann et al., 2013, p. 25).

Description.— Carapace obovate, widest at anterolateral 1/3, narrowing posteriorly; bifid rostrum with deep sulcus at axis, arcing laterally to small, inner orbital spine. Two sets of anterolateral spines on either side of rostrum, first of which possesses three short spines, the second two; followed by four anteriorly directed spines on lateral margin, extending half way to the posterior and ending before the carapace narrows, becoming smaller posteriorly. Orbits oval, laterally elongate extending nearly to lateral margin of carapace, vertically compressed. Dorsal carapace evenly covered with fungiform pillars ending abruptly at the proximal margin of the anterolateral spines. Branchiocardiac grooves weakly depressed arcs on either side of axis. Anterior suture of dorsal carapace and venter distinctly grooved, gently arching from lateral margin of rostrum below the slight shelf formed by anterior spines to the most posterior lateral spine; two rows of small spines between anterior spines and suture; second sinuous groove extending parallel to first and more posterior; separated by slightly domed ridge. Some sections of sternites visible.

Sternite 3/4 fused. Sternite four widest at contact with coxae of first pereiopods; forms widest portion of sternum. Cuticle of sternum also with smaller, fungiform

69 pillars. Palm of chelae with anteriorly directed spines on ventral and dorsal margin; laterally compressed and much narrower than tall; dactyl about equal in length to anterior margin of palm. Outer face of palm smooth. Fixed finger short, slightly downturned. Segments of first pereiopods with rows of anteriorly directed spines.

Material examined.— See Table 7.

Occurrence.— Late Cretaceous, Campanian-Maastrichtian. Coon Creek

Formation, Tennessee, Blue Springs locality; Coon Creek Formation, Mississippi;

Navesink Formation, new Jersey; Marshalltown/Merchantville spoils of the north side of the C and D Canal, Delaware (Feldmann et al., 2013).

Discussion.— Thirty specimens of Cretocoranina testacea were observed and measured from the Blue Springs Locality of the Coon Creek Formation. Of these 30 specimens, one specimen was preserved with a swelling in the left lateral dorsal carapace in the branchial region, a deformation attributed to infestation by an isopod

(Klömpmaker, 2014). This isopod related swelling has been reported in many decapods and has been observed previously in C. testacea (Figure 20.5)

(Klompmaker, 2014).

Specimens vary in preservation but are generally excellent, some of which have articulated claws and ventral portions of the carapace (such as sternites) and some show cuticle texture. Feldmann et al. (2013) noted that the type specimen and the material from New Jersey were incomplete, with the type specimen lacking a rostrum and the posterior portion of the carapace. The Coon Creek Formation specimens from Mississippi present the most complete examples of the species.

Superfamily RANINOIDEA De Hann, 1839

Included families.— Lyreididae Guinot, 1993; Raninidae De Hann, 1839.

Diagnosis.—“Carapace longer than wide or about as wide as long, generally ovate, usually vaulted transversely, regions poorly defined; usually with well developed rostrum and orbital spines; anterolateral margins usually with one spine but can have none or more than one; posterolateral margin lacking spines; cervical groove not reaching ventral edge of carapace; branchiocardiac groove indistinct, developed as boundary of urogastric region; branchial ridges absent; junction between sternum and pterygostome usually wide; coxa of third maxilliped small, flattened; sternum narrow, sternites 1-3 generally fused, sternites 5 and 6 with lateral extensions, sterna suture 6/7 complete, sternites 7 and 8 often reduced and at lower level than other sternites; where known, pleon narrow in males and females, showing reduced but clear dimorphism, never reaching sternite 4; genital openings coxal, spermatheca present on endosternite 7/8; branchiostegite reduced; gymnopleuran condition present,” (Karasawa et al., 2014, p. 243).

Family LYREIDIDAE Guinot, 1993

Included Subfamilies.—Bicornisranininae Karasawa et al., 2014; Lyreidinae

Guinot, 1993; Macroacaeninae Karasawa et al., 2014; Marylyreidinae van Bakel,

Guinot, Artal, Fraaije, and Jagt, 2012; Rogueinae Karasawa et al., 2014.

Diagnosis.—“Carapace much longer than wide, oblanceolate; dorsal surface smooth or punctate, regions undefined; anterior margin narrow or wide; rostrum trifid, middle spine generally much longer than other two which serve as inner orbital spines; orbit with intra- and outer orbital spines; anterolateral margins may be entire or with one or two spines; sternum-pterygostome junction poorly to well-developed

71 or absent; gymnopleuran condition present; sternites 1-3 fused, forming a cap-like shape; sternite four large, with lateral extension anteriorly, concave laterally; sternite

5 of similar shape but smaller, with lyreidid hook (Guinot, 1979) and double peg pleonal locking mechanism; sternite 6 much smaller, sometimes with ridge; sternites

7 and 8 much reduced in size, median one reaching sternite 7; pleon narrow in both males and females, telson short, somite 6 long; pleon sexually dimorphic, somites 2 and 5 proportionally wider in females than males, somite 6 proportionally longer in females than males; spermatheca placed on sternite 7, separated by wall; merus of maxilliped three longer than ischium,” (Karasawa et al., 2014, p. 243-244).

Subfamily MARYLYREIDINAE Van Bakel, Guinot, Artal, Fraaije, and Jagt, 2012

Included genera.—Bournelyreidus Van Bakel, Guinot, Artal, Fraaije, and Jagt,

2012; Hemioon Bell, 1862; Heus Bishop and Williams, 2000; Marylyreidus Van

Bakel, Guinot, Artal, Fraaije, and Jagt, 2012.

Diagnosis.—“Carapace much longer than wide, oblanceolate, widest at about mid-length; dorsal surface smooth, regions undefined, ornamented with upright nodes or fungiform nodes; fronto-orbital margin wide, ranging from about two-thirds to

80% maximum carapace width; rostrum trifid, middle spine generally much longer than other two serving as inner orbital spines, middle spine bifid; orbit with intra- and outer orbital spines; anterolateral margins with one or two spines; sternum- pterygostome junction absent; gymno-pleuran condition present; sternites 1-3 fused, forming a cap-like shape; sternite 4 narrow, with blunt-triangular episternite directed laterally; sternite 5 with very large lateral projections, arcuate, extending as far as lateral margin of carapace, with lyreidid hook (Guinot, 1979) and double peg pleonal

72 locking mechanism composed of short pegs; sternite 6 much smaller, sometimes with ridge; sternites 7 and 8 much reduced in size; pleon narrow in both males and females, telson short, somite 6 long; spermatheca placed on sternite 7, separated by wall; merus of maxilliped 3 longer than ischium,” (Karasawa et al., 2014, p. 254).

Range.—Early Cretaceous (Albian) - Miocene (Karasawa et al., 2014, p. 254).

BOURNELYREIDUS Van Bakel, Guinot, Artal, Fraaije, and Jagt, 2012

1992. Hemioon eysunesensis. Collins and Rasmussen, Grønlands Geologiske

Undersogelse, Bulletin 162: p. 19.

Type species.— Hemioon eysunesensis Collins and Rasmussen, 1992.

Included species.—Bournelyreidus carlilensis (Feldmann and Maxey, 1980);

Bournelyreidus laevis (Schlüter in Von der Marck and Schlüter, 1868);

Bournelyreidus oaheensis (Bishop, 1988d); Bournelyreidus n. sp.

Diagnosis.—“Carapace much longer than wide, oblanceolate, widest at about mid-length; dorsal surface smooth, regions undefined; fronto-orbital margin wide, ranging about half to two-thirds maximum carapace width; rostrum overall trifid, middle spine generally much longer than other two which serve as inner orbital spines, central spine bifid at tip; orbit with intra- and outer orbital spines; anterolateral margins with two spines; sternum-pterygostome junction absent, sternite 4 articulating with third maxilliped; sternites 1-3 fused, forming a cap-like shape; sternite four narrow, with short arcuate projections anteriorly, concave laterally, with blunt triangular episternites directed laterally; sternite 5 with moderate lateral projections, arcuate; chelae flattened, lower margin with spines,” (Karasawa et al.,

2014, p. 254).

Discussion.—Bournelyreidus and Macroacaena as described by Van Bakel et al. (2012) and maintained by Karasawa et al. (2014) have very subtle differences; those of the dorsal carapace that are most apparent are the anterolateral spines and anterior margin. Bournelyreidus has two anterolateral spines and Macroacaena has one spine and may have small protuberance between the anterolateral spine and the outer orbital spine, as well as longer spines with an anterior margin that is flared anterolaterally. The ventral morphology of these genera is difficult to compare; the type material of all species of Bournelyreidus lack preservation of the venter entirely.

Macroacaena is limited to, “those species with a fronto-orbital width of about 50% maximum carapace width; a carapace reaching maximum width at about the position of the last anterolateral spine and maintaining that width to about 50 or 60% the distance posteriorly on carapace; a fourth sternite with small projections anteriorly; and a fifth sternite with wide projections anteriorly,” (Karasawa et al., 2014, p. 246).

Bournelyreidus has “sternite four narrow, with short arcuate projections anteriorly, concave laterally, with blunt-triangular episternites directed laterally; sternite 5 with moderate lateral projections, arcuate,” (Karasawa et al., 2014, p. 254). As can be seen, the preserved sternal morphology that is diagnostic of each genus is subtle. That said, sternite 1-3 is crown-like in Macroacaena and cap-like in Bournelyreidus;

Bournelyreidus has two anterolateral spines and Macroacaena may have nodes between anterolateral spines, much longer spines, and a flared frontal margin;

Macroacaena has a longitudinal keel on the dorsal carapace (sometimes) and

Bournelyreidus does not. The dorsal surface is smooth in Bournelyreidus and coarsely punctate in Macroacaena.

BOURNELYREIDUS n. sp.

Figures 20.3, 20.9, 20.16, 20.17

1991. Raninella tridens. Bishop, Mississippi Geology, 12(1-2): p. 13.

2012. Bournelyreidus tridens (Roberts, 1962). Van Bakel et al., Zootaxa, Monograph 3215, p. 118, fig.

38C, 38D (part).

Description.— Carapace small, narrow. Widest medially and nearly equal in width at anterior and posterior margin. Anterior outer-orbital spines forming continuous shelf to rostrum. Rostrum not preserved. Length of anterior spines unknown. Two pairs of small anterolateral spines directed anteriorly; carapace widest at most posterior anterolateral spine. Cuticle smooth to finely punctate. Dorsal carapace axially inflated with weak longitudinal keel; Branchiocardiac grooves bordering cardiac region as weakly depressed arcs on either side of axis. Sternite 1-3 fused, cordate. Sternite four long, narrow; sternite five wide with anterolateral to anteriorly projected and rounded episternites; episternite five with posterior projected hook-like, convex inward processes.

Material examined.— MPPM 1972.46.157, length: 20.64 mm, width: 14.16 mm; GAB 37-834, length: 18.95 mm; GAB 37-835, length: 12.66 mm, width: 9.5 mm; GAB 37-833 (specimen and cast), length: 12.84 mm, width: 7.7 mm; GAB

37-1071, length: 14.22 mm, width: 8.36 mm; GAB 37-832 (specimen and cast), length: 12.86 mm, width: 8.42 mm; KSU 6060, length: 15.56 mm, width: 18.42 mm;

KSU 6061, length: 13.79 mm, width: 16 mm; KSU 6062, length: 13.94 mm, width:7.72 mm; KSU 5101, length: 15.09 mm, width: 9.86 mm, frontal width: 4.56 mm; MMNS-IP 3140, 3 specimens: MMNS-IP 3140a, width: 8.32 mm, MMNS-IP

3140b, length: 11.2 mm, width: 7.1 mm, MMNS-IP 3140c, length: 14.56 mm, width:

8.66 mm.

Occurrence.— GAB 37, Blue Springs Locality, Coon Creek Formation, New

Albany, Mississippi, Maastrichtian.

Discussion.— Raninella tridens was originally described by Roberts, 1967, from a single specimen from New Jersey that lacked sternal morphology and lacked details of the anterior margin. Bishop (1991) later referred specimens from

Mississippi to this species as well. The species was later assigned to Bournelyreidus tridens by Van Bakel et al. (2012), who referenced the type material of R. tridens in the diagnosis, as well as referred to specimens from Mississippi. Morphology of the sternum of the Mississippi specimens was included in the diagnosis Bournelyreidus tridens, though the sternal morphology was not available on the type specimen of

Bournelyreidus tridens. Karasawa et al. (2014, p. 246) assigned the type specimen to

Macroacaena tridens (Roberts, 1962) as a new combination, which was considered

Bournelyreidus tridens by Van Bakel et al. (2012) and Feldmann et al. (2013).

Karasawa et al. (2014) did not include the Mississippi material in this diagnosis and suggested that the placement of this material needed to be assessed and was beyond the scope of their study at the time.

Karasawa et al. (2014) expanded on their explanation, saying that this assignment was Macroacaena tridens sensu Roberts (1962) and that Macroacaena is restricted to those species with fronto-orbital widths of about half the maximum carapace width, that the maximum carapace width is at about the last anterolateral spine, and that the width is maintained about 50 or 60% of the way down the carapace

76 posteriorly, as well as having sternite 4 with small projection anteriorly and sternite five with wide projections anteriorly. They noted that specimens from Mississippi possessed a hook-like structure on sternite 5, and had an articulation of sternites 4 and

5 similar to that of Lyreidinae and not Marylyreidinae. Specimens observed herein also have a hook-like structure on sternite five extending posteriorly to six, but sternite five is widest and extends laterally, and sternite 4 is quite narrow, as is typical of Lyreidinae and Marylyreidinae.

The assessment by Karasawa et al. (2014, p. 248-249) is puzzling: though the type specimen of Macroacaena tridens sensu Roberts (1967) from New Jersey has not been studied herein. The casts referred to by Karasawa et al. (2014, p. 248) of

Mississippi material have been observed and are of specimens GAB 37-832 and GAB

37-833. Examination of illustrations by Karasawa et al. (2014), the casts of

Mississippi material, and Van Bakel et al. (2012) are not congruent with statements of

Karasawa et al. (2014, p. 248-249) and generic diagnosis of Karasawa et al. (2014, p.

246, p. 254), nor with specimens of Mississippi material at hand. Karasawa et al.

(2014) did not place the Mississippi material in Macroacaena tridens “because they lack a very long extension on sternite 5, have a hook-like structure on sternite 5 that does not appear to be present in Bournelyreidus, and have an articulation of sternites

4 and 5 similar to that of the Lyreidinae and not the Marylyreidinae […] we place

Raninella tridens sensu Roberts within Macroacaena, based upon its possession of an anterior margin about half the maximum carapace width, two anterolateral spines, and what appears to be a flared anterior margin,” (p. 248-249). This assessment is curious because some of these morphologic characters attributed to Macroacaena are in fact

77 characters diagnostic of Bournelyreidus (two anterior spines), or possessed by both genera (anterior margin about half the maximum carapace width) as are attributed to the Lyreidinae that are actually characters of Marylyreidinae (very large lateral projections on sternite 5 that extend as far as lateral margin of carapace; though it is not specified that these are the structures being referred to by Karasawa et al. (2014).

Type material of Bournelyreidus eysunesensis possesses a prominent tubercle on the fourth somite and the presence or absence of a lyreidid hook is not discussed.

Illustrations of the paratype MGUH 21.594 illustrate that the ventral material becomes fragmentary, posteriorly. If the lyreidid hook is present on this species, it is not preserved.

Because the type material of Macroacaena tridens does not posess a venter, the placement of the New Jersey specimen in Macroacaena tridens (Roberts, 1967) is questionable. It should be considered separately from the placement of the

Mississippi specimens until more material becomes available from New Jersey.

The three described species of Bournelyreidus include one species from the

Upper Cretaceous Pierre Shale of South Dakota, Bournelyreidus oaheensis (Bishop,

1978). Bournelyreidus oaheensis also has a narrow fourth sternite but has no other sternites preserved. This species also has two tubercles between the anterolateral spine and outer orbital spine and the branchiocardiac grooves continue posteriorly to a weakly concave outward depression on either side of the axis, unlike

Bournelyreidus n. sp. It is noteworthy that this species of Bournelyreidus has tubercles between the anterolateral spines, as does the diagnosis of Macroacaena by

Karasawa et al. (2014), which states that Macroacaena “may [have] a small

78 anterolateral protuberance between anterolateral spine and outer orbital spine, (p.

246).

Bournelyreidus laevis (Schlüter in Von der Marck and Schlüter, 1868) of

Germany also possesses a relatively un-ornamented carapace and two anterolateral spines. The specimen illustrated is a line drawing of the carapace. A brief description of the claw is also included; no sternal morphology is preserved. This species does not possess any morphology that is preserved to differentiate it from Bournelyreidus n. sp., but the lack of substantial evidence to confirm it is the same species coupled with the locality of the specimen in Europe suggests that this assessment is not most parsimonious.

Bournelyreidus carlilensis (Feldmann and Maxey, 1980) from the Carlile

Shale (Turonian) of Kansas, has poorly preserved sternites, but also possesses two anterolateral spines and the slightly sinuous carapace ornamentation of the cardiac region similar to B. oaheensis. Specimen GAB 37-333 of Bournelyreidus n. sp. has a faint depression around the cardiac region on some of the larger specimens, but the

There are no other genera of Lyreidinae described by Karasawa et al. (2014) with the form of the posterior projection of the “lyreidid hook” that is present on the

Mississippi specimens, including Macroacaena, the genus to which Bournelyreidus tridens sensu Roberts (1967) was reassigned. The morphology of the hook on the

Mississippi specimens is most similar to Rogueus orri Berglund and Feldmann, 1989

(Karasawa et al., 2014, p. 243, fig. 10), but this species ranges from the Paleocene –

Eocene, has a completely different anterior margin, and an incomplete suture between

79 sternites 4 and 5 that “coils” at lateral margins, though this last character superficially looks like episternite 4 on Bournelyreidus n. sp.

Specimens referred to Marylyreidinae possess the “lyreidid hook” of Guinot

(1979) and Karasawa et al. (2014) but Bournelyreidus n. sp. has a hook that is neither quite in the same placement (the hook on Bournelyreidus n. sp. is closer to the sixth sternite) nor of the same morphology (Bournelyreidus n. sp. is less pointed and does not extend as far from the carapace) as illustrated by Guinot (1993) and does not possess a distal or subdistal peg. As such, the Mississippi specimens possess unique dorsal and sternal morphology for the genus while conforming to Bournelyreidus most obviously by possessing two anterolateral spines. It is not apparent by the phylogeny of Karasawa et al. (2014) that the presence of the lyreidid hook is a particularly indicative morphologic character for any particular genus within the

Marylyreidinae, therefore supporting the placement of Bournelyreidus n. sp. in the genus based on all other characters except for the slight axial keel on the dorsal carapace.

Superfamily CALAPPOIDEA De Haan, 1833

Family AETHRIDAE Dana, 1851

Genus PREHEPATUS Rathbun, 1935a

Type species. – Prehepatus cretaceous Rathbun, 1935.

Included species.— Prehepatus cretaceous Rathbun, 1935; Prehepatus dilksi

Roberts, 1962; Prehepatus mexicanus Schweitzer et al., 2006; Prehepatus hodgesi

Bishop, 1983; Prehepatus harrisi, Bishop, 1985; Prehepatus pawpawensis Rathbun,

1935.

Generic diagnosis.— “Left merus triangular, with large nodes on outer surface; carpus longer than high, with nodes arranged into rows of 3 or 4 each parallel to distal margin. Right and left chelae longer than high, becoming higher distally; upper, lower, and outer surfaces ornamented with low, conical spines; proximal margin with distinctive collar, forming articulation with carpus; spines on outer surface generally arranged into rows; inner surface mostly smooth, sometimes with few small tubercles; fixed finger short, directed weakly downward with respect to lower margin of manus, with small tubercles; movable finger with small tubercles at proximal end, keel generally extending from tubercles at least half the distance of the finger,” (Schweitzer et al., 2006, p. 33).

PREHEPATUS HARRISI Bishop, 1985

Figures 10.4, 10.7, 10.14, 10.15

1985. Prehepatus harrisi. Bishop, Journal of Paleontology, 59(5):1028.

1991. Prehepatus harrisi. Bishop, Mississippi Geology, 12(1-2): 13.

2006. Prehepatus harrisi. Schweitzer et al., Bulletin of the Mizunami Fossil Museum, 33: 33.

Diagnosis.— Chelae only. Merus triangular with large nodes on outer surface; outer surface convex, inner surface flattened with few, smaller nodes. Proximal margin with smooth collar. Upper margin slightly rounded. Outer surface ornamented by five or six approximate rows of tubercles. Tubercles becoming smaller toward upper and lower margins. Fingers short and stout; fixed finger continuing straight from lower margin; dactyl strongly downturned, smooth on inner and outer surface, tubercles present on upper surface.

Material examined.— KSU-013 (right chelae), length (l): 19.44 mm, height

(h): 18.9 mm; MPPM 1972.46.449 (right chelae), l: 27.2 mm, h: 23.16 mm; MPPM

1972.46.448 (left chelae), l: 26.38 mm, h: 24.2 mm.

Occurrence.— GAB 37, Blue Springs Locality, Coon Creek Formation, New

Albany, Mississippi, Maastrichtian; the early Maastrichtian of Northeast Mexico

(Vega et al., 1995, p. 247); and the Prairie Bluff Formation in sec. 3. T8S, R3E,

Union County Mississippi, Maastrichtian (Bishop, 1985).

Discussion.— Material from the Blue Springs locality consists of three specimens, one of which is abraded and no longer possesses tubercles or fingers. One specimen is also noteworthy for its smaller size. The largest specimen possesses a nearly complete dactyl, illustrating tubercles on the upper margin, not previously described or preserved in the species.

Schweitzer et al. (2006, p. 32) discussed the placement of Prehepatus within higher taxa. The recognition of Hepatidae Stimpson, 1871 as a junior synonym of

Aethridae Dana, 1851 is confirmed in Schweitzer et al. (2010, p. 85) and retained in this study. The genus is known only from right and left chelae and one carpus, and is known only from North America. It was originally named by Rathbun (1935) for specimens collected in the Cretaceous Gulf Coastal Plain (Schweitzer et al., 2006).

Prehepatus dilksi Roberts, 1962, is known from the Cretaceous of New Jersey

(Feldmann, et al., 2013).

The only record of the genus not from North America is Prehepatus werneri

Fraaye and Collins, 1987, from the Netherlands. However, Jagt et al. (2010) proposed that P. werneri is a junior synonym of Necrocarcinus ornatissimus Forir, 1887 and

82 recommended the combination Roemerus ornatissimus (Forir, 1887) after the genus described by Bishop (1983), but which Jagt et al. (2010) refer to as a ‘form genus’.

Prehepatus werneri may indeed be a junior synonym of Necrocarcinus ornatissimus, as the claws of P. werneri are very similar to those found in association with

Necrocarcinus ornatissimus carapaces (Jagt et al., 2010, fig. 1). Necrocarcinus ornatissimus is herein considered a senior synonym of Prehepatus werneri. This assessment by Jagt et al. (2010) is noteworthy because it limits the genus to North

America. Discovery of associated carapaces with material of Prehepatus claws in

North America will clarify the identification of Prehepatus as a uniquely American genus.

Section HETEROTREMATA Guinot, 1977

Superfamily RETROPLUMOIDEA Gill, 1894

Family RETROPLUMIDAE Gill, 1894

Type Genus.—Retropluma Gill, 1894.

Included Genera.—Archaeopus Rathbun, 1908; Bathypluma de Saint Laurent,

1989; Costacopluma Collins and Morris, 1975; Cristipluma Bishop, 1983;

Loerenthopluma Beschin, Busulini, De Angeli, and Tessier, 1996; Retrocypoda Via

Boada, 1959; Retropluma Gill, 1894.

Family diagnosis.— “Carapace rectilinear or ovoid, wider than long, ornamented with three transverse ridges, usually positioned on protogastric, epibranchial, and mesobranchial regions. Front narrow, downturned, axially sulcate; orbits broad, margins sinuous, often with blunt projection at midlength; orbits

83 terminating in sharp spine. Sternum with transverse ridges mimicking those on the dorsal carapace,” (Schweitzer and Feldmann, 2001, p. 201).

Genus CRISTIPLUMA Bishop, 1983

Type species. – Cristipluma mississippiensis Bishop, 1983.

Included species.— Cristipluma mississippiensis Bishop, 1983.

Generic diagnosis.— As for species.

CRISTIPLUMA MISSISSIPPIENSIS Bishop, 1983

Figures 20.2, 20.7, 20.8

1983. Cristipluma mississippiensis. Bishop, Journal of Crustacean Biology, 3(3): 428.

1985. Cristipluma mississippiensis. Bishop, Journal of Paleontology, 59(5): 1028.

1986. Cristipluma mississippiensis. Bishop, Crustacean issues, 4: 111-142.

1991. Cristipluma mississippiensis. Bishop, Mississippi Geology, 12 (1-2): 12-13.

Diagnosis.— “Carapace ovoid, wider than long, gently arched and poorly differentiated by grooves. Two prominent, continuous transverse ridges crossing carapace; one at midpoint just behind urogastric region, forming widest part of carapace, other behind subtle, crescentic branchiocardiac grooves,” (Bishop, 1983, p.

428).

Description.—Carapace ovoid, slightly wider than long, gently arched with steep lateral margins. Two wide, coarsely punctate, transverse ridges intersect urogastric and cardiac regions as well as cardiac and intestinal regions at the midline, continuing laterally and curving slightly anteriorly at margins. Perpendicular and anterior to transverse ridge is a ridge of equal height, continuing anteriorly through mesogastric region, punctate, tapering to smooth surface halfway to anterior margin.

Margins of mesogastric region demarcated weakly as elevated, pyriform protogastric

84 region, vanishing anteriorly. Carapace with irregularly distributed nodes on depressed sections. Weak grooves defining cardiac region, nearly equi-dimesional, lateral margins concave axially, bounded by arcuate swellings. Posterior margin straight with narrow rim.

Material examined.— Seven specimens from the Blue Springs Locality, Coon

Creek Formation, collected December 17 and 18, 2012. Specimen KSU-301, length:

17.11 mm; width: 23.6 mm; KSU-012, length: 16.24 mm, width: 17.88 mm, frontal width: 11.14 mm; KSU- 302, length: 18.9 mm, width: 20.23 mm; KSU-303, length

15.3 mm, width 18.2 mm; KSU-304, length: 16.38 mm, width: 18.2 mm; KSU-305, length: 12.49 mm, width: 16.23 mm; and two specimens on loan: MMNS IP-2786a, length: 20.9 mm, width: 25.3 mm; MMNS IP-2786b, length: 20.22 mm, width: 21.84 mm; MMNS IP-194, width: 19.78 mm.

Occurrence.— GAB 37, Blue Springs Locality, Coon Creek Formation,

Union County, Mississippi, Maastrichtian.

Discussion.— Cristipluma mississippiensis is the only known species of the genus. The genus differs from others in the family in having the anterior most ridge being developed axially and on to protogastric region. It is also noteworthy that other genera in the family and previous descriptions of C. mississippiensis are referred to as being granulated; specimens of C. mississippiensis observed herein are punctate on raised portions of the carapace cuticle and smooth to very weakly and irregularly granulated on the remainder of the dorsal carapace. Few specimens of Cristipluma mississippiensis exist; Bishop first described a single specimen from the Blue Springs

Locality in 1983. Collection on December 17th and 18th, 2012 yielded seven

85 specimens of the species. The above description includes observations of the preserved posterior margin of a carapace of one of these new specimens. Preservation of cuticle is common but specimens are fragmented.

Bishop (1983) discussed the presence of a retroplumid in the Coon Creek

Formation at the Blue Springs Locality in New Albany, Mississippi, and suggested that the specimen was likely allochthonous because modern species in the family are deep-dwelling. The presence of more specimens of Cristipluma mississippiensis found at the Blue Springs Locality suggests that it is unlikely that these crabs were in fact living in deeper environments. The preservation is fragmentary which could well indicate extensive transport. It is not parsimonious to hypothesize transport of numerous crab carapaces up the continental shelf from deep waters. It therefore seems likely that this monospecific genus from the Cretaceous could have lived in shallower environments than its modern relatives; a change in preferred habitat within a family is not uncommon in the fossil record.

Superfamily PORTUNOIDEA Rafinesque, 1815

Included families.— Carcineretidae Beurlen, 1930; Geryonidae Colosi, 1923;

Mathildellidae Karasawa and kato, 2003a; Portunidae Rafinesque, 1915.

Diagnosis.— “Carapace hexagonal, subhexagonal, rectangular, or transversely

ovate; carapace usually wider than long, usually widest at position of last

anterolateral spine; flattened or weakly vaulted; regions poorly or moderately

defined; anterolateral margins entire or with up to nine spines including outer

orbital spine; front entire or spined; lobe on endopod of first maxilliped (‘portunid

lobe’) sometimes present; chelipeds usually robust; last pair of pereiopods may

have ovate dactyls; sternal sutures 4/5, 5/6, 6/7, and 7/8 usually incomplete; sternite

8 usually visible in ventral view, with penial groove (Portunidae); male abdominal

somites all free or 3-5 fused, sutures may be visible; gonopod 1 strongly curved,

with inflated, strongly hooked base,” (Karasawa and Schweitzer, 2006, p. 59).

Family CARCINERETIDAE Beurlen, 1930

Included genera.— Branchiocarcinus Vega, Feldmann and Sour-

Tovar, 1995; Cancrixantho Van Straelen, 1934; Carcineretes Withers, 1922;

Mascaranada Vega and Feldmann, 1991.

Diagnosis.— “Carapace quadrate or ovate, wider than long, with one or more transverse ridges; transverse ridges located on protogastric, hepatic, or branchial regions; front narrow, may be entire or with blunt spines; supraorbital margin long, with spines and notches; fronto-orbital width nearly equal to maximum carapace width; posterior margin narrow; epibranchial region inflated, arcuate. Male sternum ovate, sternites 1/2 fused with no evidence of suture; sterna suture 2/3 a deep groove; sterna suture 3/4 incomplete; sternite 8 not visible in ventral view. Male abdomen narrow, somites apparently all free, entirely filling space between coxae of fifth pereiopods, reaching to about middle of somite 4. Chelipeds more robust than pereiopods 2-5, manus may have keels on outer surface; propodi and dactyli of fourth and/or fifth pereiopods flattened into paddle-like shape,” (Karasawa and Schweitzer,

2006, p. 59).

NEW GENUS

Type species. – New genus new species (Rathbun, 1926) (Branchiocarcinus flectus sensu Phillips et al., 2013).

Generic description.— As for species.

NEW GENUS NEW SPECIES (Rathbun, 1926)

Figures 10.2, 10.5, 10.8, 10.10, 10.13

2013. Branchiocarcinus flectus. Phillips et al., Paläontologische Zeitschrift, 87(3): p. 14.

Description.— Carapace small, inverted sub-trapezoidal; transversely elongated with greatest width at anterolateral spines. Anterior margin straight.

Rostrum long, downturned and slightly wider at most distal end. Anterolateral margin rounded to posterolateral and finely serrate; strong triangular anterolateral spine divides anterolateral from posterolateral margins. Dorsal carapace regions marked by strong transverse ridges. Protogastric regions flat and intersected at mid-section by transverse ridge. Mesogastric process strongly marked, projecting toward frontal margin. Mesogastric region ovate with a median, transverse ridge, funnel-shaped and extending forward toward frontal margin. Urogastric region transversely subovate with median ridge; cardiac region with strong transverse ridge. Intestinal region flat with weakly defined grooves on lateral margin of the region extending and connecting to weakly defined grooves of the cardiac region. Cervical groove deep and parabolic from outer margin of orbital spines along lateral margin of protogastric regions extending to the posterior margin of the urogastric region. Epibranchial region with posterolaterally inclined short ridge; mesobranchial with slightly longer ridge and metabranchial region with transverse ridge that extends to the base of the anterolater spine. Posterolateral margin posteriorly inclined with a pronounced tubercle at midlength. Posterior margin rimmed and nearly straight.

Endopodite of third maxilliped subtrapezoidal with median longitudinal groove. Right chela slightly larger than left. Merus subrectangular with row of

88 tubercles on lower margin and four strong ridges on outer surface; ridge on dorsal surface of merus with eight tubercles and inner row of three spines. Inner surface of merus smooth; fixed finger triangular with longitudinal groove on outer surface at midline. Occlusal surface dentition of variable size, dactyli curved toward fixed finger. Sternum wide, transversely subovate. Sternite 1 and 2 fused, sternite 3 fused axially and open laterally. Sternite 4 largest; sternites 5, 6, and 7 of same form, rotating posteriorly, sternite 8 smaller. Episternites not visible. Penial groove present and biconcave. Pleon groove extending slightly onto sternite 3. Male abdomen triangular and narrow with triangular telson. Fifth pereiopod reduced.

Material examined.— Three specimens: KSU-3062, length 17.1 mm, width

15.42 mm; MPPM 1972.46.932, length 15.17 mm, width 22.68 mm; MPPM

72.46.459, length 13.82 mm, width 20.02 mm.

Occurrence.— GAB 37, Blue Springs Locality, New Albany, Mississippi,

Maastrichtian. Material examined by Phillips et al. (2013) was collected from the Owl

Creek type locality in Tippah County, Mississippi, the Nixon Sands and the Prairie

Bluff formation in Goodfood, Mississippi, and the Tinton Formation in Monmouth

County, New Jersey, also Maastrichtian.

Discussion.— The Coon Creek specimen, Branchiocarcinus flectus (sensu

Phillips et al., 2013) most notably resembles the dorsal carapace of Lithophylacidae

Van Straelen, 1936 (Lithophylax) and the Goneplacidae MacLeay, 1838

(Icriocarcinus). The specimen shares characters of the Goneplacoidea and the

Portunoidea, but sternal morphology preserved in available material confirms the

placement within the Portunoidea. The lack of sternal morphology for the genus

Branchiocarcinus, originally represented by a single specimen of a single species,

Branchiocarcinus cornatus, has made the placement of this genus difficult.

Likewise, the placement of Icriocarcinus has been questioned.

In 1988, Bishop assigned the new genus and new species, Icriocarcinus xestos to the Carcineretidae. In 1995, Vega, Feldmann, and Sour-Tovar described

Branchiocarcinus cornatus and assigned it to the Carcineretidae. In 2002,

Feldmann and Villamil described a new species Ophthalmoplax triambonatus and revised the Carcineretidae to include five genera: Carcineretes, Branchiocarcinus,

Mascaranada, Ophthalmoplax, and Woodbinax. Also in 2002, Schweitzer et al. transferred Ichriocarcinus into the Goneplacidae. In 2003, Longusorbis was moved by Schweitzer et al. into the Carcineretidae and included Branchiocarcinus,

?Cancrixantho, Carcineretes, Longusorbis, Mascaranada, Opthalmoplax,

Woodbinax, and tentatively Lithophylax. In 2005, Števčić erected the subfamily,

Icriocarcininae. In 2006, Karasawa and Schweitzer conducted a phylogenetic study of the Xanthoidea sensu lato and maintained Icriocarcinus within the Goneplacidae and Branchiocarcinus in the Carcineretidae, with the included genera: Carcineretes,

Longusorbis, Mascaranada, Ophthalmoplax, Woodbinax, possibly Cancrixantho, and Lithophylax. In 2007, Schweitzer et al. revised the Carcineretidae and removed

Branchiocarcinus without assigning the genus to a higher taxon. The Carcineretidae retained Carcineretes, Mascaranada, and ?Cancrixantho. Ophthalmoplax and

Longusorbis were assigned to the Portunidae. In 2008, Karasawa et al. revised the

Portunoidea and retained the Carcineretidae as assigned in 2007. In 2010,

Schweitzer et al. included Branchiocarcinus in the Carcineretidae. In 2013, Bishop

90 et al. assigned Branchiocarcinus and Icriocarcinus to a newly erected family, the

Icriocarcinidae. As should be evident, much work has been done surrounding the placement of Branchiocarcinus and Icriocarcinus, but their placements remain unclear.

The characters that Branchiocarcinus flectus shares with the Goneplacoidea include having a straight front that sometimes has a median notch or projection

(though, this is also observed in the Portunoidea), the anterior end of the sterno- abdominal cavity reaches the anterior of thoracic sternite 4 (in B. flectus it actually extends beyond sternite 4 slightly), and goneoplacids are usually heterochelous (B. flectus is weakly heterochelous). The species also lacks some characters that are distinctive within the Portunoidea, including incomplete sternal sutures of 4/5, 5/6,

6/7, and 7/8. Other characters are merely not observed in the available material and cannot be confirmed or rejected at present in B. flectus because these specific parts are not preserved. Those characters of Portunoidea that are not preserved include a

‘portunid lobe’ (sometimes) on the endopod of the first maxilliped, male pleomere 3

(almost always) with transverse keel, the last pair of pereiopods having (sometimes) ovate dactyls, and the gonopod being strongly curved, with an inflated, strongly hooked base. It is no wonder Phillips et al. (2013) suggested new higher taxa at the family level for the species.

Branchiocarcinus flectus should remain in the Portunoidea, as Phillips et al.

(2013) suggested, because of the abundance of supporting morphologic evidence including the hexagonal carapace that is widest at the last anterolateral spine, regions being poorly to moderately well defined (arguably well defined in B.

flectus), arcuate epibranchial ridges, male sternite 8 visible posteriorly, penial

groove present, and sternite 8 visible in ventral view.

Branchiocarcinus flectus (Rathbun, 1926) sensu Phillips et al., 2013 was assigned to a new family, Icriocarcinidae, which embraced Branchiocarcinus and

Icriocarcinus (Phillips et al., 2013). Bishop (1988) originally assigned Icriocarcinus to the Carcineretidae within the Portunoidea and Vega et al. (1995) subsequently assigned Branchiocarcinus to that family. Icriocarcinus was later assigned to the

Gonioplacidae in the Goneplacoidea in a phylogentetic analysis utilizing extant and extinct families (Karasawa and Schweitzer, 2006). The coded characters that set the included genera apart from those of other similar families and documented assignment to the Goneoplacidae were: front of the carapace straight, without medial notch; upper orbital margins entire with notch between frontal margin and the orbits indistinct; the width of the male pleon fills entire space between the 5th coxae; all male somites free; sternum broad and ovate; sternite 8 not visible in ventral view; sterno-abdominal cavity reaching anterior of sternite 4, and gonopods stout and sinuous (Karasawa and Schweitzer, 2006). Števčić (2005) erected the subfamily,

Icriocarcininae to embrace Icriocarcinus, and retained this subfamily within the

Goneplacidae. Phillips et al. (2013) suggested that this subfamily should be elevated to family level, Icriocarcinidae, within which they also included Branchiocarcinus.

Phillips et al. (2013) referred the newly erected family to the Portunoidea.

Phillips et al. (2013) cited nine characters in common between Icriocarcinus and Branchiocarcinus to support the placement of the two genera into a new family, but these characters are quite generalized. Characters include: an inverted

92 subtrapezoidal carapace; slender eyestalks; narrow and projected “pseudorostrum;” anterolateral spines; transverse ridges on the dorsal carapace; small tubercles along the posterolateral margin; subovate sternum with interrupted sutures 4/5; longitudinal carinae on outer surface of chelipeds, two rows of spines on the dorsal surface of the palms, and a short, curved dactylus; and slender pereiopods 2/5 with unciform dactyli. Many of these characters are also shared with the Lithophylacidae, a

Cenomanian family.

Dorsal carapace, cheliped, and sternal morphology of Branchiocarcinus flectus (Rathbun, 1929) are of the appropriate morphology for placement of the species within the Carcineretidae in most characters. This is where the genus was originally placed and is fitting except for a few characters on the sternum. The extension of the pleon to the third sternite is present in B. flectus, but it is a character that is not observed in the Carcineretidae, Goneplacidae, Lithophylacidae, or

Icriocarcinus. The family Carcineretidae also specifies that the eighth sternite is not visible in ventral view (though it is in the superfamily), but this is not the case in B. flectus, because the 8th sternite is in fact visible in ventral view. The 8th sternite is visible in Lithophylacidae and is also reduced and the male telso also reaches the margin of the 3rd sternite.

This is the only observable character that is not congruent with the diagnosis of the Carcineretidae but present in the Goneplacidae. Branchiocarcinus flectussensu

Phillips et al., 2013 also possesses a lateral groove on the third sternite, a character unique to the Carcineretidae and Lithophylacidae. The nature of the pereiopods as described by Phillips et al. (2013) are also appropriate for the Carcineretidae and are

93 confirmed through illustrations provided by Phillips et al. (2013). Pereiopods cannot be observed in the material at hand. Phillips et al. (2013) cited that the dactyli of pereiopod 2 through 5 were hooked and slightly curved, but neither feature can be verified by their illustrations. Whether or not the pleonal segments, particularly 3 through 5, are fused or free is also important but cannot be discerned from the description by and illustrations in Phillips et al. (2013) or material at hand. Because little observed morphology can set Branchiocarcinus flectus apart from the

Carcineretidae and the preservation of Branchiocarcinus cornatus does not permit a thorough understanding of the genus, Branchiocarcinus flectus sensu Phillips et al.,

2013 is herein retained within the Carcineretidae.

Because Branchiocarcinus cornatus Feldmann and Vega, 1995 was the first and only species assigned to the genus, as described from an external mold of a carapace. Workers including Phillips et al. (2013) have noted that some characters could be artifacts of preservation. Branchiocarcinus flectus (Rathbun, 1926) sensu

Phillips et al., 2013 is the second species to be included in the genus and has representation by a number of specimens, some of which possess sterna. Because the type species lacks sternal morphology and the carapace is of poor preservation, one might take caution in assigning B. flectus to the genus and should be considered further evidence to take caution when assigning it to higher taxa. Because of this, as well as the shared characters with other genera closely resembling the dorsal carapace of Branchiocarcinus flectus sensu Phillips et al., 2013, the Coon Creek specimens are assigned to a new genus.

The included description of Branchiocarcinus flectus utilizes described characters from Phillips et al. (2013) and those observed from three samples from loaned material. Described morphology by Phillips et al. (2013), such as the nature of pereiopods 2 through 4 and pleon characters, were not included when they could not be verified.

Rathbun, (1926) named a single, poorly preserved claw from the Coon Creek

Formation Eryma flecta. Since then, most researchers (Bishop 1983, 1985, 1986a,

1986b; Phillips et al., 2013) have suggested their uncertainty at the assignment of the claw to Eryma. Phillips et al., (2013) determined that the claw Rathbun (1926) assigned to Eryma was articulated with an undescribed species from the Blue Springs

Locality, creating the new combination, Branchiocarcinus flectus. Having observed the claw Rathbun (1926) named Eryma flecta, this claw lacks much of the detail of claws preserved and articulated with Branchiocarcinus flectus. However, this specimen will never be able to provide more information than it already has with such poor preservation. The original claw has a narrower palm and no spines or carina preserved to confirm it is that of B. flectus sensu Phillips et al., 2013. The proximal margin of the palm of the “Eryma flecta” Rathbun (1926) specimen possesses a dorsally up-turned process, where as the Coon Creek specimens do not.

The lack of ornamentation on the Rathbun (1926) specimen is not necessarily of consequence: close attention must be paid to the layer of cuticle preserved, as exemplified in specimen MHN LM 2005.1.14 of Lithophylax trigeri (Guinot and

Breton, 2006). This specimen is of nearly identical form and size of Branchiocarcinus flectus sensu Phillips et al., 2013, but lacks the spinose ornament. Though Guinot and

Breton, 2006 suggest that this specimen possesses exocuticle, thin section images of other specimens (Guinot and Breton, 2006, p. 598, Fig. 4B) do not indicate exocuticle. The specimen illustrated in Fig. 3 also does not appear to have exocuticle and endocuticle appears smoothed from abrasion. Because the proximal margin of the original specimen “Eryma flecta” does not match that of Branchiocarcinus flectus sensu Phillips et al., 2013, the Coon Creek Formation specimens are assigned to a new species.

INCERTAE SEDIS

MAJID (?) FRAGMENT 1

Figure 10.12

Description.—Fragmented anterior portion of carapace, only. Frontal margin narrow, extending into wide rostrum formed by two wide, projections flared outward.

Frontal margin comprising 1/3 of total carapace width. Medial carina with two spines most anteriorly followed by a pair of two spines directly anterior to well defined cervical groove; groove concave forward axially, turning posteriorly at lateral margins. Branchial region slightly inflated. Mandibles (?) with inward-directed processes.

Occurrence.— Coon Creek Formation, Locality MS.73.033a, Union County,

Mississippi.

Material examined.—MMNS 4108, height: 26 mm, width: 20.3 mm.

MAJID (?) FRAGMENT 2

Figure 10.11

Description.— Specimen slightly wider than long, pleonal somite. Indication of single horizontal suture, rounded on all sides. Vertical keel along axis with well defined depressions on either side; inflated laterally.

Occurrence.— Coon Creek Formation, Locality MGS173, Union County,

Mississippi.

Material examined.—MMNS 4016, height: 14.66 mm, width: 15.38 mm.

Discussion.—Both specimens are a single occurrence from the Coon Creek

Formation. This is the first potential documentation of Majidae in the Coon Creek

Formation and is a significant occurrence because majids are un-common in the

Cretaceous.

DISCUSSION

i. Taphonomy

Many specimens of fossil lobster are preserved with the pleon in the contracted state. This is observed as the pleon curled underneath the body and the telson closest to the most anterior portion of the animal. Most specimens of Linuparus are not preserved like this but with the pleon outstretched straight behind the body in a single plane. The specimen from Keyes’s personal collection is very peculiar because it has both the telson and the most anterior end of the pleon upturned to point dorsally (Fig. 11.5, 11.6, 11.7).

This concave upward flexure of the pleon is not possible with the ventral tissue intact.

Because of this, fossilization must have occurred post rigor mortis. As the more flexible tissue was broken down by enzymes (bacteria), the pleon would be physically capable of

97 bending in the upturned direction. The hypothesis is difficult to confirm because the venter of the pleon is not preserved.

Variation in preservational style is very apparent in specimens of Tetracarcinus subquadratus as well as a few specimens of Linuparus sp. Most specimens of Coon

Creek Formation decapods are preserved as internal molds or are replaced by phosphate.

Few specimens have silica associated with their preservation, and the majority of this type is confined to T. subquadratus. Thin sections of T. subquadratus and D. australis indicate the possibility of more diagenetic processes effecting T. subquadratus because of this aluminum/silica film on the exterior of the cuticle. Elemental mapping of T. subquadratus indicates that specimens preserved with white, iridescent cuticle possess silica and aluminum (Fig. 21), whereas most other specimens are phosphorous and calcium (Figs. 22 - 25). Specimens also fluoresced in long and shortwave ultraviolet light with a vibrant yellow and orange color in localized places. These specimens also had small flecks that fluoresced with a bright blue. The presence of silica and aluminum is not observed in thin section (Figs. 26 and 27). The location of these minerals in elemental maps indicates that the cuticle is not replaced but rather the exterior of the cuticle is covered in a thin layer of clay minerals (Fig. 21). Though preservational style appears variable in hand sample, these differences are likely relicts of weathering of phosphatized remains to clay and do not indicate variability due to epibionts, exo- vs. endo-cuticle, taxon, etc.

Fossils do not have any preferred orientation. Carapaces of larger crabs sometimes have partial legs attached: it is also common to find more complete crabs with

98 disarticulated leg sections nearby at the Blue Springs Locality. Some carapaces lack sturnum but have claws and legs concentrated in the cavity of the underside of the carapace where the sternum would be. Many claw and leg fragments are found weathered out of the outcrop.

Bishop and Williams (1991) discussed the potential for preservation of molt as well as corpses in the fossil record. They suggested that carapaces preserved with attached sterna indicate that they are the remains of corpses, not molts (Bishop and

Williams, 1991, pg. 458). Bishop also described a configuration of the carapace, where it is somewhat articulated and indicative of a molt as well, referred to as Salter’s position, where the “carapace is lifted and flipped forward, making an angle of about 90° with the sternum. Molts are distinguished from corpses by noting that the carapace has split along the pleural sutures, and the carapace lying outside of the pleural sutures are still attached to the sterna plastron,” (Bishop, 1972 pg. 633). There are no specimens from the Coon

Creek Formation observed in Salter’s position, but many specimens are internal molds of the carapace and therefore may indicate preservation of a molt.

SEM images suggest that internal, microscopic structure of cuticle is well preserved in most specimens, regardless of species and physical variability in cuticular morphology or apparent preservation differences (Figs. 21 – 28). This can be seen as light and dark laminations of cuticle, fibrous, vertical minerals perpendicular to “laminations” of in-filled spaces between pore canals, and microscopic patterns on external surface of cuticle of Dakoticancer australis (Fig. 28).

Though some decapods are preserved in concretions and some cuticle illustrates concoidal fracture using scanning electron microscopy, concretions do not illustrate similar qualities to concretions containing decapods found in correlative beds in the Bear

Paw Shale of the Western Interior Seaway, indicating dissimilarity in preservation and environment (Feldmann et al., 2012). X-ray florescence and reflectance data lacked utility and are therefore not illustrated herein.

ii. Paleoecology

The Coon Creek Formation is fossiliferous, with excellent preservation of a diverse assemblage of decapod taxa, and assemblages of so many fossil decapods are a rarity, yet the Coon Creek fauna has been underutilized for its paleoecological potential.

The vertebrate taxa (Carpenter et al., 2003; Gibson et al., 2008; Gallagher et al., 2012;

Stringer, 2003; Whetstone, 1977), ichnofossils (Hall and Savrda, 2008), and associated sediments have been addressed from the formation or correlative units within the associated seas of North America, but little has been done to qualify or quantify the taxa at the community level. Trophic level interactions between invertebrates have been addressed in broad schemes, but these studies have largely excluded the (Finnegan, 2013;

Bush and Pruss, 2013).

The Coon Creek Formation ecology was first discussed in a thesis by Moore

(1974). Wade (1927), Rathbun (in Wade, 1927) and Bishop (1983; 2003a; 2003b) have also worked on the Coon Creek fauna and specifically its decapod fauna. Other invertebrates studied include the cephalopods (Larson, 2012), the oysters (Lerman, 1965;

Griffin and Gibson, 1999), echinoderms (Ciampaglio, 2003), foraminifera (Lobegeier and

Lusk, 2008; Frederick, 2003), and ostracodes (Granata, 1960). These works do not assess

100 community structures, nor do they suggest interpretations of trophic interactions. Specific food web interactions within the Maastrichtian tend to be isolated as single-taxon papers

(Kruta et al., 2011).

Interpreting trophic interactions at the community level at the Blue Springs

Locality, and possibly Mesozoic aged marine fossil assemblages in general, pose a few unique challenges. Mesozoic fauna included large, nektonic predators, such as mosasaurs, which must be accounted for in the trophic scheme – a level of interaction that is not observed in more primitive associations, such as the Ordovician of Cincinnati, Ohio.

Because of this, these assemblages are by definition composed of allochthonous fossils that should not be omitted from the study.

Taphonomic biases are of great importance in the invertebrates as well. Many mollusks at the Blue Springs Locality are brittle and poorly preserved, creating an inaccurate assessment of their biomass, diversity, and number of individuals. In contrast, the decapods pose a taphonomic bias skewed toward overrepresentation of individuals due to possible preservation of multiple molts per individual.

A challenge when assessing any fossil assemblage for its potential as a complete ecosystem is, of course, the absence of most primary producers and meiofauna. Workers have recently addressed and proposed a way to quantify the “soft and squishy” of the paleo realm, (Sperling, 2013), but these methods are not practical for paleo-community assessments.

Personal observations include foraminifera, seen in thin section and sediment samples. Some foraminiferal forms are globose or planktonic while others are more characteristic of benthic forms. Frederick (2003) sampled the type section of the Coon

101

Creek Formation and identified 34 species of foraminifera, 31 of which were benthic and three planktic. The benthic species are dominated by the genera Cibicides, Anomalina, and Bolivina. Epibenthic forms indicate a normal shelf condition. Planktic foraminifera are long ranging taxa, Pseudoguembelina globosa and Archaeoglobigerina cretacea, and therefore cannot be used as age indicators for the Coon Creek Formation.

Shark teeth and mosasaur vertebrae were identified via oral communication with

Dr. George Phillips on site (2012, 2013). Turritellid and naticid gastropods, bivalves, and many oysters (Exogyra sp.; see Griffin and Gibson, 1999 on the bioimmuration of

Exogyra) were also collected, though gastropod and bivalve preservation was very poor at the Blue Springs Locality. Otoliths and very small, button shaped corals were also collected. Internal molds of burrows, mosasaur vertebrae, and coprolites are tentatively identified (personal communication, Richard Keyes, 2013) from collecting at the Blue

Springs Locality as well.

Ammonites are known from correlative units and those of the Campanian

(Cobban and Kennedy, 1994; Larson, 2012; Ebersole, 2009). Bruce Wade (1926) compiled the first thorough assessment of the Coon Creek fauna. Recent paleoecological synthesis by Moore (1974) included systematics and paleoecology, but did not assess the decapods. Many abstracts have been published on the Coon Creek fauna in the last ten or

20 years (Stringer, 2003; Markin and Gibson, 2010; Lobegeier and Lusk, 2008; Larson,

2003, Jones and Gibson, 2000; Gibson et al., 2008; Frederick, 2003; Dockery and

Bandel, 2003; Ciampaglio, 2003) and include invertebrate, vertebrate, micro, larval forms, and macro fossils.

102

iii. Paleobiogeographic Comparisons

Genera of retroplumids include Costacopluma Collins and Morris, 1975;

Archeopus Rathbun, 1908; Loerentheya Beurlen in Lőrenthey and Beurlen, 1929;

Loerenthopluma Beschin, Busulini, De Angeli and Tessier, 1996a; Retrocypoda Via,

1959, Retropluma Gill, 1894; Serrablopluma Artal et al., 201;, and Gaudipluma

Artal et al., 2013. The fossil genus Costacopluma has been reported to occur in the

Late Cretaceous rocks of Greenland, Mexico, and West Africa (Feldmann and Portell,

2007). It continues into the Paleocene, occurring in Argentina, Brazil, Venezuela,

Kashmir, and West Africa but does not occur in the United States until the Eocene

(Feldmann and Portell, 2007). Costacopluma is thought to be derived from Archeopus and to have originated in Mexico, where it radiated into the North Atlantic and

Tethys. The only known occurrence of Costacopluma is in the U.S. from the Eocene of southern Alabama (Feldmann and Portell, 2007). Two other genera, Gaudipluma and Serrablopluma, are reported from the Eocene of northern Spain (Artal et al.,

2013). The retroplumids are common and diverse in the Eocene of northern Spain, the

Eocene and Oligocene of northern Italy, and the Eocene of southern France (Artal et al., 2013). Cristipluma mississippiensis does not occur in any other proximal geographic location.

Decapod taxa are compared at the species level throughout correlative units in

North America. They are compared here on the basis of general geographic range during the Maastrichtian. Species found only in the Coon Creek Formation (Mississippi

Embayment) include: Hoploparia tennesseensis; Hoploparia mcnairiensis; Linuparus new species; Linuparus sp.; Palaeopetrochirus enigmus; Seorsus wadei; Bournelyreidus

103 new species; Cristipluma mississippiensis; Species that occur in the Coon Creek

Formation as well as the Atlantic Coastal Plain: Hoploparia georgeana; Mesostylus mortoni; Tetracarcinus subquadratus; Avitelmessus grapsoideus; Cretacoranina testacea;New genus new species (Carcineretidae); Latheticocarcinus atlanticus; Species that occur in the Coon Creek Formation as well as the Western Interior Seaway:

Tetracarcinus subquadratus; Avitelmessus grapsoideus; Species that occur in the Coon

Creek Formation as well as the Gulf Coastal Plain: Mesostylus mortoni; Dakoticancer australis; Avitelmessus grapsoideus; Prehepatus harrisi (see Fig. 30). Shared species from all surrounding North American localities supports the hypothesis that the

Mississippi Embayment acted as an ecotone during the Maastrichtian.

Schweitzer and Feldmann (2005) discussed the presence of most lineages across the Cretaceous – Paleogene boundary. The extinctions present in decapods at this time are rather pseudoextinction. In fact, the Late Cretaceous is a time of adaptive radiation in the Central Americas. This may be attributed to the composition of decapod exoskeletons, which are more robust to fluctuations in the Carbonate Compensation

Depth as aposed to mollusks (Schweitzer and Feldmann, 2005). The Late Cretaceous was rather a time of rapid evolution and faunal turnover, with the greatest number of last occurances of genera (1/3 of the total genera known from the Late Cretaceous) taking place in the Maastrichtian. About half of these last occurances were in the Central

Americas (Schweitzer and Feldmann, 2005). New species described herein are both first and last occurances as there are no known specimens from other formations. Finding species that survive across the K – Pg boundary will continue to be difficult due to the paucity of Paleocene rocks.

104

The presense of species at high and low latitudes, such as Avitelmesus grapsoideous and Tetracarcinus subquadratus, may be a good example of ecological generalists or eurytypic species surviving both in the Western Interior Seaway as well as

Atlantic Coastal Plain, and Mississippi Embayment. Though geographically diverse, these mono-specific genera do not survive the K – Pg extinction.

Cope et al. (2005) described the fauna of the Clayton Formation (Paleocene,

Danian) from the northeast of the Mississippi Embayment in Olmsted, Pulaski Co.,

Illinois. Four genera: Linuparus, Hoploparia, Paguristes, and Mesostylus, are reported from the Clayton Formation and also occur in the Coon Creek Formation. Of those four genera, two species, Hoploparia tennesseensis and Mesostylus mortoni, occur in the

Maastrichtian and Danian of the Mississippi Embayment.

Another noteworthy biogeographic comparison is the absence of intersex characters in the Dakoticanceridae in all other localities and species. Bishop (1974) first described these qualities in Dakoticancer overanus of the Western Interior Seaway, which was subsequently studied further by Jones (2014). The species is found only in the

Western Interior Seaway and is the only species in the family with this morphological variation.

The intersex specimen was first found at Sitting Bull Locality near Mobridge,

South Dakota (specimen number USNM #173542) (4-2007) (Bishop, 1974, p. 215). A new example of Dakoticancer intersex was discovered in 1976 by Mr. Harry Mendryk at the Mobridge Locality. This specimen had female gonopores at the base of the fourth leg and male gonopores at the base of the fifth leg which is different from the previously

105 described intersex configuration (Bishop, 1976-78, pg. 196). Jones (2014) described other forms of intersex specimens, which were also from Mobridge. Coon Creek Formation specimens of Dakoticancer australis yielded 78 specimens with sterna preserved, of which 30 were identified as female and 58 were identified as male. This ratio of more males than females also occurs in the Dakoticancer overanus population in Mobridge

(Jones, 2014). Of these 78 specimens, none possessed intersex characters. The preservation of Coon Creek specimens was quite different from those of Mobridge; the

Mobridge specimens are generally preserved in individual concretions and the Coon

Creek Specimens for the majority weathered out of the rock unit readily. Because of this, gonopores were often damaged or the cuticle from the area was missing entirely. Missing gonopores from the record of Coon Creek material could be forming a preservation bias and could be the reason that this intersex pattern is not observed in Coon Creek specimens of Dakoticancer australis at this time.

CONCLUSIONS

 Two new species, Linuparus new species and Bournelyridus new species are

described from the Coon Creek Formation.

 The Coon Creek Formation decapod specimens are very well preserved,

possessing ornamentation and cuticle.

 Decapod specimens are generally phosphatized, but some specimens, specifically

Tetracarcinus subquadratus and fragments of Linuparus sp. are replaced with

silica. These specimens also illustrate fine laminations of cuticle in cross section.

106

 The preservational style of decapods in the Coon Creek Formation do not possess

the same microbial sheath structure described by Feldmann et al. (2012) from the

Western Interior Bear Paw Shale specimens.

 Seventeen species are described from the Coon Creek Formation, including

Hoploparia tennesseensis; Hoploparia mcnairiensis; Linuparus new species;

Linuparus sp.; Palaeopetrochirus enigmus; Seorsus wadei; Bournelyreidus new

species; Cristipluma mississippiensis; Hoploparia georeana; Mesostylus mortoni;

Tetracarcinus subquadratus; Avitelmessus grapsoideus; Cretacoranina testacea;

New genus new species (Carcineretidae);Dakoticancer australis; Prehepatus

harrisi; Latheticocarcinus atlanticus.

 Eight species of decapod occur exclusively in the Coon Creek Formation.

 Seven species of decapod occur in the Coon Creek Formation and the Atlantic

Coastal Plain.

 Four species of decapod occur in the Coon Creek Formation and Gulf Coastal

Plain.

 Two species of decapod occur in the Coon Creek Formation and Western Interior

Seaway.

 The presence of a diverse decapod fauna in the Coon Creek Formation is

congruent with the assessment of Schweitzer and Feldmann (2005): evolution and

extinction rates were high in the Late Cretaceous. Rapid faunal turnover,

adaptation, and radiation were commonplace in Central America during the

Maastrichtian.



107

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Figure 1. Maastrichtian outcrops of North America, black. Intercontinental seaway outlined, gray (modified from Schuchert, 1955).

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Figure 2. Late Cretaceous facies along the Mississippi Embayment, with dominantly siliciclasitic sediments in the Selmer – Tupelo area (Mancini et al., 1995).

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Figure 3. Regional Upper Cretaceous stratigraphy (Smith, C. C., 1989).

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Figure 4. Interpretation of stratigraphic context of the Coon Creek beds (Mancini et al., 1995).

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Figure 5. Blue Springs Locality of the Coon Creek Formation near New Albany, Mississippi (Google Earth).

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A B C

D E F

G H I Figure 6. Thin section images, magnification 10 X 0.4. A) Burrow at center of flat concretion with larger grains; B) Cuticle in light concretion; C0 outer edge of concretion with dark, fine grained rind and larger sediment lightly cemented to the outside; D) glauconite grains in sediment; E) fine grains in concretion; F) cross section of burrow; G) fine grained rind of a concretion with coarse grains internally; H) concretion from image A with calcite infilling crack to the burrow at center; I) light colored concretion with coarse grains.

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A B C

D E F

Figure 7. Thin section images, magnification 10 X 20. A) glauconite grain; B) concretion in plain light; C) concretion in cross polarized light; D) cuticle and surrounding grains; E) coarse grains; F) glauconite grain.

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A B

D E

Figure 8. Thin section images, magnification 10 X 40. A) foraminifera test with glauconite infilling chambers; B) weathering mineral grain into mica; C) gastropod protoconch; D) weathering mineral grain into mica in cross polarized light; E) mineral grains in cross polarized light without preferred orientation.

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Figure 9. Figure 9.1, 9.2. Hoploparia tennesseensis Rathbun, 1926. Specimen MPPM 1972.46.446. Scale bar = 1 cm. 1) view of inner surface of left lateral cutting claw and segments of pleon; 2) view of outer surface of left lateral cutting claw. Figure 9.3. Hoploparia tennesseensis Rathbun, 1926, crushing claw. Specimen MPPM 005. Scale bar = 1 cm. Figure 9.4. Hoploparia tennesseensis Rathbun, 1926, crushing claw. Specimen MPPM 004. Scale bar = 1 cm. Figure 9.5. Hoploparia tennesseensis? Rathbun, 1926, cutting claw. Specimen MPPM 1972.46.447. Scale bar = 1 cm. Figure 9.6. Hoploparia tennesseensis Rathbun, 1926, carapace and pleon. Specimen B-001. Scale bar = 1 cm. Figure 9.7, 9.8, 9.9. Hoploparia georgeana Rathbun, 1935, claw. Specimen GAB 37-1147. Scale bar = 1 cm.

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Figure 10. Figure 10.1. Mesostylus mortoni (Pilsbry, 1901). Specimen MPPM 72.46.456; Figure 10.2. New genus new species. Specimen MPPM 72.46.459; scale bar = 1 cm. Figure 10.3. Bournelyreidus new species. Specimen MPPM 1972.46.432; Figure 10.4. Prehepatus harrisi Bishop, 1985. Specimen MPPM 1972.46.449; Figure 10.5. New genus new species. Specimen MPPM 1972.46.432; Figure 10.6. Mesostylus mortoni (Pilsbry, 1901). Specimen MPPM 1974.35.16; Figure 10.7. Prehepatus harrisi Bishop, 1985. Specimen MPPM 1972.46.448; Figure 10.8. New genus new species. Specimen 1972.46.432; Figure 10.9. Bournelyreidus new species. Specimen MMNS IP 3140; Figure 10.10. New genus new species. Specimen MPPM 012 ; Figure 10.11. Majid (?) fragment 2. Specimen MMNS 4016; Figure 10.12. Majid (?) fragment 1. Specimen MMNS 4108; Figure 10.13. Branchiocarcinus flectus (Rathbun, 1926). Specimen MPPM 012; Figure 10.14. Prehepatus harrisi Bishop, 1985. Specimen number KSU – 013; Figure 15. Prehepatus harrisi Bishop, 1985. Specimen number KSU – 013; Figure 10.16. Bournelyreidus new species; specimen GAB 37-832; Figure 10.17. Bournelyreidus new species; specimen GAB 37-832, arrow indicating hook. All Scale bars = 1 cm.

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Figure 11. Figure 11.1, 11.2, 11.3. Linuparus new species. Specimen MMNH 2000.1.11; scale bars = 1 cm. Figure 11.4. Linuparus new species. Specimen MPPM 001; scale bar = 1 cm. Figure 11.5, 11.6, 11.7. Linuparus new species. Specimen from the private collection of Richard Keyes; scale bar = 1 cm.

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Figure 12. Figures 12.1 and 12.2. Tetracarcinus subquadratus. Specimen KSU 008-1; Specimen MPPM 1972.46.51; Figures 12.3 – 12.9. Dakoticancer australis. 12.3: Specimen B-011; 12.4: Specimen B-004; 12.5: Specimen B-007; 12.6: Specimen B 004-1; 12.7: Specimen 37-414; 12.8: Specimen 37-915; 12.9: 37-939. Scale bars = 1 cm.

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Size proportions of Dakoticancridae in the Coon Creek Formation 45 y = 0.9764x + 0.9754 R² = 0.793 40

D. australis 25 T. subquadratus

20 Length(mm) Linear (D. australis)

Linear (T. 15 subquadratus)

10 y = 0.552x + 5.4512 R² = 0.2977

n = 114 5

0 0 10 20 30 40 50 Width (mm)

Figure 13. Species of the Dakoticancridae found at the Blue Springs Locality of the Coon Creek Formation illustrating similar ratios of length to width in the Dakoticancer australis and Tetracarcinus subquadratus throughout ontogeny.

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Dakoticancer australis

Male, 47

Female, 28 Number specimens Number of

n = 75

Figure 14. Number of male and female specimens of Dakoticancer australis in this study. Specimens of Tetracarcinus subquadratus do not have sternum preserved and thus cannot have their sex determined.

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18 16y = -0.2066x + 17.53 14 R² = 0.1963 Female 12 10 Male 8

w:fw (mm) w:fw 6 y = 0.3094x + 1.3568 Linear 4 R² = 0.5406 (Female) 2 Linear 0 (Male)

10 30 n(Female)=6 n(Male)= 13 Width (mm)

Figure 15. Male vs. female ratios of Dakoticancer australis do not illustrate sexual dimorphism when comparing width against ratio of width to frontal width.

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0.26 y = -0.0012x + 0.24 0.2125 R² = 0.0174

0.22 Female 0.2 0.18 Male 0.16

fw :fow (mm) :fow fw Linear 0.14 y = -0.0022x + 0.255 (Female) 0.12 R² = 0.1708 Linear 0.1 (Male)

10 30 n(Female)=6 Width (mm) n(Male)= 13

Figure 16. Male vs. female ratios of Dakoticancer australis do not illustrate sexual dimorphism when comparing width against ratio of frontal width to fronto-orbital width.

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2.5 y = 0.0211x + 1.3223 2.3 R² = 0.4974 Female

2.1 1.9 Male 1.7 y = 0.0119x + 1.5086 1.5 R² = 0.0324

w :fow (mm) :fow w Linear 1.3 (Female) 1.1 0.9 Linear 10 30 (Male) n(Female)=17 Width (mm) n(Male)=28

Figure 17. Male vs. female ratios of Dakoticancer australis do not illustrate sexual dimorphism when comparing width against ratio of width to fronto-orbital width.

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Claw size vs. Granule Size 15

10 5 Right

per mm2 per 0 Left 0 20 40

Number of Granules Granules of Number Height (mm)

Figure 18. Dakoticancer australis claw height vs. granule size.

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3 2.8 2.6 y = 0.0757x + 0.6754 2.4 R² = 0.527 2.2 2 T. subquadratus 1.8

w:fow (mm) w:fow 1.6 Linear (T. 1.4 subquadratus) 1.2 1 9 19 n= 12 w (mm)

0.24

0.22

0.2 0.18 T. subquadratus 0.16

fw:fow (mm) fw:fow 0.14 Linear (T. subquadratus) 0.12 y = -0.0093x + 0.3461 R² = 0.2635 0.1 10 15 20 25 n=26 w (mm)

18 16 y = 0.7958x - 2.5916 R² = 0.7002 14 12 T. subquadratus 10

w:fw (mm) w:fw 8 Linear (T. 6 subquadratus) 4 n=18 9 14 19 24 w (mm)

Figure 19. Tetracarcinus subquadratus measurements of width against ontogenetic ratios illustrating frontal width to fronto-orbital width decreases with increase in size, an inverse relationship of width against frontal width or width against fronto-orbital width.

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Figure 20. Figure 20.1. Cristipluma mississippiensis. Specimen MMNS IP-2786; Figure 20.2. Cretacoranina testacea. Specimen GAB 37-860; Figure 20.3. Cretacoranina testacea. Specimen GAB 37-837; Figure 20.4, 20.6. Cretacoranina testacea. Specimen KSU-011; Figure 20.5. Letheticocarcinus atlanticus (Roberts, 1962). Specimen B-003; Figure 20.7 – 20.9. Cristipluma mississippiensis. Specimen KSU 301; Figure 20.8 and figure 20.9: Specimen KSU 012. Scale bars = 1 cm.

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Sample 5: Tetracarcinus Aluminum Calcium subquadratus

Iron Phosphorous Silica

Figure 21. Elemental maps of cuticle thin section of Tetracarcinus subquadratus. Aluminum and silica are concentrated on the outer margin of the cuticle. Calcium and phosphorous are also present in a relatively high abundance in the surrounding sediment as well as traces very small traces of iron. The absence of data on the lower margin of the cuticle is likely caused by a shadowing effect from uneven sampling surface.

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Sample 2b: Aluminum Calcium Linuparus sp.

Potassium Phosphorous Silica

Sulfur Iron

Figure 22. Elemental maps of cuticle thin section of Linuparus sp. Calcium and phosphorous are concentrated in the cuticle, silica and aluminum are also present in a relatively high abundance in the surrounding sediment as well as traces of sulfur , potassium, and iron.

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Sample 3: Aluminum Calcium Bournelyridus

Phosphorous Silica

Figure 23. Elemental maps of cuticle thin section of Bournelyridus n. sp. Calcium and phosphorous are concentrated in the cuticle but preservation is of a lesser degree than illustrated in other specimens, silica and aluminum are also present in a lower abundance in the surrounding sediment .

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Sample 2b: Hoploparia Aluminum Calcium tennesseensis

Silica Potassium Phosphorous

Sulfur Iron Figure 24. Elemental maps of cuticle thin section of Hoploparia tennesseensis. Calcium and phosphorous are concentrated in the cuticle. Large white spot has formed due to charging of the specimen and is therefore causing a shadow in the elemental data. Silica and aluminum are also present in a relatively high abundance in the surrounding sediment as well as traces of sulfur, potassium, and iron.

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Sample 7: Tetracarcinus Aluminum Calcium subquadratus

Potassium Phosphorous Silica

Iron Magnesium

Figure 25. Elemental maps of cuticle thin section of Linuparus sp. Calcium and phosphorous are concentrated in the cuticle, silica and aluminum are also present in a relatively high abundance in the surrounding sediment as well as traces of sulfur, potassium, and iron.

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Figure 26. Thin section of Tetracarcinus subquadratus carapace cuticle, illustrating phosphatization of endocuticle with laminations and pore canal visible. Magnification 10 X 40.

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Figure 27. Thin section of Dakoticancer australis carapace cuticle, illustrating phosphatization of endocuticle with laminations visible. Magnification 10 X 40.

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A B

C D

E F

Figure 28. SEM images of Dakoticancer australis claw. A) cross section illustrating cuticle on right- hand margin of specimen; B) outer-most layer of endocuticle illustrating micro-ornamentation on lower half of image and pore infilling of endocuticle on upper half of

image; C) pore canal opening, dark spot at center of image; D) endocuticle ornamentation; E) mineralization of spaces between pore canals, high magnification; F) mineralization of spaces between pore canals, low magnification.

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Figure 29. Fifteen species of decapods in the Coon Creek Formation of the Mississippi Embayment. Of these 17, 8 of them are endemic and 8 occur in correlative units on the shelf of North America.

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T. subquadratus

D. australis

P. harrisi H. georgeana M. mortoni

A. grapsoideous

Figure 30. Distribution of species in common with the Coon Creek fauna.

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Table 1. Percent composition of Coon Creek Formation lithology and included concretions.

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Table 2. Table of Linuparus species characters of the most similar species to Linuparus n. sp.

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