Caridea, Anomura, Axiidea, Brachyura) from Alabama and Mississippi, USA

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Journal of Crustacean Biology Advance Access published 4 April 2019 Journal of Crustacean Biology The Crustacean Society Journal of Crustacean Biology 39(3) 279–302, 2019. doi:10.1093/jcbiol/ruz002

Paleogene Decapoda (Caridea, Anomura, Axiidea, Brachyura) from Alabama and Mississippi, USA

Rodney M. Feldmann1, , Carrie E. Schweitzer2 and George Phillips3, Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021 1Department of Geology, Kent State University, Kent, OH 44242, USA; 2Department of Geology, Kent State University at Stark, 6000 Frank Ave. NW, North Canton, OH 44720, USA; and 3Mississippi Museum of Natural Science, 2148 Riverside Dr., Jackson, MS 39202, USA

Corrspondence: R.M. Feldmann; e-mail: [email protected] (Received 17 November 2018; accepted 16 January 2019)

ABSTRACT Investigation of decapod crustaceans from Paleogene sediments of Mississippi has yielded one new Eocene brachyuran species, Matutites periosus n. sp., and six new Oligocene brachyuran species, Lophoranina lincki n. sp., Notopus adinae n. sp., Calappilia granulosa n. sp., C. perreaulti n. sp., C. gableorum n. sp., and Acanthocarpus osborni n. sp. Other occurrences include previously described caridean shrimp, axiids, pagurids, and raninids. Mesostylus, a callianassid, is described for the first time from Danian deposits, all other occurrences being Late Cretaceous. The Oligocene decapod fauna from Gulf Coastal North America, including localities in Alabama and Mississippi, is most like that of Pacific Coastal Baja California and most different from Pacific coastal localities from Oregon northward. Key Words: Alpheidae, Calappidae, Callianassidae, Lyreididae, Matutidae, Paguridae, Raninidae, southeastern USA

INTRODUCTION cyclical fluctuations in carbonate content (Hart et al., 2013). The Clayton Formation is Danian and the Pine Barren beds are early The fauna of the Gulf Coastal Plain of the United States has Danian (Mancini & Tew, 1988). In the superjacent “Sucarnoochee been the subject of several studies of decapod crustaceans; Clay” (Danian-Selandian), Rathbun (1935) reported a prodigious however, continued collection of specimens in Alabama and diversity of decapods. Now known as the Porters Creek Formation Mississippi have yielded many new occurrences including new (MacNeil, 1946), where the clay beds of the lower member are species. Among these, several species of calappid crabs have been exposed east of Camden, Wilcox County, AL, they preserve discovered in Oligocene strata deposited in a variety of shallow- exceptional decapod crustaceans and corals (Rathbun, 1935; shelf environments. We describe and illustrate the new species and Toulmin, 1977). Further to the west, west of Camden, the upper the new occurrences of decapod crustaceans from Alabama and member of the Porters Creek Formation, the Matthews Landing Mississippi and contrast the sedimentary environments in which Marl, contains numerous species of pristinely preserved mollusks the calappid species lived. (Toulmin, 1977). Matutites periosus n. sp. is from the Potterchitto Member of the Cook Mountain Formation (upper Claiborne Group) of Newton GEOLOGICAL SETTING County, MS. The Potterchitto Member is a glauconitic, calcare- The specimens described herein were collected in the eastern ous, clayey sand in Newton County (Merrill et al., 1985). It is Gulf Coastal Plain, Mississippi (MS) and Alabama (AL) (Fig. 1), in equivalent to the upper Lisbon Formation of Alabama; both are sediments of early Paleocene, middle Eocene, and Oligocene age. Bartonian (late Eocene) (Mancini & Tew, 1991). Only a few of Most of the new material is Vicksburg age, or early Oligocene other decapod taxa have been reported from the Cook Mountain (Fig. 2), and most are calappids. and upper Lisbon formations. Neozanthopsis americanus (Rathbun, Giulianolyreidus johnsoni (Rathbun, 1935) (Raninidae) is from 1928) of the Cook Mountain Formation of Texas was reported the Pine Barren Member of the Clayton Formation of Lowndes by Stenzel (1935) from the same unit in Mississippi (an unfigured County, AL, where it occurs with Costacopluma grayi Feldmann “complete specimen and one separate right cheliped”), presum- & Portell, 2007 (Retroplumidae), and Stevea martini Feldmann, ably the Potterchitto Member, and in the vicinity of M. periosus Schweitzer, and Portell, 2014 (Hexapodidae). At this locality n. sp. from the Cook Mountain Formation of Clarke County, (AL.43.002) the Pine Barren Member is a silty mudstone with Mississippi, Rathbun (1935) attributed a single male chela (also

© The Author(s) 2019. Published by Oxford University Press on behalf of The Crustacean Society. All rights reserved. For permissions, please e-mail: [email protected] R. M. FELDMANN ET AL. Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021

Figure 1. Geologic map of localities. Upper portion is a generalized distribution of Paleogene sedimentary groups in the southeastern USA, based on King & Beikman (1974). Lower portion (inset boxed in upper map) emphasizes the units referred to and is based on Bicker (1969) for Mississippi and Szabo et al. (1988) for Alabama. Locality numbers given in text. Alabama units were mapped illustrating modern stream dissection, whereas Mississippi units were not. Projected in State Plane North (NAD83) using ArcMap 10.3 © 1999–2015 Esri, Inc. This figure is available in color at Journal of Crustacean Biology online. unfigured) to “H. americanus Rathbun,” and Dockery (1980) fig- 1992). The Mint Spring Marl, the first lithofacies in the sequence, ured an eroded right propodus as “Harpactocarcinus sp.” Rathbun is a glauconitic, frequently micritic, quartz sand to sandy Marl (1935) attributed an eroded chela from the Cook Mountain representing a destructional phase early in a transgression over Formation (“Wahtubbee horizon”) of Clarke County to Zanthopsis the Forest Hill Delta (Coleman, 1983). The sedimentary struc- errans Woods, 1922, known from the Eocene of Peru. The Cook tures and fauna indicate high-energy deposition in shallow depths Mountain Formation of Texas has yielded a slightly greater (Tew, 1992). Abrasion of phosphatic and calcitic skeletal clasts also diversity of decapods (Stenzel, 1935; Rathbun, 1935) than in suggests a relatively high-energy system, at least compared to the Mississippi. superjacent Marianna and Glendon formations. The frequency of A single decapod has been described from the lower Lisbon Ophiomorpha-type burrows is consistent with the occurrence of the Formation (Lutetian) of Alabama. Originally reported from the numerous callianassid chelae. The Marianna Formation, however, Tallahatta Formation, the “glauconite-rich quartz sands to mica- is largely a carbonate mud, although the purer muds often alter- ceous sandy claystones” yielding the type of Costacopluma grayi nate with intervals rich in pelletal glauconite and skeletal sand. Feldmann & Portell, 2007, as well as abundant marine vertebrate Bryozoans are abundant in the Marianna Formation, particularly fossils, have since been attributed to the basal Lisbon Formation the erect cheilostome Metrarabdotos Canu, 1914. The Marianna (Clayton et al., 2013). In the lower Lisbon (Winona Formation) of Formation also abounds with large, thin, nummiform tests of the Mississippi, isolated callianassoid chelae were attributed to four benthic orbitoid foraminiferan Lepidocyclina Gümbel, 1870, par- different species by Stenzel (1935), one of which Rathbun (1935) ticularly in the glauconitic wackestone interbeds. Like most orbit- established as a subspecies of Callianassa ulrichi White, 1880. oid taxa, Lepidocyclina hosted photosynthetic endosymbionts and The bulk of the present material is from fossil-rich marine thus required relatively shallow and/or well-illuminated depths phases of the Oligocene Vicksburg Group of Mississippi, and most (Tappan & Loeblich, 1988; BouDagher-Fadel & Price, 2010). The specimens can be assigned to several species of calappid crabs Marianna Formation was largely deposited below wave base as it originating from different calcareous lithofacies. The Vicksburg lacks sedimentary structures suggestive of sustained traction cur- Group is entirely Rupelian, or early Oligocene (Mancini & Tew, rents, thus indicating relatively low-energy bottom currents (Davis 1991). The sediments of interest include (in ascending order) the & Heydari, 2015). A positive correlation between orbitoid test Mint Spring, Marianna, and Glendon formations, the first two size and water depth (Bryan, 1995) suggests the large Marianna representing a transgressive sequence and the Glendon beginning Lepidocyclina, when compared to the smaller orbitoids of the sub- the final regression of the Vicksburg Sea (Dockery, 1982; Tew, jacent Mint Spring, were possibly due to decreased light with

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Figure 2. Stratigraphic chart of select Paleogene units of the eastern Gulf Coastal Plain, USA. Integrated correlation of numerical, geochronologic and biozonal reference scales produced using TSCreator 7.3 © 2005–2018, which is based on Ogg et al. (2016), which in turn is based on a host of earlier stratigraphic contributions globally. Regional chronostratigraphic correlation with the reference columns is based on similarly numerous stratigraphic works as compiled by Mancini & Tew (1988, 1991), Tew (1992), Berggren et al. (1995), and Dockery (1996). All units are presented in terms of their surface expression only. Post-Marianna Vicksburg units are treated as formations in Mississippi, but are treated as members of the “Byram Formation” in Alabama (review by Mancini & Tew, 1991). Chart truncated twice internally (horizontal gray bars) for topical focus. greater depth, the larger size required to capture more diffuse or energy than the subjacent Marianna Formation, the Glendon weaker light. The lower frequency of orbitoids in the Mint Spring Formation represents onset of regression that produced, at least in Formation may be a function of the higher energy. The Glendon much of the Mississippi outcrop belt, a shoaling facies dense with Limestone is a bedded, glauconitic grainstone and floatstone pectinid valves, namely Pecten byramensis Gardner, 1945. populated by bivalves, including pectinids (especially toward the Four decapods and a paguroid ichnofossil previously were top), cardiids, and the deep infaunal suspension feeders Panopea described from the early Oligocene of Alabama and Mississippi. Ménard de la Groye, 1807, and Pholadomya Sowerby, 1823. Among Glypturus berryi (Rathbun, 1935) (Callianassidae), Necronectes vicks- the three Limestones, Lophoranina Fabiani, 1910 (Raninidae) and burgensis (Stenzel, 1935), and Lophoranina georgiana (Rathbun, 1935) Necronectes A. Milne-Edwards, 1881 (Portunidae), seem to occur are from the Glendon Limestone of Mississippi. Specimens only in the Glendon Formation. Generally coarser and higher identified as L. georgiana should be inspected in light of the new

281 R. M. FELDMANN ET AL. species described herein. The paratype (USNM 372809), from Oligosella Ciampaglio & Weaver, 2008 the Glendon Limestone of St. Stephens, AL, was collected much closer geographically to L. lincki n. sp. than to the type locality Type species: Oligosella longi Ciampaglio & Weaver, 2008, by original of L. georgiana, which is in southwest Georgia. Callinectes alabamen- designation. sis Rathbun, 1935 was erected for an incomplete chela reportedly from the Byram Formation of Monroe County, AL. The spionid Oligosella cf. O. longi Ciampaglio & Weaver, 2008 (Polychaeta) boring trace Helicotraphichnus Kern, Grimmer & Lister, 1974, which is a known associate of pagurized shells, was reported (Fig. 3A) by Walker (1992) in Galeodea (Mambrinia) brevidentata (Aldrich, Material examined: MMNS IP-6960 and 8795. 1885) from the early Vicksburg Red Bluff Formation. Crustaceans are previously unreported from the Catahoula Occurrence: Late Oligocene (Chattian) basal beds of the Catahoula Formation. Paralic sediments in the basal beds, however, have Formation (MMNS location MS.77.011), Wayne County, MS. recently yielded a diversity of terrestrial, paludal, estuarine, and Remarks: The claw tips here referred to Oligosella cf. O. longi exhibit shallow marine taxa at the base of a delta-building sequence the “triangular morphotype with a short hook” of Hyžný et al. (Albright et al., 2016). The caridean remains described below are (2017, 2018). This suggests that they are most likely movable thus far the only identifiable crustaceans observed in these depos- finger tips. They are quite similar to those described as Oligosella Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021 its, although degraded decapod cuticle of other, unidentifiable longi, exhibiting the same shape and a row of setal pits on the distal forms is present. margin. Oligorostra alabami Ciampaglio & Weaver, 2008, exhibits a longer hook and one large setal pit near the distal end. Material referred to Alpheus sp. have a row of setal pits on the distal margin MATERIALS AND METHODS that alternate in size between small and large (Hyžný et al., 2018). Specimens were prepared, where necessary, with PaleoTools The morphotype assigned by Hyžný et al. (2018) to females has a microjacks (Paleo Tools, Brigham City, UT, USA) under a Leica longer hook on the tip of the finger than the male morphotype; MZ6 (Leica, Wetzlar, Germany) binocular microscope. Specimens Oligosella longi exhibits the male morphotype. were whitened with ammonium chloride and photographed with a Nikon D3100 camera (Nikon, Tokyo, Japan) with an AF-S micro Infraorder Axiidea de Saint Laurent, 1979 Nikkor 60 mm lens. Images were corrected for brightness and contrast with Adobe Photoshop CC2018 software. Family Callianassidae Dana, 1852 Genus Mesostylus Bronn & Roemer, 1852 Institutional abbreviations: MMNS, Mississippi Museum of Natural Science, Jackson, MS; UF, University of Florida Museum of Type species: Pagurus faujasi Desmarest, 1822, by monotypy. Natural History, Gainesville, FL; USNM, United State National Other species: Mesostylus mortoni (Pilsbry, 1901). Museum (Smithsonian Institution), Washington, D.C. Diagnosis: See Mourik et al. (2005) and Schweitzer & Feldmann (2012).

SYSTEMATIC PALEONTOLOGY Mesostylus cf. M. mortoni (Pilsbry, 1901) Order Decapoda Latreille, 1802 (Figs. 3G, H) Infraorder Caridea Dana, 1852 Material examined: MMNS IP-7944. Superfamily Alpheoidea Rafinesque, 1815 Diagnosis: See Schweitzer & Feldmann (2012), Feldmann et al. (2013), Kornecki et al. (2017). Family Alpheidae Rafinesque, 1815 Occurrence: Paleocene (Danian) Clayton Formation, Pine Barren Included fossil genera: Alpheus Fabricius, 1798; Oligorostra Ciampaglio & Member, MMNS loc. 43.028, Lowndes County, AL. Weaver, 2008; Oligosella Ciampaglio & Weaver, 2008. Remarks: Schweitzer & Feldmann (2012) provided an overview Remarks: Hyžný et al. (2017) reviewed the occurrences of Alpheidae, of Mesostylus and M. mortoni, including synonymies, a diagnosis, commonly called snapping shrimp or pistol shrimp, in the fossil and occurrences. The species is widespread on the east and Gulf record. They are known as fossils exclusively from the well- of Mexico coasts of the United States in the Late Cretaceous calcified tips of the fingers and have a moderate fossil record. (Feldmann et al., 2013; Kornecki et al., 2017). The material here Several occurrences were originally referred to cephalopod taxa, tentatively referred to the species exhibits diagnostic characters but Hyžný et al. (2017) identified them as claw tips.Ciampaglio including a serrate upper margin of the carpus, a distinct oblique & Weaver (2008) recognized two new genera and species of what keel on the lower distal corner of the carpus, and a granular merus. are now known to be alpheid finger tips of the alpheids Oligorostra The major chela is not preserved, but the minor chela exhibits alabami Ciampaglio & Weaver, 2008 and Oligosella longi Ciampaglio long fingers as seen in Mesostylus. Although the specimen is clearly & Weaver, 2008 from Oligocene (Chattian) sediments of Alabama. referable to Mesostylus, species attribution is made difficult by the Oligorostra alabami appears to include fixed finger tips and O. longa absence of the major chela. This Danian occurrence extends the seems to be composed only of movable finger tips. They are from range of the genus into the Paleogene. Most other occurrences the same locality in the same formation and it is likely that they of Mesostylus to date are Late Cretaceous. The region in which represent the same species. Differences between finger tips in these the Clayton Formation was deposited has been suggested to be a two genera may also be a result of sexual dimorphism, recognized possible refugium, as the formation contains a high percentage of in fossil and extant Alpheus (Hyžný et al. 2018). Cretaceous survivors (Cope et al., 2005). The material referred herein is composed only of movable finger tips. They are quite similar in morphology to those of Oligosella but are slightly younger, from the Chattian Catahoula Formation Genus Neocallichirus Sakai, 1988 in Mississippi. This occurrence thus extends the range of the family only slightly. Type species: Neocallichirus horneri Sakai, 1988, by original designation.

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Figure 3. Miscellaneous cheliped fragments. Alpheidae, Oligosella cf. O. longi Ciampaglio & Weaver, 2008, three specimens from MMNS IP-6960, tips of movable fingers (A). Inner (D) and outer (B) surfaces of Neocallichirus matsoni (Rathbun, 1935), two specimens (MMNS IP-9578) (B, D). Paguridae incertae sedis, outer surface of chela (MMNS IP-9572) (C). Callianassa zeta Rathbun (1936) (MMNS IP-7255), minor? chela (E) and major? chela (F). Mesostylus mortoni (Pilsbry, 1901) (MMNS IP-7944), right major cheliped and other appendage fragments (G) and minor cheliped (H). Scale bar A = 5 mm, scale bars B-H = 1 cm.

Included fossil species: See lists in Schweitzer et al. (2010), Hyžný & Neocallichirus matsoni (Rathbun, 1935) Karasawa (2012), and Hyžný & Klompmaker (2015). (Figs. 3B, D) Remarks: Many fossil species have been placed within Neocallichirus over the past 30 years, based upon the nature of the carpus and Material examined: Two left chelae, MMNS IP-9578; two left and manus. Notably, the manus in Neocallichirus has a serrate distal one right chela, MMNS IP-9048; photographs of the holotype of margin as seen in Callianassa matsoni Rathbun, 1935. That feature Neocallichirus matsoni (USNM 371470; https://collections.nmnh. prompted Schweitzer & Feldmann (2002) to place C. matsoni si.edu/search/paleo/ under Callianassa matsoni). within Neocallichirus. This placement later was considered doubtful (Hyžný & Karasawa, 2012) but alternative placements Description of material: Manus rectangular, overall smooth; proximal have not been suggested. The problem of generic placement of margin at about 90° to upper and lower margins, appearing to axiids has been extensively discussed (summarized by Hyžný & have been sinuous; upper, lower margins nearly straight; distal Klompmaker, 2015). We therefore retain the placement for now, margin also sinuous, with long serrate projection above fixed finger, considering that the material herein referred to the species is followed by arcuate indentation in distal margin just above fixed quite fragmental. finger; outer surface smooth with few granules just proximal to

283 R. M. FELDMANN ET AL. arcuate indentation in distal margin; inner margin with rows of proximal corner; upper margin weakly concave, with double row setal pits paralleling upper, lower margins, serrate projection also of robust tubercles; lower margin nearly straight, wide, flattened, developed on inner surface of distal margin; fixed finger slightly with 3 rows of small tubercles that extend onto fixed finger; distal upturned, with broad triangular tooth at about midlength on margin oblique to upper margin, making about 100° angle, with occlusal surface, with few setal pits distally. swellings at base of movable finger; outer surface convex, 2 rows of tubercles centrally plus scattered tubercles below tubercles; inner Occurrence: Early Oligocene (Rupelian) Mint Spring Marl (MMNS surface weakly convex, with scattered, transverse short scabrous IP-9048, 9578, MMNS location MS.65.004), Smith County, MS. ridges centrally. Remarks: The new material is quite similar to the holotype of Fingers arched toward inner margin. Fixed finger with con- Neocallichirus matsoni. The holotype has a rectangular manus with spicuously strong, molariform teeth on occlusal surface, smaller the serrate extension on the distal margin and arcuate indentation at proximal end, larger at distal end, tip edentulous; outer sur- about the fixed finger as well as an upturned fixed finger as seen in face with row of setal pits below teeth, remainder of finger the referred specimens. There is not much evidence to go on, so we including lower margin with rows of tubercles. Movable finger place the material within that species. with strong molariform teeth except at edentulous tip, row of The type material for this species was collected from the setal pits parallel to occlusal surface; upper, outer surfaces with Chipola Formation, now of the Miocene Alum Bluff Group, rows of tubercles. Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021 and the late Oligocene to early Miocene Tampa Member of Occurrence: Early Oligocene (Rupelian) Mint Spring Marl (MMNS the Arcadia Formation (Scott, 2001), all in Florida. The new loc. MS.65.004), Smith County, MS. specimens were recovered from the early Oligocene Mint Spring Formation in Mississippi. Remarks: We refer the single chela described here to Paguridae, only on the basis of it being a right claw. The right claw is much Genus Callianassa Leach, 1814 sensu lato larger than the left in most species of this family. Several species of Paguroidea have been described from Cretaceous and Cenozoic Callianassa zeta Rathbun, 1936 strata of the Gulf coastal region. The material described here is similar to the type material of Pagurus alabamensis Rathbun, 1935, (Figs. 3E, F) but that species has shorter fingers, more rows of tubercles on the Callianassa zeta Rathbun, 1936: 37. outer margin of the manus, and does not show evidence of the Callianassa alpha Rathbun, 1935: 67, pl. 15, figs. 23–25, 27. Non strong molariform teeth on the occlusal surfaces of the fingers. Callianassa alpha Stenzel, 1935: 391, pl. 15, figs. 1, 2, 7, 8. Paguristes chipolensis Rathbun, 1935, is known from finger fragments which lack large molariform teeth, and this genus is referred to Material examined: MMNS IP-7255, two specimens. Diogenidae Ortmann, 1892, which generally possess stronger Description of material: Minor? chela longer than high, rectangular, left chelae than right. We therefore leave this material in open slightly smaller than major? chela; fixed finger short, with triangular nomenclature. patch of granular ornament on outer surface, occlusal surface with sharp ridge; movable finger stouter, with keel on outer surface. Pagurized gastropods Major? chela with triangular granular patch at base of fixed finger; elongated, granular swelling parallel to distal margin; fixed finger Fig. 4 arcuate, curved up towards distal end; movable finger with keel on outer surface; both fingers with setal pits on outer surfaces. Material examined: MMNS IP-8382, 9033, 9162, 9388, and 9878. Occurrence: Paleocene (Danian) Clayton Formation, Pine Barren Occurrence: All known material is from the early Oligocene Mint Member (MMNS loc. AL.43.002), Lowndes County, AL. Spring Marl, MMNS loc. MS.65.004, Smith County, MS. Remarks: Rathbun (1935) named Callianassa alpha, whereas C. alpha Remarks: There are five specimens of moderately high spired, Stenzel, 1935 was published first. Rathbun (1936) thus created fusiform gastropods bearing about six spiral striae per whorl and an C. zeta as a replacement name for C. alpha. The diagnostic feature apical angle of about 45º. Examination of illustrations in Dockery for C. zeta is the triangular patch of granules on the manus near (1977) and MacNeil & Dockery (1984) suggest a resemblance to the fixed finger, exhibited by the new material. The type material Latirus Montfort, 1810 from the upper Eocene Moodys Branch was collected from the Sucarnoochee beds, now known as the Formation in Mississippi. The shells are covered by variable Danian Porters Creek Formation. Vega et al. (2007) attributed a thickness, multilamellar, cheilostome bryozoans in a manner that portion of a manus to this species (as C. alpha Rathbun, 1935), but is interpreted to represent occupancy by pagurid hermit crabs that specimen seems to lack the triangular granular patch. More (Walker, 1992; Taylor, 1994). No remnants of the crabs remain, complete material of this taxon will be needed to place it within which is commonly the case. a family and genus, so for now we refer it to Callianassa sensu lato. Infraorder Brachyura Linnaeus, 1758 Infraorder Anomura MacLeay, 1838 Superfamily Raninoidea De Haan, 1839 Family Paguridae Latreille, 1802 Family Lyreididae Guinot, 1993 Paguridae incertae sedis Subfamily Lyreidinae Guinot, 1993 (Fig. 3C) Genus Giulianolyreidus Karasawa, Schweitzer, Material examined: MMNS IP-9572. Photographs of the holotype Feldmann & Luque, 2014 of Pagurus alabamensis, (USNM 371713), https://collections.nmnh. Type species: Symethis johnsoni Rathbun, 1935, by original designation. si.edu/search/paleo/ under Pagurus alabamensis). Other species: Giulianolyreidus bidentatus (Rathbun, 1935). Description of material: Right manus longer than high; proximal margin with articulating rim centrally, large tubercle at upper Diagnosis: See Karasawa et al. (2014).

284 PALEOGENE DECAPODA (CARIDEA, ANOMURA, AXIIDEA, BRACHYURA)

Giulianolyreidus johnsoni (Rathbun, 1935) Genus Lophoranina Fabiani, 1910

(Fig. 5) Type species: Ranina marestiana König, 1825, by original designation. Material examined: MMNS IP-7252 (two specimens); MMNS Other species: See Karasawa et al. (2014). IP-8792. Diagnosis: Carapace widest in anterior one-quarter to one-half, Diagnosis: See Karasawa et al. (2014). narrowing posteriorly; front trifid with axial single or trifid spine, lateral spines serving as inner orbital spines. Anterolateral Occurrence: Paleocene (Danian) Clayton Formation, Pine Barren margin generally with 2 prominent spines that may be bifid Member (MMNS loc. AL.43.002), Lowndes County, AL. or ornamented with granules or spinelets, possibly intervening Remarks: The type material of this species was collected from the small spines. Postfrontal region depressed slightly below level of Paleocene Sucarnoochee beds, now recognized as the Porters remainder of carapace, can be granular, scabrous, terraced, or Creek Formation (MacNeil, 1946; Renken, 1996). The specimens smooth. Carapace surface with well-developed transverse terraces; referred here were collected from the Paleocene Clayton terraces relatively narrowly spaced, usually parallel to one another Formation, Alabama. The new specimens are broken, none with anteriorly, continuous across carapace, or often interfingered with Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021 the rostrum or orbital margins. the carapace shape, narrowed one another or intersecting posteriorly; terraces ornamented frontal margin, and stout anterolateral spine, however, are all with tiny, forward directed spines; pterygostome, pleonal somites, diagnostic for the species. appendages ornamented with terraces; sternites 1–5 with scattered One specimen (Fig. 5B) has unusual damage to the dorsal cara- scabrous ornamentation (emended and modified from Karasawa pace. The damage consists of two arcuate crushed areas, which et al., 2014). are not tubes or holes. Cuticle remains in the depressed element. We are unsure of the cause of the damage but suggest that it Lophoranina lincki n. sp. could be a bite or crush mark from a predator. (Fig. 6) Family Raninidae De Haan, 1839 Types: Holotype (MMNS IP-1980), paratype (MMNS IP-7767). Subfamily Ranininae De Haan,1839 A third specimen (UF 67098) may also be referable to the species, but this is subject to confirmation. Included genera: Alcepsina Pasini & Garassino, 2017; Lophoranina Diagnosis: Carapace with 2 moderately sized, forward directed Fabiani, 1910; Lophoraninella Glaessner, 1946; Pseudoranina anterolateral spines separated by two tiny spines; surface of Charbonnier, Garassino & Hyzny, 2017; Ranina Lamarck, 1801; carapace with uniformly broadly spaced, continuous terraced lines. Raninella A. Milne-Edwards, 1862; Remyranina Schweitzer & Feldmann, 2010 (cf 2010a), Tethysranina Pasini & Garassino, 2018; Description: Carapace moderate sized for genus. Length, 43.9 mm, Vegaranina van Bakel et al., 2012. 1.15 ×maximum width, 38.1 mm, measured at midlength at level

Figure 4. Pagurized gastropod shells. Bryozoan and mold of interior of gastropod (MMNS IP-9388) (A). Gastropod encrusted with bryozoan (MMNS IP-8382) (B). Scale bars = 1 cm.

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Figure 5. Giulianolyreidus johnsoni (Rathbun, 1935) (MMNS IP-7252), two specimens. Nearly complete specimen with narrow frontal margin and large anterolateral spine diagnostic for the genus (A). Incomplete specimen with arcuate crush marks on left side of carapace (B). Scale bars = 1 cm. of second meso-urogastric terrace; weakly vaulted longitudinally, Federal Emeregency Management Agency (FEMA) who donated strongly vaulted transversely. Carapace highest at midlength when many specimens to MMNS and assisted with field work. viewed laterally. Carapace surface with strong, spinose, terraced Occurrence: Early Oligocene (Rupelian) Glendon Limestone (MMNS ridges developed on entire surface. IP-1980, 7767, MMNS loc. MS.65.004), Smith County, MS. Front (9.6 mm) about 25% maximum width, with poorly preserved, short, forward directed triangular axial rostrum, ante- Remarks: Four genera within Ranininae, Lophoranina, Lophoraninella, rolaterally directed lateral inner orbital spines. Outer orbital Remyranina, and Vegaranina, exhibit prominent terraced lines on spine small, extending anteriorly from rectangular surface the dorsal carapace. The genera can nevertheless be readily bounded axially by small spine; 2 broad, open fissures; inner fis- distinguished from one another by the nature and distribution sure inclined axially below inner orbital spine, outer fissure at of the terraces. Lophoranina, to which the new species is assigned, orbital midwidth. Anterolateral margin smoothly convex with bears terraced structures over nearly the entire dorsal surface; anterolaterally directed, strong spine at level of frontal terrace, only a narrow frontal rim may be devoid of the structures. 2 tiny spines at anterior gastric terrace level, strong spine at mid Lophoraninella and Remyranina have terraced structures only on length of gastric region. Posterolateral margin nearly straight, the posterior half of the carapace. The anterior half of the bounded by narrow, beaded rim. Posterior margin poorly pre- carapace on Lophoraninella is poorly preserved but seems to bear served, narrow. scalloped structures on an otherwise smooth surface, and that of Regions primarily defined by variations in configuration of -ter Remyranina is finely granular. The anterior half of the carapace raced lines (Fig. 6A). Frontal region with biconcave forward ter- of Vegaranina has a broad, granular frontal region followed by race with axial apex plus about 6 spine bases regularly spaced ovoid to irregularly shaped, terraced structures. Based upon these plus 2 smaller, more widely spaced spine bases laterally. Three points of distinction alone, the new species can be assigned to sinusoidally curved, continuous terraces define gastric, hepatic Lophoranina with confidence. regions. Three short terraces in meso-urogastric region separated Regional development in Lophoranina is indistinct, and distinc- from convex forward terraces in epibranchial region by longitu- tion of regions is based only upon a frontal surface that may be dinal branchiocardiac groove. Six continuous, concave forward depressed below the remainder of the carapace or differently terraces define branchial region. Branchial terraces more closely ornamented (Fig. 6A). The level of the mesogastric and urogastric spaced (2.8 mm) than gastric terraces (3.0 m). About 14 spinel- regions is defined by the presence of short, longitudinal expres- ets/cm arise from terraces throughout carapace; spinelets parallel sions of the branchiocardiac grooves. The gastric region lies ante- to carapace surface, extending at least halfway across intervening rior to the branchiocardiac grooves and posterior to the frontal space between terraces. region. The branchial region is situated posterior to the branchio- Venter, pleon, appendages not preserved. cardiac grooves. Because terraced structures are so distinctive and so variable Etymology: The trivial name derives from the family name of the within species of Lophoranina, it became apparent that there discoverer of the specimen, Dana Linck, an archeologist from the was a clear regional pattern that made comparison of species

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Figure 6. Lophoranina lincki n. sp., holotype (MMNS IP-1980). Dorsal carapace with regions delimited (A). Close-up of anterolateral margin, arrows indicating tiny anterolateral spines (B); oblique left view, showing tiny spines on terraces (C). Scale bars = 1 cm. possible. The frontal region, extending to the level just posterior The pattern of terraced structures differs in spacing and con- to the outer orbital spine, may lack terraces or have terraces tinuity. Examination of specimens and published illustrations of with a configuration different from those in the gastric regions. all the named species of Lophoranina permits a subjective evalu- The meta-urogastric level, approximately the middle of the car- ation of spacing. Spacing can be judged as “wide” when the apace, has terraced structures interrupted by the branchiocar- intervening spaces between the terraced ridges are as broad, or diac grooves. Development of terraces in the axial region could broader, than the ridges, and “narrow” when there is only a short be similar to the lateral areas, considered to be epibranchial distance between ridges. Spacing of terraced lines may be the regions, or could differ. The branchial region may bear ter- same throughout the carapace or may vary in different regions. races of a configuration different from those in more anterior Continuity of terraces ranges from those that are “continuous” positions. across the breadth of the carapace, “discontinuous” with each

287 R. M. FELDMANN ET AL. terrace extending less than the breadth of the carapace, and Included fossil species: Notopus adinae n. sp.; N. dorsipes (Linnaeus, 1758); “intercalated,” with discontinuous terraces extending axially from N. minutus Vega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer the margin. As with spacing, continuity may differ from one region & Waugh, 2001. to another and forms an important basis for defining species. Diagnosis: See Karasawa et al. (2014). Application of these criteria to specimens in the literature, cou- pled with variations in the morphology of the rostrum, orbits, and anterolateral margin, indicate that a re-evaluation of the genus is Notopus adinae n. sp. warranted; however, that is beyond the scope of this work. Several key morphological characters distinguish species (Figs. 7, 8) within Lophoranina, and L. lincki n. sp. exhibits a unique plexus of characters. The frontal margin including the orbital region Types: Holotype (MMNS IP-2324), paratypes (MMNS IP-7124, has an apparently simple, triangular rostrum, inner and outer 4988, 6316, and 8915). orbital spines of similar size, and well-developed, broadly open, Diagnosis: Carapace with inner and outer margins of orbits defined orbital fissures. These features most closely resemble those of by small, sharp spines; short, granular terraced lines on the L. bittneri (Lőrenthey, 1902), although the rostra of the two spe- anterolateral and posterolateral margins; transverse ridge separating cies are only partially preserved. The anterolateral margin of frontal region from remainder of carapace surface confined to short Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021 L. lincki n. sp. bears two slender, anteriorly directed spines, one granular ridges along posterior margin of hepatic regions. at the level of the frontal terrace and one at the midlength of the gastric region. Two minute spines separate the larger spines Description: Carapace uniformly wide anteriorly, tapering posteriorly along the margin (Fig. 6B). Although it is difficult to make com- to narrow posterior margin (Fig. 7A). Nearly flat longitudinally (Fig. parisons with certainty because of preservational differences, 7B), strongly vaulted transversely; holotype longer (20.8 mm) than several species have much stronger anterolateral spines and none wide (14.7 mm) with greatest width in anterior half. Front about 56% has the two minute spines preserved. More detailed comparison maximum width. Fronto-orbital width about 82% maximum width. of this character must await examination of more specimens Rostrum triangular, tip needle-like, extending onto dorsal carapace and reconsideration of species placement. The spacing of ter- as keel anteriorly (Fig. 7C). Orbits bounded by sharp inner, outer raced lines in L. lincki n. sp. is wide and uniform throughout the orbital spines. Upper orbital margin finely spinose, with closed orbital length of the carapace. Although some species have narrow, uni- fissure at base of inner orbital spine (Fig. 7D). Anterolateral margin formly spaced terraces, only L. bittneri and L. georgiana (Rathbun, widens from outer orbital spine to long (ca. 4 mm), narrow, sharp, 1935) are spaced as in L. lincki n. sp. Continuity of terraced lines anterolateral spine directed anterolaterally (Fig. 7E, F). Remainder clearly sets L. lincki n. sp. apart from nearly all other species of anterolateral margin parallels midline, with short, granular within the genus. The only other species that has continuous terraced lines. Posterolateral margin arises at midlength of carapace, terraces throughout is L. avesana (Bittner, 1883); however, that tapering with nearly straight margin to narrow (6.9 mm) posterior species is widest at the anterior margin, bears extremely strong margin. Posterior, posterolateral margins rimmed. Posterolateral rim anterolateral spines, and has terrace spacing that narrows poste- finely granular below, arranged as 12 short, granular, terraced lines riorly. Lophoranina lincki n. sp. is thus unique. inclined posteroventrally. Posterior rim smooth. Lophoranina lincki n. sp. has been identified previously asL. geor - Surface of carapace with subtle axial ridge, weakly depressed giana, and a specimen, (UF 67098), has been placed in the lat- hepatic regions, plus smooth, arcuate branchiocardiac grooves. ter species. Examination of illustrations of that specimen suggests Frontal region separated from remainder of carapace only near that possession of broadly spaced, continuous terraced lines and lateral margins where denticulate ridge defines the posterior mar- two, moderate sized, forward-directed anterolateral spines place it gin of the hepatic region. Regions otherwise indistinct. Surface within L. lincki n. sp. By contrast, the terraced lines on L. georgiana of exocuticle wrinkled, punctate. Punctae smaller on surface of are discontinuous throughout most of the length of the carapace, endocuticle. as seen on the holotype (USNM 371714). Left major chela (Fig. 8) with hand wider than long, compressed; The apex of the terraced lines on all species is punctuated upper margin finely granular, describing broadly arcuate surface by small pits. In well-preserved specimens, small spines can be from point of articulation with dactylus to concave proximal articu- observed to emanate from the pits and to project over the interven- lation with carpus. Lower surface weakly sinuous terminating dis- ing spaces almost parallel to the surface (Fig. 6C). The spines have tally in smooth, curved, fixed finger with acute termination. Upper been broken off during their postmortem history in most cases, surface bearing short, finely digitate, terraced lines arrayed in con- but some are preserved on a few specimens, including on L. lincki cave distally arcs covering all but uppermost part of surface. Distal n. sp. Classic studies on terraced lines by Savazzi (1982, 1985) margin between fixed finger and dactylus with 3 small nodes, irreg- interpreted the structures as serving as gripping devices, much like ularly elevated occlusal surface. Lower surface punctate. Dactylus the structure of a carpenter’s rasp, for pressing against the surface smoothly arcuate with acute tip with long axis nearly at right angles of a burrow to avoid extraction by a predator. Presence of the to long axis of hand, nearly as long as width of hand, smoothly spines would seem to obviate this function, and it was suggested arcuate on outer surface with 2 small denticles on occlusal surface. previously (Feldmann et al., 1996) that the spines might function to Outer surface with subtle keel; inner surface smooth. snag seagrass or other marine vegetation to serve as camouflage. Sternum, pleon, appendages not preserved.

Nomenclatural statement: A life science identifier (LSID) num- Etymology: The trivial name recognizes Dr. Adina Franţescu, who ber was obtained for the new species: urn:lsid:zoobank. studied the specimen and concluded it was not assignable to any org:pub:8ADA5F9E-EF00-4B61-AF43-F8235F6D3A08. species known at that time. Occurrence: Holotype (MMNS IP-2324), Oligocene (Rupelian) Glendon Limestone (MMNS loc. MS.65.004), Smith County, Subfamily Notopodinae Serène & Umali, 1972 MS. Paratype (MMNS IP-4988), Oligocene (Rupelian) Glendon/ Marianna Limestone (MMNS loc. MS.65.003), Smith County. Genus Notopus De Haan, 1841 Paratypes (MMNS IP-8915), Early Oligocene (Rupelian) Mint Spring Marl, and IP-6316, Glendon/Marianna Limestone (MMNS Type species: Cancer dorsipes Linnaeus, 1758, by monotypy. loc. MS.65.004), Smith County, MS.

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Remarks: Placement in Notopus is based upon presence of a weak (Fig. 9); however, the frontal ridge is much better developed and longitudinal keel, orbits with inner and outer orbital spines, extends across the entire width of the carapace. Comparison of a single, closed orbital fissure at the base of the inner orbital N. adinae n. sp. with the Eocene N. minutus is not possible because spine, and a serrated or spinose upper orbital margin. A long the latter is eroded so that the cuticle is not exposed, and the front spine set close to the frontal margin and anterolateral margins of the specimen is missing altogether. parallel to the midline further confirm generic placement. The short, terraced lines preserved on the outer surface of the The only exception to the diagnosis of Karasawa et al. (2014) single chela mimics those on the carapace (Fig. 8), suggesting that, is in regard to the degree of development of the frontal although the chela was not intimately associated with the carapace ridge which is developed only near the lateral margins of the material, the chela can be assigned to N. adinae n. sp. The over- carapace. all form of the chela is the typical notopine-type chela noted by The distinctive development of short terraced lines along the Karasawa et al. (2014). lateral margins of the carapace and the development of the frontal ridge serve to distinguish N. adinae n. sp. from other fossil Nomenclatural statement: A life science identifier (LSID) num- forms. Extant specimens of N. dorsipes have orbital morphology ber was obtained for the new species: urn:lsid:zoobank. and overall outline that is consistent with those of the new species org:pub:05EA25A6-53D3-4DBC-95AE-0ECEA572D14B. Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021

Figure 7. Notopus adinae n. sp., holotype (A, B, D–F) (MMNS IP-2324). Dorsal carapace (A); right lateral view (B), left anterolateral and outer-orbital region (D), right anterolateral and outer-orbital region (E), oblique anterior view showing terraced region of frontal ridge (F). Paratype (MMNS IP-7124), anterior view (C). Scale bars A–C = 1 cm; scale bars D–F, 5 mm.

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Figure 8. Notopus adinae n. sp., paratype (MMNS IP-6316). Outer surface of left chela (A); inner surface of left chela (B). Scale bars = 5 mm.

Section Eubrachyura de Saint Laurent, 1980 moderately large node is positioned posterior to it on the branchial region. A concave forward row of granules crosses the posterior Subsection Heterotremata Guinot, 1977 part of the intestinal region and the posterior and posterolateral Superfamily Calappoidea De Haan, 1833 margin exhibits a similar row of nodes. The posterolateral margin is flared out in a posterolateral direction and bears at least three Family Calappidae De Haan, 1833 prominent spines, the first two of which are longest. Although the partial specimens bear sufficient details of -mor Genus Calappa Weber, 1795 phology to place them within Calappa, comparison with previously named species is difficult. A search of literature on North and Type species: Cancer granulatus Linnaeus, 1758, by subsequent Central American fossil species indicates that most are based upon designation of Latreille (1810). claws, primarily propodi. The nature of the flared posterolateral Included fossil species: See Schweitzer, Feldmann, Garassino, Karasawa margin and the relatively long intestinal region is reminiscent of & Schweigert, 2010. Calappa ocellata Holthuis, 1958, an extant form from Curaçao; however, that species lacks the very prominent nodes seen on the Diagnosis: Carapace ovate, wider than long, widest near posterior Mint Spring form. Given the incomplete nature of the specimens margin of carapace; front narrow, triangular; orbits directed and the general lack of comparative material from the continent, forward; anterolateral margin arcuate and crenulate, dentate, or it is prudent to assign the specimens to Calappa sp. granular; posterolateral margin with spined posterolateral flange: Most of the specimens referred to Calappidae in the Mint carapace regions poorly defined; axial regions best defined of all Spring Formation can be confidently assigned to Calappilia, species regions; carapace ornamented with large tubercles often arranged of which do not possess a flared posterolateral margin. Rather, the into rows; chelae stout, narrowing proximally. spinose margin arises directly from the edge of the carapace in Calappilia. Species of Calappilia tend to be more densely nodose Calappa sp. than species of Calappa. Species of Calappilia also are typically widest at about midlength, whereas Calappa attains its greatest width (Fig. 10) in the posterior half of the carapace.

Material examined: The illustrated specimen (Fig. 10; MMNS IP-9329) and an unillustrated specimen, (MMNS IP-8822). Genus Calappilia A. Milne-Edwards in de Bouillé, 1873

Occurrence: Oligocene (Rupelian) Mint Spring Marl (MMNS loc. Type species: Calappilia verrucosa A. Milne-Edwards in de Bouillé, MS.65.004), Smith County, MS. 1873, by subsequent designation of Glaessner (1929). Remarks: One specimen represented by the posterior two-thirds of Included species: See Schweitzer, Feldmann, Garassino, Karasawa & the carapace, the posterolateral margin, and the posterior margin, Schweigert, 2010, and C. calculosa Rumsey, Klompmaker & Portell, and another specimen of only the posteriormost posterolateral 2016; Calappilia gableorum n. sp.; Calappilia granulosa n. sp.; C. minuta and posterior margin can confidently be referred to Calappidae Beschin, Busulini, & Tessier, 2013; and Calappilia perreaulti n. sp. and most probably to Calappa. The preserved axial region consists of the posterior part of the mesogastric region along with the Diagnosis: Carapace nearly circular, widest at midlength or anterior urogastric and cardiac regions and bears four prominent nodes half; orbits small, circular directed forward or slightly upward; decreasing in size posteriorly. The posteriormost node is situated anterolateral margin finely spinose, posterolateral margin with on the weakly elevated cardiac region and is separated from a several prominent spines with the largest spines near the anterior, larger node, interpreted to be on the metabranchial region. The and lacking a projecting flange; posterior margin with axial and intervening nodes define the mesogastric and urogastric regions. lateral spines. Axial regions of carapace well differentiated, The intestinal region is long, parallel sided and lacking nodes. margins defined by prominent grooves. Entire surface with nodes The epibranchial? region bears a very large node, and a single of varying sizes.

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Figure 10. Calappa sp. (MMNS IP-9392), posterior dorsal view. Scale bar = 5 mm.

species distinguish them from the Oligocene species described herein. Calappilia diglypta, from the Texas Eocene, possesses only two nodes on the axial regions, a mesogastric node and a cardiac node. The lateral regions bear only about six nodes and numerous granules; the latter are denser posteriorly. This species possesses the fewest nodes of any North American form, which serves to distinguish it from the Mississippi species from the Oligocene Figure 9. Notopus dorsipes (Linnaeus, 1758), specimen from Senckenberg that are described below. Calappilia hondoensis from the Eocene of Forschungsinstitut und Naturmuseum, Frankfurt. Scale bar = 1 cm. Mexico has a distinctive axial region. The mesogastric and protogastric regions are bounded by deep, anteriorly-diverging, straight Remarks: Busulini et al. (2014) provided a comprehensive summary grooves, and the surface of all axial regions bear numerous very of the species of Calappilia from Europe, and noted that C. mainii small nodes as opposed to the large nodes characterizing all other Allasinaz, 1987 exhibited characters that more closely allied it North American species. The anterolateral and posterolateral with species of Stenodromia A. Milne-Edwards in de Bouillé, 1873. margins are rimmed by a discrete row of very fine nodes. We agree based on the presence of a generally smooth carapace and on the configuration of the posterolateral and posterior spines which are very large as compared with those of Calappilia. Busulini Calappilia granulosa n. sp. et al. (2014) further suggested that C. sexdentata was a nomen nudum. Although A. Milne-Edwards did not illustrate a specimen in the (Fig. 11) original work, there was a clear indication of the intent to establish a new species, a statement that it contrasted with the type species Types: Holotype (MMNS IP-1135), paratypes (MMNS IP-119, 299, of Calappilia, and a clear description (A. Milne-Edwards in de 1337a, 1337b, 2325, and 8916). Bouillé, 1876: 34). This combination of statements fully conforms Diagnosis: Coarsely nodose Calappilia with 4 prominent axial nodes; to the standard for naming a new species in the fourth edition of anterolateral margin distinctly granular with about 6 small spines; the International Code of Zoological Nomenclature (1999). We posterolateral margin with 5 prominent, granular spines with therefore retain C. sexdentata as valid. fourth spine largest. Five species have so far been recognized from the Eocene of North America, Calappilia brooksi, C. calculosa, C. diglypta, C. hon- Description: Carapace wider than long, moderately vaulted doensis, and C. sitzi. Another taxon, originally identified as transversely, strongly vaulted longitudinally. Axial regions well Aparnocondylus ocalanus (Ross, Lewis & Scolaro, 1964) and sub- defined, lateral regions not differentiated. Surface coarsely nodose sequently reassigned to Calappilia by Lewis (1969), has recently and densely granulose. been reassigned to Calappa (Rumsey et al., 2016). Among the Front narrow, about 9% maximum width measured at, or just species, all but C. calculosa can be readily distinguished from the anterior to, midline, downturned slightly, sulcate, with broadly above mentioned North American species by having a carapace triangular termination. Fronto-orbital width 26–35% maximum ornamentation dominated by small nodular structures. Calappilia width. Orbits directed forward, slightly upward; orbital rim finely brooksi and C. sitzi are quite similar to one another, but they can beaded, with 2 narrow fissures, one at midlength, one adaxial to be distinguished by several features. Calappilia brooksi bears five or it. Anterolateral margin convex, becoming more so posteriorly, six prominent blunt protuberances on the posterolateral margin finely granular with up to 8 small bluntly triangular spines, 2 of that decrease in size posteriorly, whereas the blunt protuberances which are situated just posterior to orbits, 6 in posterior half of on C. sitzi are less pronounced and seem to be largest in the mid- anterolateral margin. Posterolateral margin with 5 granular spines; dle of the series. The mesogastric node is elongated in C. sitzi but first, second shorter than third; fourth longer, stouter; fifth largest. round in C. brooksi. Although the fronts on specimens of the two Posterior margin apparently straight, 26–29% maximum width, species are not well preserved, that of C. sitzi appears to be more bounded laterally by blunt nodes. extended beyond the orbits than the front on C. brooksi. It should Overall surface of exocuticle granular; surface of endocuticle also be noted that the diagnostic features observed on these two more finely granular.

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Regions poorly defined. Frontal region smooth. Cervical Calappilia perreaulti n. sp. groove crosses midline just posterior to prominent mesogastric node, extending anterolaterally to anterolateral margin near (Fig. 12) outer orbital corner. Mesogastric and protogastric regions with field of about 9 small nodes anterior to large node. Metagastric, Types: Holotype (MMNS IP-9944), paratype (MMNS IP-4985). urogastric, cardiac regions bounded laterally by well-defined, Diagnosis: Carapace transversely ovoid, moderately vaulted sinuous branchiocardiac groove with transverse crenulations transversely, longitudinally; anterolateral margin with small, reflecting muscle or other tissue attachment sites. Metagastric upturned spines; posterolateral margin with 6 sharply pointed region with large node; urogastric node smaller. Cardiac node spines; carapace surface uniformly arrayed with moderately sized large, centrally positioned near anterior of elongate oval region. nodes and relatively few granules. Intestinal region with small, transverse ridge. Lateral regions of carapace with about 10 prominent nodes plus several smaller Description: Carapace wider (41.4 mm) than long (34.2 mm); ones. Nodes moderately large in epibranchial region, largest maximum width at midlength; moderately convex in cross section on mesobranchial region, becoming smaller on metabranchial transversely, strongly convex longitudinally, more so anteriorly. region. Axial regions well defined; lateral regions not differentiated. Sternum, pleon, appendages not preserved. Surface with numerous small, moderately sized nodes on an overall Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021 granular surface. Etymology: The trivial name refers to the distinctly granulose nature Front with downturned, sulcate, bifid rostrum, frontal width of the carapace margins and the surface of the carapace. (ca. 4.2 mm) 10% maximum width; orbits ovoid when viewed Measurements: Measurements given in Table 1. from front with long axis of orbits directed ventro-distally; fronto- orbital margin (ca. 15.7 mm) 38% maximum width. Orbital Occurrence: All known material from the Oligocene (Rupelian) margin upturned, with 2 closed fissures, one centrally placed, Vicksburg Group of Mississippi, including the Glendon Limestone second lateral to it. Anterolateral margin convex, becoming more (MMNS loc. MS.61.001), Rankin County (MMNS IP-1135); the so posteriorly, with about 10 weakly upturned, flattened spines. Marianna Limestone (loc. MS.31.001), Jasper County (IP-299); the Posterolateral margin with about 6 flattened spines directed Glendon/Marianna Limestone (IP-1337), Glendon Limestone (IP- outward, increasing in size posteriorly to fifth spine, with sixth 2325), and Marianna Limestone (IP-119, 8916), loc. MS 65.004, spine smaller. Posterior margin (16.6 mm wide), weakly convex, Smith County, MS. depressed, with smooth margin. Remarks: Calappilia granulosa n. sp. conforms to the definition Mesogastric region broadens anteriorly, bounded by distinct, of the genus in all regards. Presence of the spinose or lobose broad cervical groove passing around prominent posterior mes- posterolateral margin without the projected flange characteristic ogastric node, terminating near outer orbital corner. Anterior of Calappa, and the presence of a carapace surface lacking clear part of mesogastric and protogastric regions undifferentiated, definition of regions and possessing numerous small or large nodes with 10 small nodes arrayed in 2 transverse rows of 4 nodes are perhaps the most distinguishing features of the genus. Calappilia anteriorly, 2 nodes in advance of large node. Remaining axial calculosa is morphologically most similar to C. granulosa n. sp. in regions bounded by deep, sinuous, branchiocardiac groove. that the carapace of both species has a large number of large nodes Metagastric region with moderately large node. Urogastric and relatively fewer small nodes. The front of C. calculosa, however, node smaller than metagastric region. Cardiac region with field is depressed well below the surface of the gastric regions and the of about 5 small nodes. Intestinal region long, as wide as uro- orbits are directed forward, whereas the front on C. granulosa n. gastric region, with 3 small nodes along axis. Lateral regions sp. lies at the same level as the gastric regions and the orbits are with numerous small to moderately sized nodes, largest at directed forward and upward. The lateral margins of C. granulosa n. about midlength, decreasing in size anteriorly, posteriorly. Field sp. are densely granular as are the posterolateral spines. Calappilia between nodes finely granular. calculosa has three posterolateral spines and C. granulosa n. sp. has Sternum, pleon, appendages not preserved. five. The nodes on the branchial regions on C. calculosa are arrayed in parallel lines with coarse nodes in the more axial line and smaller Etymology: The trivial name acknowledges Ray Perreault, who nodes in the abaxial line. This pattern is not observable on the new collected the holotype specimen and permitted it to be deposited species. in MMNS. Specimens assigned to C. granulosa n. sp. are preserved in dif- Occurrence: All known material from the Oligocene (Rupelian) ferent ways reflective of different degrees of degradation. Four Vicksburg Group of Mississippi, including the Mint Spring Marl specimens (MMNS IP-299 (Fig. 11B), 1135 (Fig. 11A), 1137a (Fig. (MMNS loc. MS.61.003), Rankin County (MMNS IP-9944), 11C), and 1337b) have exocuticle and endocuticle preserved. The and the Glendon/Marianna Limestone (loc. MS.77.006), Wayne exocuticle has moderately coarse granules concentrated on the County (IP-4985), MS. nodes and fewer moderately coarse granules on the intervening carapace surface. This same array of granules is evident on the Remarks: The development of small, upturned spines on the surface of the endocuticle, where exposed; however, the endocu- anterolateral margin and sharp, horizontally directed spines on the ticular granules are less well developed. The cuticle on MMNS posterolateral margin of C. perreaulti n. sp. are distinctive among IP-2325 (Fig. 11D) has been etched and degraded so that struc- the North American species of Calappilia. The distribution of nodes tural details are less evident, but some nodes and a few gran- on the carapace and the overall outline of the carapace further ules are still visible. The surface of the mold on the interior of distinguish this species. The carapace nodes are small to medium the cuticle is exposed and appears to be relatively smooth. The sized and are more strongly developed on the anterior half than overall appearance of the ornamentation on C granulosa n. sp. on the posterior. The carapace is transversely ovate, much wider would therefore be quite different based upon which surface was than long. observed.

Nomenclatural statement: A life science identifier (LSID) num- Nomenclatural statement: A life science identifier (LSID) ber was obtained for the new species: urn:lsid:zoobank. number was obtained for the new species: urn:lsid:zoobank. org:pub:732BD6F9-276A-4187-AFED-C03F4BCAACC8. org:pub:3B06F535-AEBF-4640-933B-AD392B01CEA1.

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Figure 11. Calappilia granulosa n. sp, holotype (MMNS IP-1135) (A). Paratype (MMNS IP-299) (B); paratype (MMNS IP-1337a), (C); paratype (MMNS IP-2325) (D). Scale bars = 5 mm.

Table 1. Measurements (in mm) taken on the dorsal carapace of Calappilia granulosa n. sp. All specimen numbers are prefixed MMNS IP.

Specimen number Carapace length Carapace width Frontal width Fronto-orbital width Posterior width

1135 (holotype) 30.9 42.6 - 14.8 - 1137a - 42.8 - - - 1137b 18.1 19.9 - 6.0 5.7 299 28.0 33.3 3.0 11. 6 8.7 8162 14.9 22.5 - - -

Calappilia gableorum n. sp. Front narrow, 3.7 mm, weakly sulcate axially with small, upturned spine at tip plus 2 larger, anterolaterally directed spines at the inner (Fig. 13A-D) orbital corner. Orbits ovoid, weakly upturned, fronto-orbital width 9.9 mm. Upper orbital margin rimmed, with 2 closed fissures. Types: Holotype (MMNS IP-8375), paratypes (MMNS IP-9383, Anterolateral margin convex, arising below level of orbits; finely 9384, 9750 and 9832 (two specimens)). granular flank lies below anterolateral margin, visible in lateral Diagnosis: Front projected beyond orbits; posterolateral margin with view; anterolateral margin with about 8 minute nodes separated by 2 or 3 small spines plus elongated spine near posterior margin; 3 fine spines resulting in serrated appearance. Posterolateral margin prominent posterior spines. Surface of carapace densely granular. with 2 prominent, outward-directed spines, the posteriormost spine longest (Fig. 13A). Posterior margin with 2 lateral spines plus axial Description: Carapace ovoid, wider, 26.0 mm, than long, 24.4 mm spine; posterior width less than fronto-orbital width (Fig. 13A). (width measured at midlength on holotype); moderately vaulted Mesogastric, frontal, and protogastric regions bounded by transversely, more strongly vaulted longitudinally, particularly in V-shaped, sinuous cervical groove terminating laterally just posterior posterior portion. Axial regions well defined, lateral regions weakly to outer orbital corner. Frontal region without nodes. Mesogastric distinguished. Surface with moderate, small sized nodes separated region with large axial node posteriorly plus 2 small nodes anterior to by granular field. it. Protogastric region with 3 nodes, the posteriormost node largest.

293 R. M. FELDMANN ET AL.

Occurrence: All known material from the Oligocene (Rupelian) Mint Spring Marl (MMNS loc. MS.65.004), Smith County, MS. Remarks: This species from the Mint Spring Formation bears a conspicuously large mesogastric node and relatively broad metagastric and urogastric regions so that the branchiocardiac groove extends posteriorly from the position of the indistinct cervical groove to the intestinal region in a nearly straight line. Other species of Calappilia have a branchiocardiac groove in the shape of an hourglass. One specimen has a small bryozoan colony on the left posterolateral margin (Fig. 13B).

Nomenclatural statement: A life science identifier (LSID) number was obtained for the new species: urn:lsid:zoobank.

org:pub:ED2EADC4-5DF9-439D-A530-38C167E2A0C2. Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021

Calappilia calculosa Rumsey, Klompmaker & Portell, 2016 (Fig. 13E)

Material examined: The sole specimen from Mississippi (MMNS IP-9194). Diagnosis: “Carapace length/width ratio 0.9–1.0. Five parallel rows of nodes cross carapace from posterior to anterior. Three prominent spines lie along posterolateral margin, posteriormost strongest. Carapace widest just anterior to midpoint. Posterior margin protruding, angular at corner, flat.” (Rumsey et al., 2016: 331). Occurrence: Oligocene (Rupelian) Mint Spring Marl (MMNS loc. MS.65.004), Smith County, MS. Remarks: A single specimen from the Mint Spring Formation differs markedly from all other species of Calappilia collected from the Oligocene of Mississippi. Comparison with previously published specimens of Calappilia strongly supports placement within Calappilia calculosa. The Mint Spring specimen has an etched cuticle so that the details of surface expression are obscure. The specimen nevertheless has a length/width ratio of 0.91, the maximum width is in the anterior half of the specimen, and there are three prominent rows of longitudinally arranged nodes and an outermost pair of nodes of reduced size, also arranged longitudinally. There are a pair of prominent postero-lateral spines near the posterior margin and a large spine on the posterior margin. The carapace tapers markedly toward the narrow posterior which is one of the most distinguishing features of C. calculosa (Rumsey et al., 2016). Comparison of the features of the Mint Spring specimen and the diagnosis of the species given above strongly support assignment of the specimen to C. calculosa. Although the specimen exhibits an etched surface, it is well- enough preserved that fragments of a bryozoan colony are preserved on the axial region (Fig. 13E). Tashman et al. (2018) Figure 12. Calappilia perreaulti n. sp., holotype (MMNS IP-9944) (A), examined the grooming potential of brachyurans, demonstrating oblique anterior view of holotype (B). Paratype (MMNS IP-4985) (C). Scale that the axial regions tend to have far less potential for grooming bars = 5 mm. than do the frontal and branchial regions. Although the surface of C. calculosa is etched and some of the colony could well have been lost, the site of attachment of the bryozoans is therefore consistent Metagastric region narrow, with small node. Urogastric region nar- with the conclusions regarding epibiont location of Tashman et al. row, granular. Cardiac region inverted pentagonal with corners (2018). defined by 5 small nodes surrounding large central node. Intestinal region narrow, short, depressed, with 2 small nodes. Lateral regions ill-defined with nodes generally increasing in size posteriorly to attain Genus Acanthocarpus Stimpson, 1871 greatest size in undifferentiated branchial region. Sternum, pleon, appendages not preserved. Type species: Acanthocarpus alexandri Stimpson, 1871, by monotypy. Included species: Acanthocarpus alexandri Stimpson, 1871; A. bispinosus Etymology: The trivial name honors the Brian Gable family who has A. Milne-Edwards, 1880; A. brevispinosus Monod, 1946; A. delsolari allowed MMNS access to Smith County Lime Plant (MS.65.004) Garth, 1973; A. meridionalis Mañé-Garzón, 1980; A. obscurus for many years. (Rathbun, 1918, as Mursia obscura); A. osborni n. sp.

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Figure 13. Calappilia gableorum n. sp., holotype (MMNS IP-8375) (A); paratype (MMNS IP-9750), note bryozoan on left posterolateral margin (B); holotype, right lateral view (C); paratype (MMNS IP-9832) (D). Calappilia calculosa (Rumsey, Klompmaker & Portell, 2016) (MMNS IP-9194), arrows indicate bryozoans (E). Scale bars = 1 cm.

Diagnosis: Surface of carapace not strongly nodose; regions poorly Anterolateral margin smoothly convex, beaded rim arising from defined except metagastric, urogastric, cardiac regions, which are base of orbit above which is a weakly concave surface bounded bounded my branchiocardiac grooves (also see Rathbun, 1937). dorsally by subtle crest also arising from base of orbit, with about 3 small spines anteriorly grading into beaded crest posteriorly. Acanthocarpus osborni n. sp. Posterolateral margin beaded anteriorly; large spine directed dorsolaterally near posterior end of margin. Second larger, dor- (Fig. 14) solaterally directed spine defines posterolateral corner. Posterior margin poorly exposed, possibly straight, smooth. Holotype: MMNS IP-3780. Axial regions of carapace bounded by broad depression anteriorly, narrow, deep branchiocardiac groove posteriorly. Mesogastric Diagnosis: Carapace with well-defined axial regions bearing region poorly defined, triangular, bounded at widest point by rounded nodes; branchial regions with broadly rounded nodes; deep pits aligned transversely with prominent node. Two smaller posterolateral margin with 2 spines, the posteriormost largest. pits situated nearer midline define position where subtle cervical Description: Carapace ovoid, wider (21.6 mm), than long (> groove crosses midline. Cervical groove curves in convex forward 15.0 mm); regions defined by nodose swellings; posterolateral arc through deep pits to anterolateral margin. Metagastric region margin with 2 long spines; surface of carapace generally smooth. with moderately large node. Urogastric region arcuate, depressed. Front narrow, broken, 15% maximum width measured just ante- Cardiac region circular with prominent central node. Intestinal rior to midlength. Fronto-orbital margin 41% maximum width. region narrow, depressed. Protogastric, hepatic regions defined by Orbits transversely ovoid, directed anteriorly, slightly upward broad swellings. Branchial regions with 2 rows of about 4 nodes (Fig. 14A); upper orbital rim with 2 short, closed fissures, one at each; more axial nodes largest; lateral row with smaller nodes ter- midlength and second between midlength and outer margin. minating posteriorly in posterolateral spine.

295 R. M. FELDMANN ET AL.

Occurrence: Oligocene (Rupelian) Marianna Limestone (MMNS loc. MS.65.004), Smith County, MS. Remarks: Although placement within Calappidae is certain, assignment to Acanthocarpus is done with some reservation. Species of Acanthocarpus are diagnosed as having a posterolateral margin with one prominent spine (Rathbun, 1937). The new species has a prominent spine near the posterolateral corner and a second spine anterior to it. This latter spine is broken, so that it is not possible to be certain of its original length. Other species assigned to the genus exhibit variation in development of the posterolateral spines, as in A. bispinosus, which exhibits four or five small spines posterior to the large spine. Acanthocarpus delsolari has one short spine anterior to the large spine, which is situated near the posterolateral corner. It thus appears that the

spine arrangement of A. osborni n. sp. falls within the range of Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021 variation of the genus. In contrast, among the genera most like Acanthocarpus, species of Calappa have a broad, serrate, or spinose flange extending from the posterolateral margin, and species of Calappilia lack the flange but bear several small, triangular spines. Species of Mursia Leach in Desmarest, 1823 have a prominent spine dividing the anterolateral and posterolateral margins. None of these genera can accommodate the new species. The sole specimen is broken so that the frontal region and the posterior margin are not preserved. Furthermore, most of the left side of the specimen is missing. The carapace cuticle is bleached and the inner surface appears to have been etched, possibly during the molting process. The pterygostome is separated from the carapace, supporting the interpretation that this is a molt rather than a corpse. The only other potential fossil occurrence is Acanthocarpus obscurus from the Miocene of Panama. That specimen is quite incomplete, so like the species described here, the referral to Acanthocarpus is made with some reservation.

Nomenclatural statement: A life science identifier (LSID) number was obtained for the new species: urn:lsid:zoobank. org:pub:3A583FD6-D1CC-4137-907B-A9E4AB613744.

Calappidae incertae sedis (Fig. 15)

Material examined: Cheliped fragments (MMNS IP-1338, 1342, 2356, 9834, 9881, 9945, 9946, 9751). Occurrence: All known material from the Oligocene (Rupelian) Vicksburg Group of Mississippi, including the Glendon/Marianna Limestone (MMNS IP-1338, 1342), Marianna Limestone (IP- 9751), and Mint Spring Marl (IP-9834, 9881, 9945) (MMNS loc. MS.65.004), Smith County and the Glendon/Marianna Limestone (loc. MS.61.001; IP-2356), and the Mint Spring Marl (loc. MS.61.003; IP-9946), Rankin County, MS. Remarks: Recognition of chelae as originating from a calappid is based upon very distinctive features (see, for example, Rathbun, 1937: pl. Figure 14. Acanthocarpus osborni n. sp., holotype (MMNS IP-3780); dorsal 61). Propodi are triangular in outline, broadening distally, with a carapace (A). Image reflected to form composite image; the sulcus between short, weakly downturned fixed finger bearing short, blunt denticles. orbits is an artifact of breakage; the rostrum is unknown (B). Right lateral The upper surface often has a row of spines, the outer surface is view (C). Scale bars = 1 cm. variably arrayed with granules and nodes, and the inner surface is generally smooth and concave, conforming to the configuration of the anterior margin of the crab. The dactylus is strongly concave Pterygostome broad, weakly granular anteriorly, smooth else- on the occlusal surface and convex on the outer surface. The most where, with well-defined rim on inner margin (Fig. 14C). distinctive feature of the dactylus is a prominent, blunt, hook-like Sternum, pleon, appendages not preserved. projection extending from the proximal end of the occlusal surface and curving proximally and toward the occlusal surface. The feature Etymology: The trivial name honors Adam Osborn, the collector of functions as an anchor point when the crab attacks and fragments the the specimen. apertural margin of a gastropod (Schweitzer & Feldmann, 2010b).

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Figure 15. Calappidae: cheliped fragments (MMNS IP-9946), left chela and merus (A); left chela (MMNS IP-9834) (B); left chela (MMNS IP-9945) (C); right chela (MMNS IP-8917) (D); Right chela (MMNS IP-1338) (E); movable finger showing basal tooth (MMNS IP-9751) (F). Scale bars = 1 cm.

The specimens of propodi and dactyli attributed to identity of these specimens may be resolved when carapaces Calappidae from the Mint Spring and Marianna formations and claws are found together. conform closely to the above diagnosis. Because there are a large number of calappid species from these formations, however, it is not prudent to assign them to a particular species. Superfamily Leucosioidea Samouelle, 1819 The nodose and granular ornamentation on each specimen Family Matutidae De Haan, 1841 is distinctive so that comparisons between claws can be pro- posed, but species assignments cannot be made. Two speci- Included genera: Ashtoret Galil & Clark, 1994; Eomatuta De Angeli mens (MMNS IP-9945 (Fig. 15C) and MMNS IP-9946 (Fig. & Marchiori, 2009; Hemsut Feldmann, Schweitzer, Bennett, 15A)) bear rows of blunt nodes extending from the lower sur- Franţescu, Resar & Trudeau, 2011; Matuta Weber, 1795; Matutites face upward and distally to the upper surface. This pattern is Blow & Manning, 1996; Pseudohepatiscus Blow & Manning, 1996; distinctly different from the more randomly arrayed nodes on Szaboa Müller & Galil, 1998. the outer surface of MMNS IP-9834 (Fig. 15B) and MMNS IP-9881. These two propodi also have a distinctive granu- Diagnosis: Carapace not much wider then long, widest at about lar lower surface which is not evident on MMNS IP-9945 midlength, regions undifferentiated, ornamented with discrete and MMNS IP-9946. The propodi from the Mint Spring tubercles not arranged in rows; lateral spine may be present; Formation may well be attributable to two different species. anterolateral and posterolateral margins often with discrete spines; The sole propodus collected from the Marianna Formation front trilobate or with 4 spines; posterior margin narrow; pereiopods (Fig. 15F) has an etched and eroded surface but appears to 2–5 paddle-like; chelae with few spines on upper margin, rows of have less prominent nodes and a less strongly spinose upper tubercles on outer surface; sternum obovate, narrow, widest anterior surface than any of the Mint Spring specimens. The species to midlength; sterno-abdominal cavity extending onto sternite 4;

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Other species: Matutites americanus (Rathbun, 1935); M. miltonorum Feldmann, Bice, Schweitzer Hopkins, Salva & Pickford, 1998; M. periosus n. sp. Diagnosis: Carapace ovate, wider than long, widest at position of second to third anterolateral spine; regions broadly inflated, with sharp swellings at highest points; orbits circular, directed forward, front projecting slightly beyond orbits; anterolateral margin with 2 or 3 small, sharp spines; posterolateral margin weakly convex or nearly straight, with large, sharp spine; posterior margin narrow.

Matutites periosus n. sp. (Fig. 16) Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021 Type: Holotype (Fig. 16; MMNS IP-160). Diagnosis: Carapace narrowing considerably posteriorly; anterolateral margins with 2 large blunt spines plus smaller one; mesogastric with strong posterior swelling; hepatic region with weak transverse keel; branchial regions with transverse swelling, strong posterolateral swelling; cardiac region elongated, with strong Figure 16. Matutites periosus n. sp., holotype (MMNS IP-160). Scale anterior swelling, bar = 1 cm. Description: Carapace slightly wider than long, length about 94% maximum carapace width, widest about 35% the distance abdominal somites 3–5 fused in males (adapted from Bellwood, posteriorly at anterolateral corner; moderately vaulted transversely 1996; Schweitzer & Feldmann, 2000b). and longitudinally; regions moderately defined as broad raised areas, some with large tubercles at apices. Remarks: Blow & Manning (1996) originally placed Eriosachila Blow Front quadrilobed, outer lobe serving as inner orbital spine, & Manning, 1996, Matutites, and Pseudohepatiscus in Matutinae of axially and dorsally broadly sulcate, not projected beyond orbits, Calappidae. Feldmann et al. (1998) placed Matutites in Calappidae about 20% maximum carapace width. Orbits shallow, circular, and suggested that Pseudohepatiscus may be synonymous with it. rimmed, outer-orbital spine projected slightly upward, fronto- Schweitzer & Feldmann (2000b) placed Eriosachila and Matutites orbital width about half maximum carapace width. Anterolateral within Hepatidae Stimpson, 1871 (now Aethridae), based upon their margins rimmed, with 2 large blunt spines, one at about one-third well-defined carapace regions and spinose lateral margins. Eriosachila distance posteriorly on margin, one at anterolateral corner, weak as it is now construed is rather variable but possesses crispate, projection situated between 2 large spines. Posterolateral margins spinose anterolateral margins, usually concave posterolateral longer than anterolateral margins, large spine at about one-third margins, and a narrow front that is slightly produced beyond the the distance posteriorly on margin, margin flared laterally near orbits. Matutids have wider fronto-orbital widths, straight or weakly posterolateral corner, apparently to accommodate bases of pos- convex posterolateral margins, less broadly concave anterolateral terior pereiopods; posterior margin narrow, convex, about one- margins, and anterolateral and posterolateral margins about equal quarter maximum carapace width, with short, blunt projections at in length. We suggest that Eriosachila be retained in Aethridae based posterior corners. upon its conspicuously convex and crispate anterolateral margins, Mesogastric region with conspicuous narrow anterior pro- concave posterolateral margins, and narrow front produced beyond jection, posteriorly with wide swelling with 2 small tubercles the orbits. Matutites and Pseudohepatiscus might be better reassigned at apex. Protogastric regions with broad swelling posteriorly, back to Matutidae based upon their wide fronto-orbital width, weaker; smaller swelling at lateral margin. Hepatic region with four-lobed front not produced beyond the orbits, weakly convex broad transverse keel extending into first large anterolateral anterolateral margins, anterolateral and posterolateral margins spine. Urogastric region depressed well below level of mesogas- about the same length, convex or straight posterolateral margins, tric and cardiac regions. Cardiac region longitudinally elon- and discrete carapace spines much like extant matutids. gated, merging into weakly inflated intestinal region, with large Examination of the holotype of the type species of swelling anteriorly. Branchial regions not well-differentiated, Pseudohepatiscus, P. marinoi, as well as the holotype for Matutites with ovate, transverse swelling extending toward anterolat- anthonyae, suggests that they are very similar. Pseudohepatiscus marinoi eral corner, swelling with few apical tubercles. Mesobranchial appears to be a more weathered carapace, whereas M. anthonyae area with high, sharp tubercle at about level of posterolateral retains more carapace detail. The main difference between the spine. Pterygostomial region high, with scattered tubercles. two type species appears to be a weakly convex posterolateral Very poorly preserved fragment of sternum with sternites 4–7? margin with a large spine in Matutites, whereas that margin is preserved, sternites 5,6? with episternal projections. Chelipeds straight and entire in Pseudohepatiscus. Both are known from the slightly heterochelous, similar, left larger, length of manus of Santee Limestone in South Carolina (Blow & Manning, 1996). right cheliped about 96% length of left manus, height of manus Based upon this difference, we retain them as separate genera, of right cheliped about 93% height of left manus; merus with noting that they are very similar to one another. strong distal lower spine; carpus apparently with few spines on upper margin. Outer surface of chelae with 3 rows of large tubercles, one or two rows on lower margin, lower margin with Genus Matutites Blow & Manning, 1996 corrugated appearance in left manus, upper margin with several strong blunt spines forming crest-like structure. Fixed finger Type species: Matutites anthonyae Blow & Manning, 1996, by original extending straight from manus; movable finger stout proximally, designation. strongly arched.

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Figure 17. Correspondence Analysis (MVSP 3.13m) of decapod fauna at Oligocene localities in North America. Statistics in Supplementary material Table S2; N = number of Oligocene genera in each region (see Supplementary Material Table S1).

Measurements: Measurements (in mm) of the holotype: maximum species, Lophoranina lincki n. sp., Notopus adinae n. sp., Calappilia carapace width, 45.1; maximum carapace length, 42.3; fronto- granulosa n. sp., Calappilia perreaulti n. sp., Calappilia gableorum orbital width, 22.0; frontal width, 9.3; posterior width, 11.0; length n. sp., and Acanthocarpus osborni n. sp. Of these, Notopus and to position of maximum width, 15.5; length of left manus, 22.6; Acanthocarpus are reported from the Paleogene Gulf Coastal Plain length of right manus, 21.8; height of left manus excluding crest, for the first time. Other occurrences include previously described 13.7; height of right manus excluding crest, 12.7. caridean shrimps, axiids, pagurids, and raninids. Mesostylus, a callianassid, is described for the first time from Danian deposits, all Etymology: The trivial name is the Greek word periosus for “immense” other occurrences being Late Cretaceous. in reference to the large size of the specimen in relation to other Oligocene fossil decapods are reasonably well known from species in the genus. North America (Supplementary material Table S1). The most Occurrence: Eocene (early Bartonian) Cook Mountain Formation, diverse fauna is derived from west coastal rocks of Washington Potterchitto Member (MMNS loc. MS.51.001), Newton County, MS. and Oregon (Rathbun, 1926; Schweitzer & Feldmann, 1999; Schweitzer & Feldmann, 2000a, b, c, d; 2001; Fraaije et al., 2006; Remarks: Matutites periosus n. sp. differs from all other species of the Nyborg & Vega, 2008). Less diverse Oligocene faunas from west genus by its size. It has a stronger posterolateral spine as compared coastal North America are reported from British Columbia to all other species, and the carapace tubercles on the apices of (Rathbun, 1926; Nyborg et al., 2016), Alaska (Schweitzer & the protogastric, mesogastric, and branchial regions are larger than Feldmann, 2000a, d, 2001; Tucker & Feldmann, 1990) and those in other species. The swellings on the epibranchial region are Baja California Sur (Rathbun, 1930; Schweitzer et al., 2002, transversely elongate, also seen in M. anthonyae, but those of the new 2004). Gulf coastal localities are moderately diverse com- species are much larger. pared to other North American occurrences (Rathbun, 1935). Other species of Matutites are recovered from either Limestone Correspondence Analysis conducted in MVSP 3.13m indicates or clay of early Paleocene (M. americanus) or middle Eocene that the Gulf coastal Oligocene decapod fauna at the generic (M. anthonyae, M. miltonorum) age (Mancini & Tew, 1988; Blow level is most like that of Pacific Coastal Baja California Sur (Fig. & Manning, 1996, Feldmann et al., 1998). The Cook Mountain 17; Supplementary material Table S2). None of the localities, Formation in Mississippi, from which the new species was col- however, is particularly similar in terms of shared genera. Only lected, is composed of marls, sand, and clay of middle Eocene two genera (Calappa and Megokkos) occur at three localities, and age (Mancini & Tew, 1991). The new species does not extend the seven genera occur at two localities (Supplementary material geologic range of the genus, already known from the southeastern Table S1). Oligocene decapod occurrences in North America USA, nor does it expand the range of sedimentary environments also show a faunal gradient from lower to higher latitudes and reflecting the ecological range of the genus. from east to west (Fig. 17), indicating that faunal composition was strongly differentiated biogeographically (Schweitzer, 2001b; Nomenclatural statement: A life science identifier (LSID) num- Feldmann & Schweitzer, 2006). ber was obtained for the new species: urn:lsid:zoobank.org:pub: urn:lsid:zoobank.org:pub:56A4F26C-B2CF-4EF1-90F4-F6C1A01 B0DF5. SUPPLEMENTARY MATERIAL Supplementary material is available at Journal of Crustacean Biology DISCUSSION online. S1 Table. Oligocene occurrences of decapod crustacean genera Paleogene sediments of Mississippi have yielded one new Eocene in North America. brachyuran species, Matutites periosus n. sp. and six new Oligocene S2 Table. Correspondence Analysis results from MVSP 3.13m.

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ACKNOWLEDGEMENTS Coleman, J.L. Jr. 1983. The Vicksburg Group carbonates: A look at Gulf Coast Paleogene carbonate banks. Gulf Coast Association of Geological Numerous people contributed specimens to this project. We are Societies Transactions, 33: 257–268. especially grateful to Paul Hartfield, Josh Johns, Dana Linck, Cope, K.H., Utgaard, J.E., Masters, J.M. & Feldmann, R.M. 2005. The George Martin, Linda McCall, Meredith Montgomery, Adam fauna of the Clayton Formation (Paleocene, Danian) of southern Illinois: a case of K/P survivorship and Danian recovery. Bulletin of the Osborn, Shane Parrish, Scott Peyton, Ray Perreault, Joy Rushing, Mizunami Fossil Museum, 32: 97–108. Cathy Shropshire, Kelisha Stamps, James Starnes, Andy Weller, Davis, A. & Heydari, E. 2015. Sedimentology and petrography of Faye Whobrey, and the Leslie & Sue Pitts family. We are grateful Marianna and Glendon limestones of the Vicksburg Group in Smith to the landowners for locality access, particularly the Bryan Gable County, Mississippi. Geological Society of America Abstracts, 47: 526. family, Mac and Bryan Palmer, and Gray and James Morris. Data De Angeli, A. & Marchiori, L. 2009. Eomatuta granosa n. gen., n. sp. (Decapoda, Brachyura, Matutidae), nuovo crostaceo dell’Eocene dei for diversity analysis was compiled under NSF EAR-1223206 Monti Berici (Vicenza, Italia settentrionale). Lavori, Società Veneziana di to CES and RMF. Our thanks to the two anonymous reviewers Scienze Naturali, 34: 105–110. and the Editor-in-Chief for improving the manuscript. Data for De Bouillé, R. 1873. Paléontologie de Biarritz. Compte Rendu Travaux Congrès e diversity analysis was completed under NSF EAR-1223206 to Scientifique de France, 39 session, Pau: 11, pl. 4.

De Bouillé, R. 1876. Paléontologie de Biarritz et des autres localités des Basses Downloaded from https://academic.oup.com/jcb/article/39/3/279/5427698 by guest on 29 September 2021 CES and RMF. Partial support to cover the cost of publication Pyrénées. Lithographie Veronese, Pau, France. was provided by Carl Campbell, Linda McCall, George Martin, Desmarest, A.G. 1822. Histoire naturelle des Crustacés fossiles. Les Crustacés Rick Poropat, Fay Whobrey and the Eastern Missouri Society of proprement dits: 67–154, pls. 5–11. F. G. Levrault, Paris. Paleontology. Desmarest, A.G. 1823. Malacostracés. Dictionnaire des Sciences naturelles, 28: 138–425. REFERENCES Dockery, D.T. III. 1977. Mollusca of the Moodys Branch Formation, Mississippi. Mississippi Geological, Economic and Topographical Survey Albright, B. III, Phillips, G., Starnes, J., Stringer, G. & Weller, A. 2016. Bulletin, 120, 1–212, Appendix. The Jones Branch local fauna: An early Arikareean mammalian assem- Dockery, D.T. 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