New Cretaceous and Eocene Decapoda (Astacidea, Thalassinidea, Brachyura) From British Columbia, Canada

ANNALS

OF

CARNEGIE

MUSEUM 28 February 2009

Vol. 77, Number 4, Pp. 403–423

NEW CRETACEOUS AND EOCENE DECAPODA (ASTACIDEA, THALASSINIDEA, BRACHYURA) FROM BRITISH COLUMBIA, CANADA Carrie E. Schweitzer [Research Associate, Section of Invertebrate Paleontology, Carnegie Museum of Natural History] Department of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW, North Canton, Ohio 44720 [email protected]

Rodney M. Feldmann [Research Associate, Section of Invertebrate Paleontology, Carnegie Museum of Natural History] Department of Geology, Kent State University, Kent, Ohio 44242 [email protected]

Vlasta C´ osovic´

Department of Geology and Paleontology, Faculty of Science, University of Zagreb, Horvatovac, 102A, 10000 Zagreb, Croatia [email protected]

Richard L. M. Ross 326 Quarry Rd., Comox, BC Canada V9M3C4 [email protected]

David A. Waugh Department of Geology, Kent State University, Kent, Ohio 44244 [email protected]

Abstract A collection of Cretaceous and Eocene specimens of decapod crustaceans from southern Vancouver Island, British Columbia, Canada, has yielded the following new taxa: Cowichianassa meckerti, new genus, new species (Callianassidae); Comoxianassa haggarti, new genus, new species (Callianassidae); Preclarocarcinus parvus, new genus, new species (Homolodromiidae); and Nitotacarcinus canadensis, new species (Tumidocarcinidae). Two of the new taxa are callianassid ghost shrimp, each of which displays sexual dimorphism and is well documented from major and minor claws as well as abdomina, unusual in the fossil record and especially unusual for Cretaceous forms. Nitotacarcinus was known previously only from the Eocene of Denmark; thus, the new species extends its paleobiogeographic range dramatically. The layers in which the decapods were collected from the Paleogene Via Appia Beds were deposited in generally high-energy conditions favoring the formation of glauconite. Cuticular structures of Eucorystes platys Schweitzer and Feldmann, 2001, are described. Key Words: Astacidea, Brachyura, Canada, Cretaceous, Decapoda, Eocene, sexual dimorphism, Thalassinidea

INTRODUCTION Recent research on fossil decapods has resulted in numerous additions to the decapod fauna of western Canada and Pacific Coastal North America in general (Schweitzer 2001; Schweitzer et al. 2003, 2004, and references therein). The Cretaceous decapod fauna from coastal British Columbia is now quite robust (Table 1), and in addition, five Eocene species within four families are also known. We here also provide sedimentological data on the Via Appia Beds, which corroborate previous interpretations of the unit as shallow marine. Because of these advances, we now have a much more detailed understanding of the paleoecology, paleobiogeography, evolution, and phylogeny of the North American Decapoda. Herein, we describe several new species, most of which mark new records for the genera. LOCALITIES Eden Creek.—Via Cranberry Lane exit to Inland Island

Highway, 2 km to Oyster River Main Logging Road, then 9.5 km to Eden Main Logging Road, then 2.6 km to just past Eden 300 Branch in a ditch roadcut, lat. 49°49’39.4”N, long. 125°25’43.5”W (Schweitzer et al. 2003), from late Turonian to Campanian Comox Formation (Karafit and Stockey 2008). The rocks at the Eden Creek locality were reported to be early Coniacian in age (Karafit and Stockey 2008, and references therein). Ammonites from the locality include species of Lewesiceras Spath, 1939, Prionocycloceras Spath, 1926, and Mesopuzosia Matsumoto, 1954. Because the ages of the formations for the various localities reported herein are already well documented (Mustard 1994; Karafit and Stockey 2008), we report the associated cephalopods to supply a more complete picture of the paleoecology and associated fauna at each locality. Northwest Bay Beach.—Located 3 km south of Parksville, on the east coast of Vancouver Island, British Columbia, lat. 49°19’ 00” N , long. 124°13’ 00” W, probably early Santonian Haslam Formation. Ammonites include species

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Table 1. Updated list of Decapoda known from coastal British Columbia. List after that of Schweitzer et al. (2003); updated taxonomy after Karasawa and Schweitzer (2006) and Karasawa et al. (2008). † indicates taxa described herein; * indicates Eocene taxa; remainder are Cretaceous in age. Infraorder Astacidea Latreille, 1802 Superfamily Glypheoidea Winckler, 1883 Family Glypheidae Winckler, 1883 Genus Glyphea von Meyer, 1835 Glyphea sp. Family Mecochiridae Van Straelen, 1924 [imprint 1925] Genus Meyeria McCoy, 1849 Meyeria? harveyi Woodward, 1900 Superfamily Nephropoidea Dana, 1852 Family Erymidae Van Straelen, 1924 [imprint 1925] Genus Phlyctisoma Bell, 1863 Phlyctisoma dawsoni (Woodward, 1900) Genus Enoploclytia McCoy, 1849 Enoploclytia minor Woodward, 1900 † Enoploclytia sp. Family Nephropidae Dana, 1852 Genus Hoploparia McCoy, 1849 Hoploparia horrida Schweitzer et al., 2003 Hoploparia tshudyi Schweitzer and Feldmann, 2001

Superfamily Homolodromioidea Alcock, 1899 Family Homolodromiidae Alcock, 1899 † Preclarocarcinus new genus † Preclarocarcinus parvus new species Superfamily Raninoidea de Haan, 1839 Family Raninidae de Haan, 1839 Subfamily Palaeocorystinae Lo˝renthey in Lo˝renthey and Beurlen, 1929 Genus Eucorystes Bell, 1863 Eucorystes harveyi (Woodward, 1896) † Eucorystes platys Schweitzer and Feldmann, 2001 Subfamily Lyreidinae Guinot, 1993 Genus Macroacaena Tucker, 1998 Macroacaena chica Schweitzer et al., 2003* Macroacaena naselensis (Rathbun, 1926) Subfamily Raninoidinae Lo˝renthey in Lo˝renthey and Beurlen, 1929 Genus Raninoides H. Milne Edwards, 1837 † ?Raninoides sp.

Infraorder Thalassinidea Latreille, 1831 Superfamily Callianassoidea Dana, 1852 Family Callianassidae Dana, 1852 Genus Calliax de Saint Laurent, 1973 Calliax whiteavesi (Woodward, 1896) † Genus Comoxianassa new genus † Comoxianassa haggarti new species † Genus Cowichianassa new genus † Cowichianassa meckerti new species Genus Neocallichirus Sakai, 1988 Neocallichirus manningi Schweitzer et al., 2003 Family Ctenochelidae Manning and Felder, 1991 † Genus and species indeterminate

Section Heterotremata Guinot, 1977 Superfamily Dorippoidea MacLeay, 1838 Family Necrocarcinidae Förster, 1968 Genus Cenomanocarcinus Van Straelen, 1936 Cenomanocarcinus beardi Schweitzer et al., 2003 Family Orithopsidae Schweitzer et al., 2003 Genus Paradoxicarcinus Schweitzer et al., 2003 Paradoxicarcinus nimonoides Schweitzer et al., 2003 Superfamily Carpilioidea Ortmann, 1893 Family Tumidocarcinidae Schweitzer, 2005 Genus Nitotacarcinus Schweitzer et al., 2007 † Nitotacarcinus canadensis new species* Genus Pulalius Schweitzer et al., 2000 Pulalius vulgaris (Rathbun, 1926)* Superfamily Portunoidea Rafinesque, 1815 Family Longusorbiidae Karasawa et al., 2008 Genus Longusorbis Richards, 1975 Longusorbis cuniculosus Richards, 1975 Family Mathildellidae Karasawa and Kato, 2003 Genus Branchioplax Rathbun, 1916 Branchioplax carmanahensis (Rathbun, 1926)* Family Macropipidae Stephenson and Campbell, 1960 Genus Megokkos Schweitzer and Feldmann, 2000 Megokkos alaskensis (Rathbun, 1926)* Superfamily Retroplumoidea Gill, 1894 Family Retroplumidae Gill, 1894 Genus Archaeopus Rathbun, 1908 Archaeopus bicornutus Schweitzer et al., 2003* Archaeopus rostratus Schweitzer et al., 2003 Archaeopus vancouverensis (Woodward, 1896)

Infraorder Palinura Latreille, 1802 Superfamily Palinuroidea Latreille, 1802 Family Palinuridae Latreille, 1802 Genus Linuparus White, 1847 Linuparus canadensis (Whiteaves, 1884) Linuparus vancouverensis (Whiteaves, 1895) Infraorder Brachyura Linnaeus, 1758 Section Podotremata Guinot, 1977 Superfamily Dromioidea de Haan, 1833 Family Dynomenidae Ortmann, 1892 Genus Acanthodiaulax Schweitzer et al., 2003 Acanthodiaulax mclayi Schweitzer et al., 2003 Superfamily Homoloidea de Haan, 1839 Family Homolidae de Haan, 1839 Genus Latheticocarcinus ludvigseni Schweitzer et al., 2004

of Canadoceras Spath, 1922, Pseudoschloenbachia Spath, 1921, and Damesites Matsumoto, 1942. Trent River.—Float shale blocks at the site of a former railway bridge over the Trent River, 7.2 km west of Highway 19A on Bayton Main Logging Road at lat. 49°35’30.5” N, long. 124°59’28.4” W from the lower Haslam Formation of early Santonian age (Schweitzer et al. 2003). The ammonite fauna includes species of Hauericeras de Grossouvre, 1894, Polyptychoceras Yabe, 1927, Neophylloceras Shimizu, 1934, and Kitchinites Spath, 1922.

Dove Creek Road.—Blast piles (and remaining roadcut) from construction at the intersection of Inland Island Highway 19 and Dove Creek Road at lat. 49°43’42.4” N, long. 125°06’15.1” W, from the Pender Formation of early Campanian age (Schweitzer et al. 2003). New Island Highway.—Collected from construction debris of New Island Highway, north of Courtenay, British Columbia, 300 m north of Hydro crossing. Rocks exposed there are the early Campanian Pender Formation (Mustard 1994).

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Fig. 1.—Photomicrographs of concretions from the Via Appia beds. A–B, specimen 1 from lower Via Appia beds (note sorting and subangular to subrounded shape); C–E, specimen 2 from lower Via Appia beds (note gastropods with geopetal infillings, suggesting that they were reworked (D) and infillings of mollusk shells which are the same composition as surrounding matrix (E)); F–G, Appian Concretion 1 from upper Via Appia beds, including clasts of wood, mollusks, and echinoid spines; H, Appian Concretion 2 from upper Via Appia beds; I, Appian Concretion 3 from upper Via Appia beds.

Puntledge River Fish Hatchery.—Just downstream from fish hatchery along Puntledge River, lat. 49°41’21.2”N, long. 125°2’7.5”W, early Campanian Pender Formation (Mustard 1994). Ammonites collected from the Island Highway and Puntledge River sites include species of Eupachydiscus Spath, 1922, Urakawites Matsumoto, 1955, Eubostrychoceras Matsumoto, 1967, and Glyptoxoceras Spath, 1925. Von Ark’s.—On Puntledge River, 3 km northwest of Courtenay with access via the end of Cessford Road and a farm road to the top of the bank through Von Ark’s dairy farm. The trail down to the shale exposure follows a cable support to a cable car used by the farm and the bottom portion is accessed by rope; lat. 49°41’ 16.6” N, long. 125°01’ 40.1” W; in rocks of the early Campanian Pender Formation.

Via Appia Beds.—Beach exposures at low tide south of Shelter Point, on Highway 19A to Appian Way Road, 14 km south of Campbell River, BC; lower bed at lat. 49°54’44.8”N, long. 125°10’46.9” W (Schweitzer et al. 2003) and upper bed at lat. 49° 54’43.5”N, long. 125°10’39.4” W, from unnamed rocks assigned to a middle-late Eocene age by some (Haggart et al. 1997; in Mindell et al. 2006; in Elliott et al. 2006), and to a Paleocene to Eocene age by Sweet (1997), in Mindell et al. (2006), and in Elliott et al. (2006). Data presented by Mindell et al. (2006) and Elliott et al. (2006) are consistent with a Paleogene age. Nautiloid cephalopods have been collected from the unit. SEDIMENTOLOGY OF VIA APPIA BEDS Because the age of the Via Appia beds is somewhat uncertain,

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we conducted a micropaleontological analysis of several concretions collected from the unit. Unfortunately, few foraminiferans were recognized in the rocks, but sedimentological observations allowed an interpretation of the depositional environment in which decapods were collected. Lower Via Appia beds.—Specimen 1 (Figs. 1A, B) grains are more or less similar in size (0.1 mm to 0.5 mm in diameter) and shape (subangular to subrounded), indicating good sorting (long transport or high-energy environment). The grains include quartz, plagioclase, and glauconite. Grains are immersed in a brown calcite matrix, and the plagioclase grains have margins that show calcification (dissolution and cementation). The environmental setting was most likely shallow with a large influx of terrestrial material. The composition of the grains of Specimen 2 (Figs. 1C– E) is the same as the other specimens with glauconite less than 1% of total grains. They are held together by micrite and clay. Fossils include crab carapaces, sponge spicules, gastropods, and mollusk bioclasts. Gastropod shells show geopetal infillings, suggesting that they were reworked. The remnants of crab carapaces show bioerosional drilling holes, so that before being buried they were exposed to bioeroders. The infillings of mollusk shells are the same composition as surrounding matrix. The depositional environment was likely shallow, agitated water. Upper Via Appia beds.—Appian Concretion 1 (Figs. 1F, G) contains scattered quartz and feldspar grains dispersed throughout a silty calcareous muddy matrix. Glauconite grains comprise around 1% of total inorganic grains. The fossil content includes pelecypods, gastropods, crabs (abundant), echinoid spines, probably brachiopods, and smaller benthic foraminiferans (probably a shallow infaunal form, cf. species of Vaginulina d’Orbigny, 1826 or Vaginulinopsis Silvestri, 1904). The mollusk shells are complete, whereas the crab remains show a high degree of fragmentation. Dissolution and bioerosion effects are absent and sorting is poor. This suggests a somewhat sheltered, calm sedimentary environment, where the crab debris was transported due to its lower density (buoyancy effect). Organisms that depend on light are missing and glauconite grains are present, suggesting light-deprived conditions due to terrigenous influx. The conditions in the environment in which this concretion originated differ from those in which the other concretions formed. It was collected from the upper beds of the Appian Way unit. Appian Concretion 2 (Fig. 1H) contains glauconite grains comprising 2–3.5 % of the total inorganic content. Remnants of crabs and bryozoans are present. The quartz grains and feldspars are densely packed, more or less of the same size, and angular in shape. A silty, calcareous matrix is present in a considerably smaller amount than in Appian Concretion 1 and sorting is better. Occasionally, muddy matrix is concentrated in forms similar to muddy balls. High energy is suggested by sorting and by grains

being in contact where the matrix is “swept” out. Appian Concretion 3 (Fig. 1I) is composed of wellsorted, angular quartz and feldspar grains within a muddy, silty matrix. Glauconite grains comprise about 1% of all inorganic grains. The grains are partly weathered (brown margins), suggesting diagenetic processes. Fossils include crabs, but they comprise fewer than 5% of all grains in the sample. Fossils are fragmented into small pieces. The rocks were formed in a high-energy setting, too energetic for foraminifera. These interpretations are consistent with previous data on the Via Appia beds. Mindell et al. (2006) and Elliott et al. (2006) each reported the unit to be shallow marine and described plant material from it. Our results suggest a largely shallow, high-energy environment for both the lower and upper Via Appia beds. Notably, Appian Concretion 1 is distinctly different from the other concretions in terms of appearance, sorting, and depositional environment. It was collected from the upper Appian Way beds as were Appian Concretions 2 and 3, which were formed in what was apparently the more general environment for the Via Appia Beds (high-energy, shallow). This mixture of depositional environments is consistent with a setting in which a mixture of plant and marine material could be preserved together, in a near-shore, tectonically active area receiving an influx of terrestrial sediments which was then sorted by wave energy. Downslope mixing of concretions and fossils could explain the differences in sedimentology in the concretions we examined. SYSTEMATIC PALEONTOLOGY Order Decapoda Latreille, 1802 Infraorder Astacidea Latreille, 1802 Family Erymidae Van Straelen, 1924 [imprint 1925] ?Enoploclytia McCoy, 1849 Type species.—Astacus leachi Mantell, 1822, by original designation. ?Enoploclytia sp. (Fig. 2) Material examined.—GSC (Geological Survey of Canada, Eastern Paleontology Division, Ottawa, Ontario, Canada) 27174 and 27177. Occurrence.—Northwest Bay Beach and Dove Creek Road localities. Discussion.—Two well-preserved, but incomplete, macruran specimens can tentatively be assigned to Enoploclytia, although they show none of the characters necessary to assign the material to a known species or to document a new species. One of the specimens (Fig. 2A) consists of part and counterpart of the right side of the branchiostegite,

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Fig. 2.—?Enoploclytia sp. A, GSC 27174, right lateral view of cephalothorax; B, right lateral view of cephalothorax, GSC 27177; C, posterior view of abdomen, GSC 27177; D, left lateral view of abdominal somites, GSC 27177; E, right lateral view of abdominal somites, GSC 27177. Scale bars = 1 cm.

measuring 38.9 mm from the cervical groove to the posterior margin. The cervical groove is partially exposed and is broad, smooth, and much stronger than the postcervical and branchiocardiac grooves, which are about equally developed and subtle. The postcervical groove extends anteroventrally beyond the termination of the branchiocardiac groove joining an inferior groove that is also weakly developed. The posteromarginal carina and postmarginal groove are well defined. The entire surface of the branchiostegite is covered by uniform sized and evenly distributed fine pustules. The second specimen (Figs. 2B–E) exhibits the right branchiostegite with morphology similar to that of the other specimen but is somewhat smaller, approximately 30 mm from the cervical groove to the posterior margin. It also exhibits the first three abdominal somites. The first somite is small, 5.2 mm long, poorly exposed, and appears to lack a well-developed pleuron. The second somite is 6.0 mm long, excluding the articulating ring, axially keeled, with a boss defining the separation between the tergum and pleuron, and a rectangular pleuron which is slightly drawn out on the posterolateral corner. The pleuron has a broad rim and the entire somite is pustulose. The third somite is 4.9 mm long, with a tergum similar to that of somite two but with a narrow, triangular pleuron with a tiny

posterolateral spine. These characters conform to the descriptors of the genus (Förster 1966: 146) but, because the specimens are so fragmentary, comparison with described species cannot be done with confidence. Examination of the illustrations of British representatives of the genus (Woods 1925–1931: pl. 24, fig. 4) tempts comparison with the type species. The ornamentation on the branchiostegite is similar, but this is an unreliable basis for comparison because many erymid taxa exhibit similar ornamentation. The only species of Enoploclytia that has been described from the Pacific Northwest is Enoploclytia minor Woodward, 1900, from Hornby Island, British Columbia. The sole specimen of that species is extremely poorly preserved (Feldmann and McPherson 1980: pl. 1, figs. 1, 2) and a reliable comparison is not possible. Thus, it is prudent to retain the specimens questionably in Enoploclytia until better and more complete material is discovered. Infraorder Thalassinidea Latreille, 1831 Superfamily Callianassoidea Dana, 1852 Family Callianassidae Dana, 1852 Discussion.—The two new genera described herein are most closely allied with the Callianassidae based upon

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Fig. 3.—Callianassidae. A, Cowichianassa meckerti, major and minor cheliped, female, GSC 27162, paratype; B, Cowichianassa meckerti, major cheliped and abdomen, male, GSC 27160, paratype; C, Comoxianassa haggarti, major and minor chelae and abdomen, female, GSC 27157, paratype; D, Cowichianassa meckerti, major cheliped, male, GSC 27159, holotype; E, Comoxianassa haggarti, cheliped and abdomen, gender indeterminate, GSC 27158, paratype; F, Cowichianassa meckerti, inner surface of major chela, male, GSC 27161, paratype; G, Comoxianassa haggarti, major cheliped, male, GSC 27156, holotype; H, Comoxianassa haggarti, abdominal somites 3, 4, and 5, gender indeterminate, GSC 27158, paratype; I, Comoxianassa haggarti, abdominal somite 6 and portion of telson, gender indeterminate, GSC 27158, paratype; J, Exaflex® cast of Comoxianassa haggarti, major and minor chelae and abdomen, female, GSC 27157, paratype. Scale bars = 1 cm.

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several factors. Callianassidae possess large, well-developed carpi and often have unequal first pereiopods, both features seen on the specimens described herein. Other families within the superfamily and even the infraorder have carpi of reduced size compared to those seen in the Callianassidae. A review of recent literature on the Callianassoidea indicates that subfamily designations are in flux (Manning and Felder 1991; Sakai 1999, 2001, 2005; Ngoc-Ho 2003; Poore 2005); thus, we do not elect to use them here. A review of current genera in use indicates that none can accommodate the new material. This is not surprising, given the Campanian (Late Cretaceous) age of the specimens. In addition, these specimens are some of the best-preserved callianassids of Cretaceous age, yielding proximal elements of the major cheliped, distal elements of the minor cheliped, and some elements of the abdomen. Cretaceous callianassids are rarely known from such varied material. Examples include specimens of various species of Protocallianassa Beurlen, 1930, from which the new material is distinguished based upon the shape of the manus and carpus. Callianassa masanorii Karasawa, 1998, retains many elements of the carapace, abdomen, and appendages; however, such preservation is unusual. In spite of the differences of the new callianassid material with younger and extant callianassids, there are amazing similarities between the Cretaceous material and extant Callianassidae. In at least one of the Cretaceous specimens, some elements of what appear to be pleopods are preserved (Fig. 4B), and these elements are nearly identical in shape to the exopod of pleopods of extant callianassids. In other specimens, the abdominal somites are preserved, and they retain the same overall size and shape ratios as those of extant callianassids. The mani, carpi, and meri of the new fossils show ornamentation that is identical to that of modern callianassids, although the ornamented elements do not occur in the same combinations with one another as seen in any existing named genera. Thus, although the genera are different, the basic callianassid body plan has changed little since at least the Late Cretaceous. Other Cretaceous callianassids have been reported from the Pacific Northwest of North America. Calliax whiteavesi (Woodward, 1896) is known from various localities in coastal British Columbia (Schweitzer et al. 2003, and references therein). That species possesses very fine serrations on the lower margin of the merus as well as a fixed finger that is downturned at a marked angle to the proximal portion of the manus; neither condition is present in the two new genera named below. Neocallichirus manningi Schweitzer et al., 2003, has been reported from the Cretaceous of Vancouver Island. That taxon has fine serrations on the lower margin of the manus and a row of setal pits along the lower margin of the carpus, neither of which is present in the new material described here. Thus, the new material is unique. Indeed, it should be mentioned that both C. whiteavesi and N. manningi are known from different formations of different ages than the two new

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genera. As far as the two new genera, it is not uncommon to have two species of two genera of callianassid living in the same environment. Just a cursory scan of the biological literature of the group demonstrates that even species within the same genus may inhabit the same general area. Cowichianassa, new genus Type species.—Cowichianassa meckerti, new species, by monotypy. Diagnosis.—As for species. Etymology.—The genus name is derived from Cowichan, the name of indigenous peoples of Vancouver Island and British Columbia and the name of numerous geographical landmarks in the region. The gender is feminine. Discussion.—Cowichianassa, new genus, differs from all other genera within the family in possessing a longer-thanhigh ischium with a longitudinal groove and a bulbous distal end; a merus with marked, small spines on the lower margin; a carpus that is about as long as high; a manus with a proximal margin at a slightly oblique angle to the lower margin; markedly heterochelous first pereiopods; markedly sexually dimorphic chelipeds; and abdominal somites with large lower distal projections overlapping the succeeding somite. No other genus exhibits this combination of characters. Calliax de Saint Laurent, 1973, a genus to which numerous fossil species, including some Cretaceous ones have been assigned, possesses a marked spine on the ischium of the first pereiopod which is lacking in the new specimens. Neocallichirus Sakai, 1988, also embracing numerous fossil species, has a bulbous projection parallel to the distal margin of the manus which is not present in the new material. Protocallianassa, an extinct genus, exhibits a carpus with a markedly oblique lower margin with a distally directed projection on the lower distal corner, and it has the proximal margin of the manus oriented at about a 110 to 120 degree angle to the lower margin of the manus, not seen in the new material. Thus, the new genus is warranted. Cowichianassa meckerti, new species (Figs. 3A–B, 3D, 3F, 4) Diagnosis.—Ischium longer than high, with longitudinal groove, with bulbous distal end; merus with marked, small spines on lower margin; carpus about as long as high; manus with proximal margin at slightly oblique angle to lower margin; markedly heterochelous and sexually dimorphic chelipeds; abdominal somites with large lower distal projections overlapping succeeding somite. Description.—Ischium of male major cheliped longer than high, becoming higher distally, upper margin sinuous, distal margin with concave depression for articulation with merus, outer surface with longitudinal groove. Merus longer than high, height about 80% of length, upper

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Table 2. Measurements (in mm) taken on specimens of Cowichianassa meckerti, new species and Comoxianassa haggarti, new species. L1, manus length; H1, manus height; L2, movable finger length; L3, fixed finger length; L4, carpus length; H2, carpus height; L5, merus length; H3, merus height

Specimen Number

L1

H1

GSC 27159 (Holotype) GSC 27162 GSC 27161 GSC 27160

12.2

11.0

10.0 9.0 8.6

8.4 6.6 7.7

10.3 9.3

GSC 27157 GSC 27156 (Holotype)

GSC 27159 (Holotype) GSC 27162 GSC 27161 GSC 27157 GSC 27158 GSC 27156 (Holotype)

L2

L3 L4 Major Cheliped Cowichianassa meckerti, new species 7.1 4.3 9.2

H2

L5

H3

10.4

8.8

6.9

7.1 — 7.0

9.2 — 5.5

4.8 — —

9.2 7.9

9.3 7.2 5.0 — — — 6.7 — — Comoxianassa haggarti, new species 7.3 5.3 — — — 3.6

— 7.0

— 5.3

— 3.6

7.1

3.9

Minor Cheliped Cowichianassa meckerti, new species — — —

—

—

—

10.3 10.7

3.2 4.5

3.3 3.7

— —

— —

12.1 3.5 9.6

4.3 3.7 —

4.1 3.2 —

— — —

— — —

— — 4.4 — — 3.7 Comoxianassa haggarti, new species — — 3.8 3.5 3.2 2.6 — — —

surface weakly convex, proximal margin short, lower margin convex, serrate; outer surface longitudinally keeled, keel about one-third the distance from upper margin. Carpus higher than long, length about 90% of height, proximal margin with short projection for articulation with merus, then forming concave arc, followed by convex portion which merges with convex lower margin; upper margin weakly convex, distal margin weakly concave. Manus slightly longer than high, height about 90% length, stout, outer surface very convex; distal margin with projection centrally for articulation with carpus, oriented at about 100 degree angle to lower margin of manus; upper margin weakly convex; lower margin weakly convex, possibly with serrations or setal pits; distal margin initially straight, perpendicular to upper margin, then becoming oblique and merging with fixed finger. Fixed finger stout, shorter than movable finger, straight, apparently edentulous. Movable finger arcuate, stout, with large setal pits paralleling occlusal surface, possibly with one blunt tooth distally. Manus of male minor chela longer than high, distal margin straight, at about 115 degree angle to lower margin, upper and lower margins straight; fixed finger straight. Male abdominal somite 4 poorly known. Somite 5 short, with angular projection on distal lower corner which articulates with somite 6. Somite 6 about same size as 5, with concavity in proximal margin to receive posterior projection of somite 5, with thickened lower distal swelling which articulates with telson. Telson narrowing distally, blunt tipped. Endopod of male pleopod 2 (?) longer than wide, obovate, widest at midlength, narrowest at base, narrowing distally, blunt tipped, with small projection at inner edge for articulation with distal projection; with distal projection that may be appendix masculina, appendix interna, or both fused together, structure longer than wide, deeply grooved axially so as

to superficially appear to be two structures or two fused structures, inner half narrow, outer half wider, flattened (Fig. 4B). Endopod of male pleopod 3(?) longer than high, markedly bent at 45 degree angle at midlength, tip unknown, with appendix interna near base. Other elements of carapace and appendages unknown or too poorly preserved to be described. Ischium of female major cheliped longer than high, becoming higher distally, upper surface sinuous, with longitudinal groove on outer surface. Merus longer than high, with longitudinal keel, incompletely preserved. Carpus higher than wide, length about 70 % height, upper margin nearly straight, proximal margin arcuate and arcing into convex lower margin, lower margin with small, forward directed flange on lower distal corner; distal margin apparently nearly straight. Manus longer than high, height about 84 % length, less inflated and less convex than male, proximal margin oriented at about 98 degrees to lower margin, upper and lower margins nearly straight; distal margin initially perpendicular to upper margin, then arcing to merge with fixed finger. Fixed finger long, narrowing distally. Movable finger arcuate, with small projection at about midlength. Carpus of minor chela about as long as high, with long projection on upper proximal corner for articulation with merus, upper margin paralleled by setal pits. Manus of minor chela longer than high, rectangular, fingers apparently relatively long and slender. Other elements of female unknown.

Measurements.—Measurements (in mm) taken on elements of the major and minor chelipeds of C. meckerti, new species are presented in Table 2. Etymology.—The trivial name honors Dirk Meckert,

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Fig. 4.—Cowichianassa meckerti, GSC 27159, holotype. A, Outer surface of major cheliped of male; B, closeup of pleopods 2 (P2) and 3 (P3). DP indicates distal projection of pleopod 2 that may be appendix masculina, appendix interna, or both. Scale bars = 1 cm.

Courtenay, British Columbia, who has donated numerous specimens for our study of decapods of the region. Types.—GSC 27159, holotype, and paratypes GSC 27160–27162. Occurrence.—The specimens were collected from the Dove Creek Road locality.

Discussion.—The new species includes some specimens interpreted to be males and some to be females. The specimens considered as male specimens possess stout mani with very convex outer surfaces and stout fingers. The specimen interpreted to be a female possesses a more flattened manus and more slender fingers. We interpret it to be a member of the same species as the males referred to C. meckerti, because it has the ischium of the same shape and with a longitudinal groove, it has a sizable merus with

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Fig. 5.—Ctenochelidae, genus and species indet., GSC 27175. A, right lateral view of abdomen and cephalothorax; B, dorsal view of abdominal somites and cephalothorax; C, left lateral view of abdomen and cephalothorax; D, dorsal view of telson and uropods; E, left lateral view of telson and uropods. Scale bars = 1 cm.

a longitudinal keel although the lower surface is broken so that the presence of spines cannot be confirmed, and it possesses a long carpus as in the male specimens. Sexual dimorphism is well documented in fossil and extant callianassids, and it is typical for the female to possess more slender, less stout claws than the male (Schweitzer

et al. 2006a and references therein). In addition, both the specimens referred to as males and the specimen referred to as a female possess minor chelae that are very much smaller than the major chela. The extension on the end of the pleopod structure (Fig. 4) interpreted to be possibly the appendix masculina,

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appendix interna, or both, does not appear in any extant Callianassidae; Micheleidae Sakai, 1992; or Axiidae Huxley, 1879 sensu lato, we examined at the United States National Museum of Natural History or in any illustrations or descriptions of Thalassinoidea we could find in recent literature (e.g., Manning and Felder 1991; Poore 1994, 2005; Sakai 1999, 2001, 2005; Ngoc-Ho 2003). The two structures were described as fused at times but in no illustrations, descriptions, or examined material did they extend from the distal end of the pleopod. We considered the possibility that the pleopod and the distal structure were indeed two different structures superimposed on one another during the process of fossilization; however, careful examination with camera lucida and high magnification suggests that they were indeed attached to one another. Thus, we describe and illustrate the structure in hopes that it is recognized in other fossils. There appears to be potentially a preference for handedness in the specimens. Two of the males have right major chelipeds and one is a left major cheliped. The sole female is a left major cheliped. In Melipal chilensis Schweitzer et al., 2006a, and in Callianopsis clallamensis (Withers, 1924), nearly all of the major chelae were right chelipeds, showing no pattern in terms of gender (Schweitzer-Hopkins and Feldmann 1997; Schweitzer et al. 2006a). Unfortunately, the sample size for C. meckerti is too small to make conclusive statements about handedness. Comoxianassa, new genus Type species.—Comoxianassa haggarti, new species, by monotypy. Diagnosis.—As for species. Etymology.—The genus name is derived from the word Comox, a group of indigenous peoples from British Columbia and the name of a Cretaceous formation in the region. The gender is feminine. Discussion.—Comoxianassa new genus differs from all other genera within the Callianassidae in possessing a major cheliped with a merus with a longitudinal keel and apparently an entire lower margin; a relatively narrow carpus with a marked indentation in the proximal margin below the articulation with the merus; a large spine on the distal margin of the manus; a large spine on the movable finger that articulates with the spine on the distal margin of the manus; a minor cheliped that is smaller than the major but still retains a moderate size, especially in females; and abdominal somites with rounded projections on the lower distal corners that overlap and articulate with the following somite. Calliax possesses a marked spine on the ischium of the first pereiopod which is lacking in specimens of Comoxianassa. Neocallichirus has a bulbous projection parallel to the distal margin of the manus which is not present

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in the new genus. Protocallianassa exhibits a carpus with a markedly oblique lower margin with a distally directed projection on the lower distal corner and a proximal margin of the manus oriented at about a 110 to 120 degree angle to the lower margin of the manus, not seen in the new specimens. Examination of a variety of genera at the United States National Museum suggests that modern callianassids do not appear to have a longitudinal groove on the ischium of the major cheliped. Thus, the new genus is warranted. Comoxianassa haggarti, new species (Figs. 3C, 3E, 3G–3J) Diagnosis.—Major cheliped with merus with longitudinal keel and apparently an entire lower margin; relatively narrow carpus with marked indentation in proximal margin below articulation with merus; large spine on distal margin of manus; large spine on movable finger articulating with spine on distal margin of manus; minor cheliped smaller than major but still retaining moderate size, especially in females; abdominal somites with rounded projections on lower distal corners overlapping and articulating with following somite. Description.—Ischium of major cheliped of male longer than high, with longitudinal groove, poorly known. Merus of major cheliped of male longer than high, height about 70% of length, ovate; proximal margin nearly straight; upper margin convex; lower margin convex, apparently entire; distal margin short, nearly straight; outer surface convex, longitudinally keeled, keel sharpest distally, becoming more blunt proximally. Carpus of major cheliped of male higher than long, length about 60% of height; proximal margin with projection on upper edge for articulation with merus, followed by deep, concave reentrant, then forming convex arc and merging with lower margin which slopes obliquely distally; upper surface nearly straight; distal margin weakly concave. Manus of major cheliped of male rectangular, height about 85% length; proximal margin straight, at about 90 degree angle to lower and upper margins; upper and lower margins nearly straight; distal margin oriented at about 100 degree angle to upper margin, with sharp tooth just above base of fixed finger. Fixed finger apparently straight, short. Movable finger straight, with row of setal pits parallel to occlusal surface, with spine at about midlength that occludes with spine on distal margin of manus. Minor chela of male much smaller than major; apparently with long, slender, fixed finger. Remainder of elements of male unknown or too poorly preserved to be described. Carpus of major cheliped of female higher than wide, poorly known. Manus of major cheliped of female rectangular; proximal margin straight, at about 90 degree angle to lower margin; lower margin weakly convex; upper margin nearly straight; distal margin at about 100 degree angle to upper margin, with spine above base of fixed finger. Fixed finger straight. Minor cheliped of female slightly smaller than major cheliped. Merus of minor cheliped of female longer than high, ovate, with longitudinal keel, upper and lower surfaces apparently entire. Carpus of minor cheliped of female about as long as high; distal margin with short extension on upper edge articulating with merus, followed by concave indentation, followed by abrupt convex extension which curves into rounded lower margin; upper margin nearly straight; distal margin nearly straight, with small, forward-directed projection on lower edge. Manus of minor cheliped of female about as long as high, distal margin at about 100

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degree angle to lower margin, straight; lower margin nearly straight; upper margin weakly convex; distal margin at about 100 degree angle to upper margin. Fingers very long; fixed finger straight, becoming very slender distally; movable finger stouter, narrowing distally. Abdomen preserved in specimen of indeterminate gender with right minor chela. Somite 2 longer than other somites; with pronounced, blunt, rounded projection at distal lower corner overlapping somite 3. Somite 3 shorter than high in lateral view, appearing to have had blunt projection as in somite 2 overlapping somite 4. Somites 4 and 5 similar in shape; with pronounced, blunt, rounded projection at distal lower corner overlapping next somite, projection defined by shallow grooves on anterior and posterior edges. Somite 6 apparently without lateral projections, with rim paralleling proximal margin, rim separated from remainder of somite by wide groove. Telson poorly known; uropods arcuate. Remainder of elements unknown or too poorly preserved to be described.

Measurements.—Measurements (in mm) taken on elements of the major and minor chelipeds of C. haggarti, new species are presented in Table 2. Etymology.—The trivial name honors James Haggart, president of British Columbia Paleontological Association and Geological Society of Canada researcher. Types.—GSC 27156, holotype, and paratypes GSC 27157 and 27158. Occurrence.—The specimens were collected from the Dove Creek Road locality. Discussion.—Comoxianassa haggarti new species is known from both male and female specimens. The single male specimen available possesses a more robust manus and movable finger than do the females. It also has a somewhat smaller minor chela than do the females. It is not uncommon for males to exhibit more marked dimorphism between the two first pereiopods than females in extant callianassids (Ngoc-Ho 2003:487; illustrations in Sakai 2005). The female specimens possess less robust mani of the major cheliped and larger minor chelae than does the male. The minor cheliped of one of the female specimens possesses a merus and carpus that are very similar in shape to those of the male (Fig. 3J), thus, our consideration of specimens with this form as females of the same species as the male specimen. The male specimen exhibits a right major cheliped, and the two females exhibit left major chelipeds. Family Ctenochelidae Manning and Felder, 1991 Genus and species indeterminate (Fig. 5) Material examined.—GSC 27175. Occurrence.—Eden Creek locality. Discussion.—A single specimen of decapod represented by a crushed, partial cephalothorax and a nearly complete abdomen possesses morphology which suggests placement

within the Ctenochelidae. However, callianassoids in the fossil record are almost exclusively represented by chelipeds which are lacking on this specimen. When carapaces and abdomina are present, they tend to be crushed and difficult to interpret in detail so that it seems prudent to assign this specimen to the Ctenochelidae genus and species indeterminate. The specimen from the Comox Formation is moderate in size, total length of partial cephalothorax and abdomen estimated to be 52 mm; the cephalothorax is narrow, smooth, and badly crushed suggesting that it was very weakly calcified; there is no evidence on the specimen of grooves or lineae. The abdomen is somewhat abraded; and arrayed with somites 1–4 lying in a single plane; somites 5 and 6, as well as the telson curve ventrally and anteriorly to lie below the proximal part of the abdomen. Somite 1 is smooth, narrower than the distal somites, and crushed. Somites 2–5 are equally broad, about 8 mm, and approximately equal in length, 6 mm. Each bears an axial and three lateral longitudinal keels, the outermost of which separates the terga from the smooth, short, quadrate pleurae. Somite 6 is abraded, about as long as the preceding somites, tapering in width posteriorly, exhibiting shallow depressions extending antero-ventrally from about midlength on the flanks of the tergum. The telson is quadrate, truncate posteriorly, slightly narrower, 3.8 mm, posteriorly than anteriorly, 4.7 mm, and relatively short, 4.9 mm, compared to the length of the uropods, ca. 6.6 mm. The uropods broaden uniformly distally, have straight distal margins, and keels on the exopod. No diaresis is evident. In terms of characters that can be evaluated on this specimen, the Ctenochelidae are characterized by possession of keels on the exopod of the uropods and a cardiac prominence (Manning and Felder 1991:766). A dorsal plate is absent. By contrast, the Callianassidae lacks a cardiac prominence, have smooth exopods on the uropods, and possess a dorsal plate. There is no evidence of a dorsal plate on the specimen under consideration, the exopods of the uropods are distinctly keeled, and the area of the cardiac prominence is badly crushed and fractured so that its presence cannot be detected. Further, Manning and Felder (1991) indicated that this latter feature is typically, but not always, present. Thus, placement within the family Ctenochelidae seems quite reasonable until additional, more complete material is discovered. The Ctenochelidae have a fossil record extending into the late Eocene (Nagao 1941; Schweitzer and Feldmann 2001) in the form of several species of Callianopsis de Saint Laurent, 1973, known from Japan and possibly northwestern Washington state, U.S.A., respectively. Records of the genus are also known from the Oligocene of western Argentina (Casadío et al. 2004). The Pacific distribution of the fossil ctenochelids and the assignment of a Late Cretaceous specimen to the family suggests an origin of the group in the North Pacific and adds to the growing list of decapod taxa that arose within the Cretaceous and survived the K/T extinction event.

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Fig. 6.—Homolodromiidae (A, B) and Homolidae (C, D). A, Preclarocarcinus parvus, GSC 27176, holotype, dorsal carapace and pereiopods; B, P. parvus, oblique view showing lack of linea homolica, GSC 27176, holotype; C, Latheticocarcinus ludvigseni Schweitzer et al., 2004, GSC 12484, holotype, dorsal carapace; D, L. ludvigseni, GSC 12484, oblique view, arrows indicate linea homolica. Scale bars = 1 cm.

Infraorder Brachyura Linnaeus, 1758 Section Podotremata Guinot, 1977 Superfamily Homolodromioidea Alcock, 1899 Family Homolodromiidae Alcock, 1899 Type genus.—Homolodromia A. Milne Edwards, 1880 Included genera.—Antarctidromia Förster, Ga´zdzicki, and Wrona, 1985 (fossil); Dicranodromia A. Milne Edwards, 1880 (fossil and extant); Eoprosopon Förster, 1986 (fossil); Homolodromia A. Milne Edwards, 1880 (fossil and extant); Palehomola Rathbun, 1926 (fossil); Preclarocarcinus new genus (fossil); Rhinodromia Schweitzer, Nyborg, Feldmann, and Ross, 2004 (fossil). Diagnosis.—The family has recently been diagnosed (Schweitzer et al. 2004), and that will not be repeated here.

Preclarocarcinus, new genus Type species.—Preclarocarcinus parvus, new species, by monotypy. Diagnosis.—Carapace longer than wide, slightly arched transversely and longitudinally, carapace regions well defined, surface finely granular; linea homolica absent, carapace flanks distinct; rostrum downturned; orbits absent; anterolateral spines prominent, extending at least to level of end of rostrum; anterolateral and posterolateral margins gently convex; cheliped (P1) stout, propodus longer than high; carpus and propodus with prominent longitudinal sulcus on outer surface; P5 reduced, dorsal. Etymology.—The generic name is derived from the Latin adjective preclarus = very beautiful, splendid, and the Latinized version of the Greek masculine noun karkinos = crab. The gender is masculine.

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Discussion.—The single specimen herein referred to a new genus and species bears all the characteristics typical of the family. The carapace is longer than wide, and does not possess lineae homolicae. Orbits are absent but there are prominent anterolateral spines. The rostrum is downturned and, although the bifid nature of the termination has not been determined, the variable nature of rostra on other representatives of the family range from bearing a pair of distinct spines to a reduced bifid front that may or may not have an axial spine. These variations were illustrated by Schweitzer et al. (2004: fig. 6). Members of the family have two well-developed transverse grooves and exhibit variable development of the carapace regions; fossil forms tend to have more clearly defined carapace regions. Pereiopod 5 tends to be reduced and is carried in a dorsal position. All of these characters are well exhibited on the new genus. The genus is readily distinguished from other genera within the family in several ways. The outline of the carapace is most like that of Homolodromia; however, Preclarocarcinus is not nearly as vaulted transversely and has more distinctly differentiated lateral flanks. Although the frontal region of Eoprosopon is not well known, it is longer than wide, whereas Preclarocarcinus is equant. Several other members of the family are already known from the Pacific Northwest of North America. Rhinodromia, known from Cretaceous rocks of British Columbia (Woodward 1896), is quite different from Preclarocarcinus in possessing a bifid rostrum and a nodose, bulbous carapace. Palehomola from Oligocene rocks of Oregon (Rathbun 1926) is ovate in shape and overall quite different in ornamentation than Preclarocarcinus. Thus, Preclarocarcinus is unique and readily distinguishable from other genera within the Homolodromiidae. It is important to note that the new genus and species bears a superficial resemblance to another member of the Late Cretaceous fauna of British Columbia, Latheticocarcinus ludvigseni Schweitzer et al., 2004. The carapace outline of the sole specimen of L. ludvigseni is quite similar to that of Preclarocarcinus; however, that outline is defined by the lineae homolicae rather than the true edge of the carapace; the flanks of L. ludvigseni are missing (Fig. 6D). Details of the carapace regions are also superficially similar which has led to past confusion in familial assignment of other taxa and specimens within the Homolodromiidae and the Homolidae de Haan, 1839, in the superfamily Homoloidea de Haan, 1839 (Schweitzer et al. 2004). However, in this case the specimens are sufficiently well preserved to indicate that Latheticocarcinus and Preclarocarcinus are members of two distinct families. Although the Homoloidea are typified by the presence of lineae homolicae, two families within the superfamily Latreillidae Stimpson, 1858, and Poupiniidae Guinot, 1991, lack the structure. However, representatives of these families, each embracing only two genera, are so distinctly different from Preclarocarcinus that they cannot be considered viable families in which the new genus can

be placed. The Latreillidae have a carapace which is very long, pyriform, and drawn out into a long, bifid rostrum. The Poupiniidae exhibit a carapace which widens posteriorly, has distinct cervical and branchiocardiac grooves, and has prominent divergent rostral spines and outer orbital spines. Thus, placement within the Homolodromiidae is strongly supported. Preclarocarcinus parvus, new species (Figs. 6A, B) Diagnosis.—As for genus. Description.—Small homolodromiid, outline rectangular, longer (14.1 mm) than wide (12.1 mm), weakly vaulted transversely and longitudinally; rostrum prominent, downturned; anterolateral spines large; regions well defined. Rostrum broad at base, 26% of carapace width measured about 8 mm from front, axially sulcate and flaring upward laterally, downturned, narrowing distally; termination of rostrum not exposed. Anterolateral spines long, extending to level of end of rostrum, pointed sharply, directed anteriorly. Fronto-orbital width 71% of maximum width. Anterolateral margin smoothly curved to cervical groove. Posterolateral margin straight to approximately midlength of metabranchial region, then curving toward distinct posterolateral corner. Posterior margin convex axially and weakly concave laterally, with thin elevated rim, 8.1 mm wide. Mesogastric region generally narrow, 30% of maximum width at posterior termination; anterior process long, parallel sided, extending onto base of rostrum; posterior element with acute outer corners and weakly convex posterior margin. Metagastric region transversely reniform, widest at axial region, with ovoid swellings laterally and axial depression. Urogastric region narrow, smooth, depressed. Cardiac region pentagonal with apex directed posteriorly, with broad swellings at widest points. Intestinal region not defined. Epigastric regions not clearly defined. Protogastric regions narrow anteriorly, elongate, broadening posteriorly to moderately swollen posterior element. Hepatic region broadly swollen, separated from protogastric region by broad, smooth depression. Cervical groove nearly straight, prominent laterally, becoming obscure and disappearing axially. Epibranchial region transversely triangular, broadest laterally and terminating at level of inner margin of hepatic region. Mesobranchial region swollen, widest axially and tapering to termination near lateral margin. Branchiocardiac groove well developed, weakly concave laterally, extending to bulbous swelling at level of urogastric region, then curving posteriorly to completely encircle cardiac region. Entire surface of carapace finely granular except carapace grooves which are smooth. Granules are clearly distinct on outer surface of cuticle and more subtly expressed on mold of interior of cuticle. Ventral surface and abdomen not preserved. First pereiopod (P1) strong. Merus obscure, appearing to be short. Carpus ovoid, longer than wide; outer surface longitudinally sulcate, with prominent boss at upper articulation with propodus. Manus strong, larger than high, highest at articulation with dactylus; inner and upper surfaces smooth; outer surface convex, with sulcus extending from upper articulation with carpus to boss at articulation with dactylus. Lower surface not exposed. Fixed finger broad, with convex lower surface and straight occlusal margin, denticles not exposed; outer surface with longitudinal keel paralleled by shallow grooves. Dactylus similar in size and form to fixed finger. Termination of fingers missing. Pereiopods 2–4 (P2–P4) long, slender, slightly compressed. Entire length of P3 exceeds 23.2 mm, about twice carapace width; propodus possessing the longest and most slender segment. Last pereiopod (P5) preserved only as small basal segments which lie at posterolateral corners and are directed upward so that P5 would have been carried dorsally.

Etymology.—The trivial name is derived from the Latin

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Fig. 7.—Raninidae. A, Eucorystes platys Schweitzer and Feldmann, 2001, dorsal carapace with good preservation of cuticular nodes, GSC 27167; B, E. platys, dorsal carapace with good preservation of carapace grooves, GSC 27165; C, E. platys, dorsal carapace with unusual preservation suggestive of a molted carapace, GSC 27166; D, E. platys, dorsal carapace with unusual preservation suggestive of a molted carapace and a bopyrid isopod in right branchial region, GSC 27168; E, ?Raninoides sp., fragment of dorsal carapace, right anterior portion including anterolateral spine, outer-orbital spine, and intra-orbital spine, GSC 27164. Scale bars = 1 cm.

parvus, meaning little, in reference to the relatively small size of the species compared to most homolodromiids. Type.—The holotype, and sole specimen, is GSC 27176. Occurrence.—The specimen was collected from the Dove Creek Road locality. Discussion.—As discussed above, the distinguishing features of the species are those of the genus. Of particular note on the specimen of P. parvus is the preservation

of several elements of the pereiopods. They are usually not preserved in fossils. The preserved pereiopods of P. parvus are comparable to those of modern homolodromiids in all regards. The exposed cheliped is moderately stout, longer than high, and the carpus/propodus articulation is positioned such that the claw could be readily directed downward toward the substrate. Examination of a preserved specimen of Homolodromia robertsi Garth, 1973, LACM 80–158.6 (Los Angeles County Museum of Natural History, Research and Collections/Crustacea, CA, USA), documents a similar mode of carrying P1.

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Eucorystes platys Schweitzer and Feldmann, 2001 (Figs. 7, 8)

Fig. 8.—Cuticular surface of Eucorystes platys, GSC 27170. A, plan view, cervical groove, cuticle is present to the left and absent from the right revealing the surface of the interior mold, note that the cervical groove is deeper on the mold of the interior than on the surface of the cuticle; B, oblique photograph of the same cervical groove as Fig. 8A; note increase in cuticle thickness directly above the groove and thinning to the left of the groove.

Comparison of the remaining walking legs of P. parvus with H. robertsi also shows similarities in the long, slender form of P2–P4 and the reduced size and dorsal position of P5. Unfortunately, the terminations of P4 and P5 are not preserved on P. parvus, so it is not possible to determine whether or not they bear tiny chelae. Description of this new genus and species of homolodromiid from Campanian rocks of British Columbia is significant because it brings to three the number of Cretaceous occurrences of the family, two of which are from the Pacific Northwest. In addition to P. parvus, Rhinodromia richardsoni (Woodward, 1896) is known from Queen Charlotte Island, British Columbia (Rathbun 1926; Schweitzer et al. 2004). Feldmann (1993) named Homolodromia novaezelandica from Piripauan-Haumurian (Late Cretaceous) rocks of North Island, New Zealand. Thus, the family had a circum-Pacific distribution as early as the Late Cretaceous. Family Raninidae de Haan, 1839 Subfamily Palaeocorystinae Lo˝renthey in Lo˝renthey and Beurlen, 1929 Eucorystes Bell, 1863 Type species.—Notopocorystes carteri McCoy, 1854. Included species.—Eucorystes carteri (McCoy, 1854); E. eichhorni Bishop, 1983; E. harveyi Woodward, 1896; E. intermedius Nagao, 1932; E. oxtedensis Wright and Collins, 1972; E. platys Schweitzer and Feldmann, 2001. Discussion.—See recent discussions by Schweitzer and Feldmann (2001) and Schweitzer et al. (2003) for reviews of the genus. In an analysis of raninid cuticle, Waugh et al. (in press) suggested that not all species currently referred to Eucorystes may actually be members of the genus.

Description of material with unusual cuticle preservation.—Carapace shape, measurement ratios, and outline same as in previously published material. Entire axis keeled from tip of cardiac region to about mid-mesogastric region. Mesogastric region very well-defined anteriorly, widening distally, poorly defined posteriorly by cervical groove, with small round pit situated on groove marking each lateral margin, pair of pits on posterior margin also. Protogastric region broad, with oblique groove extending from outer orbital fissure toward axis and with two swellings along anterior margin. Hepatic region flattened below level of protogastric region, with prominent tubercle at anterior end. Metagastric region poorly marked, wider than base of mesogastric region, with a small pit along each lateral margin. Urogastric region narrower than metagastric region, bounded laterally by deep, laterally-concave segments of branchiocardiac grooves, confluent with cardiac region. Cardiac region triangular, narrowing distally, weakly defined. Cervical, branchiocardiac, and oblique groove between former two grooves moderately deep.

Material examined.—GSC 27165–27170. Occurrence.—GSC 27165–27168 were collected from the Trent River; GSC 27169 was collected from the Von Ark’s locality; and GSC 27170 was collected from the Puntledge River Fish Hatchery site. Discussion.—Eucorystes platys has been well differentiated from other members of the genus. The specimens examined herein are clearly referable to the species based upon the bifid rostrum, orbital ornamentation, anterolateral spines, and overall shape of the dorsal carapace. What is interesting about the specimens described here is the muted nature of the so-called strap-like ornamentation on the dorsal carapace. On the original type material (Schweitzer and Feldmann 2001: figs. 15, 16) and the subsequently described specimens (Schweitzer et al. 2003: figs. 10.1, 10.2), the straps in the cuticle are clearly observable even though the specimens are missing cuticle in some places and are incompletely preserved. On the material described here, the major grooves are visible (cervical, branchiocardiac) and some of the minor grooves are present (oblique grooves on the protogastric region and between the cervical and branchiocardiac grooves), but the “straps” are not well developed. Under a microscope, the outline of the “straps” is barely visible on the new specimens in most places. This is apparently due to the loss of the exocuticular layers in the new specimens, which has resulted in the loss of much of the exterior ornamentation on the dorsal carapace of this species. The cuticle surface in the new specimens actually appears to be extremely well preserved upon preliminary examination, but closer inspection shows that many of the surface features are damaged and incomplete. These specimens demonstrate the importance of recognizing the variability in ornamentation that the various cuticular layers confer upon a species. The cervical groove is initiated axially at the midline of the carapace by a linear muscle scar that can be seen on the internal mold of the carapace as a depression. The groove continues as a depression without the typical sharp definition of a muscle scar toward the lateral margins. The

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cuticle within the depression is thickened along its axis so that the groove can be seen when cuticle remains on the carapace. The groove is more pronounced and defined when the cuticle is missing, and it can be observed on the exterior of a mold of the interior (Figs. 8A–8B). Two of the specimens are preserved as crushed, fragmented carapaces. Both give the appearance of having had a thinner shell than the other, better preserved specimens. We hypothesize that these specimens may have been molts, lacking much of the calcite, and thus resulting in an easily crushed carapace. One of the crushed specimens exhibits a bopyrid swelling in the right branchial region (Fig. 7D). Such swellings are well-known for the Raninidae (Rasmussen et al. 2008), but this is the first such feature documented for a species of Eucorystes. Eucorystes platys is one of the more common Cretaceous species from Vancouver Island and is abundant in Cretaceous rocks of the area, using terminology of the Paleobiology Database, which “is an international scientific organization run by paleontological researchers from many institutions” and funded by the National Science Foundation in the United States of America (quote from homepage, http://paleodb. org/cgi-bin/bridge.pl). Paleobiology Database terminology for relative abundance in the sediment includes abundant, common, few, and rare. Subfamily Raninoidinae Lo˝renthey in Lo˝renthey and Beurlen, 1929 Raninoides H. Milne Edwards, 1837 ?Raninoides sp. (Fig. 7E) Discussion.—The specimen herein referred to Raninoides is only partially preserved, hence, the questionable designation. The very large, apparently bifid anterolateral spine makes the specimen superficially similar to species of Rogueus Berglund and Feldmann, 1989; however, members of that genus do not have well-developed outer- and intraorbital spines as does the specimen discussed here. Such spines are diagnostic for Raninoides. The outer-orbital spine in the specimen illustrated here is bifid, as in species of Raninoides, and the intra-orbital spine is bounded by open fissures, as is common in Raninoides. However, other species of Raninoides lack long stout, bifid anterolateral spines as seen in the new specimen. Thus, we questionably refer the specimen to Raninoides until more complete specimens can be collected. All known species of Raninoides to date are Cenozoic in age; the oldest is Paleocene (Schweitzer et al. 2006b). If the new specimen were to be confirmed as a member of Raninoides, it would extend the genus into the Cretaceous. Even if the new specimen belongs to a new genus, it appears to be a member of the same subfamily as Raninoides based upon the orbital configuration, which is important in raninid classification. Thus, the subfamily lineage may extend into the Cretaceous.

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Fig. 9.—Nitotacarcinus canadensis, GSC 27163, holotype. Scale bar = 1 cm.

Material examined.—GSC 27164. Occurrence.—New Island Highway locality. Section Heterotremata Guinot, 1977 Superfamily Carpilioidea Ortmann, 1893 Family Tumidocarcinidae Schweitzer, 2005 Nitotacarcinus Schweitzer, Artal, van Bakel, Jagt, and Karasawa, 2007 Type species.—Glyphithyreus bituberculatus Collins and Jakobsen, 2003, by original designation. Other species.—Nitotacarcinus canadensis, new species. Diagnosis.—as in Schweitzer et al. (2007). Discussion.—The type species for Nitotacarcinus, N. bituberculatus, has been well illustrated recently (Collins and Jakobsen 2003; Karasawa and Schweitzer 2004; Schweitzer et al. 2007). The new species lacks features of the sternum which are diagnostic for the family but does have a very well-preserved dorsal carapace, exhibiting diagnostic features for the genus. These include an equant carapace; four anterolateral spines excluding the outer-orbital spine with the last developed as a small swelling; two orbital fissures; long chelae; and a front projecting beyond the orbits. The new species is easily excluded from other genera within the family. Baricarcinus Casadío et al., 2004, Paratumidocarcinus Martins-Neto, 2001, and Tumidocarcinus Glaessner, 1960, each exhibit relatively smooth dorsal carapaces with poorly or undefined carapace regions. The new species has well-developed carapace regions. Lobonotus A. Milne-Edwards, 1864, possesses well-developed cardiac regions with large lateral extensions, which the new species lacks. Pulalius Schweitzer et al., 2000, has a much more highly vaulted carapace and a front with well-developed lobes which the new species lacks. Pulalius lacks orbital fissures, which the new species possesses. Titanocarcinus A. Milne-Edwards, 1864, possesses a bilobed protogastric

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region, exhibits a well-developed last anterolateral spine, and narrows distinctly posteriorly, all of which differentiate it from the new species. Finally, Xanthilites Bell, 1858, lacks orbital fissures entirely and narrows much more posteriorly than the new species. Thus, placement within Nitotacarcinus is made with confidence. Previously, Nitotacarcinus was known only from the lower to middle Eocene (Ypresian/Lutetian) of Denmark. Thus, the new species extends the geographic range considerably. Such a North Polar/North Temperate distribution is rare but occurs for Decapoda in Cretaceous through Eocene time (Schweitzer 2001). The family Camarocarcinidae Feldmann et al., 2008 [imprint 2007], is known to have dispersed from the western interior of North America toward northern Europe in late Cretaceous to Paleocene time (Feldmann et al. 2008 [imprint 2007]). Thus, such a dispersal route is unusual but well documented for decapods. Nitotacarcinus canadensis, new species (Fig. 9) Diagnosis.—Carapace nearly circular, slightly wider than long; four anterolateral spines excluding outer-orbital spine, last a blunt protuberance; regions moderately wellmarked; chelae rather large for carapace size.

Measurements.—Measurements (in mm) taken on the dorsal carapace of N. canadensis, new species: maximum carapace length, 16.0; maximum carapace width, 17.3; fronto-orbital width, 10.9; frontal width, 5.7; length to maximum carapace width, 7.0. Etymology.—The trivial name is derived from the occurrence of the specimen in British Columbia, Canada, the first occurrence of the genus in the western hemisphere. Type.—The holotype, and sole specimen, is GSC 27163. Occurrence.—The specimen was collected from the Eocene Via Appia Beds locality. Discussion.—Nitotacarcinus canadensis differs from the type species in possessing a more rounded carapace; smaller, more blunt anterolateral spines; narrower, shallower grooves; and less tumid regions. Unfortunately, only a single specimen is known so that size ranges or sexual dimorphism cannot be determined. Using the terminology of the Paleobiology Database, of the decapods recovered from the Via Appia Beds, this species is rare, whereas Archaeopus bicornutus Schweitzer et al., 2003, is abundant and Macroacaena chica Schweitzer et al., 2003, is common. ACKNOWLEDGMENTS

Description.—Carapace slightly wider than long, nearly circular in outline, length about 92% of maximum carapace width; widest at position of third anterolateral spine excluding outer-orbital spine, about 44% of the distance posteriorly; carapace weakly vaulted transversely, moderately vaulted longitudinally, especially in anterior half; regions well-marked by grooves on internal mold, more weakly marked where cuticle is preserved. Front projected weakly beyond orbits, bilobed, axially notched; with blunt, broad projections on either side of notch, then sloping obliquely to inner orbital angle; about one-third maximum carapace width. Orbits shallow, semicircular, deepest axially, weakly rimmed, with two closed fissures; outer-orbital angle weakly produced into triangular point; fronto-orbital width about two-thirds maximum carapace width. Anterolateral margins with four spines excluding outer-orbital spines; first spine small, triangular, directed anterolaterally; second spine larger than first, stouter, directed anterolaterally; third spine short but broad at base, directed anterolaterally; last spine a short blunt projection; anterolateral and posterolateral margins about equal in length (8 mm vs. 8.6 mm, respectively). Posterolateral margins convex, entire. Posterior margins nearly straight, rimmed. Protogastric regions trapezoidal, widest anteriorly, moderately inflated. Mesogastric region with long anterior process, widened posteriorly, bounded posteriorly by deep segments of cervical groove; metagastric region short, about as wide as posterior-most mesogastric region; urogastric region slightly wider than metagastric region, depressed below level of metagastric and cardiac regions, with concave lateral margins. Cardiac region pentagonal, moderately inflated, apex directed posteriorly. Intestinal region broadly triangular, flattened. Hepatic region strongly inflated, quadrate. Epibranchial region arcing anteriorly from base of last anterolateral spine and then abruptly arcing posteriorly and terminating at margin of urogastric region. Remainder of branchial region undifferentiated, broadly inflated. Chelipeds heterochelous, right larger, fingers with black tips. Ischia of pereiopods 2–4 longer than high. Remainder of carapace and appendages unknown.

D. Meckert, of Courtenay, British Columbia, and P. Bock, British Columbia, collected and donated some of the specimens for this study; we are grateful for their generosity. R. Lemaitre and K. Reed generously provided access to the Crustacea collection at the United States National Museum of Natural History, Smithsonian Institution, Washington, DC. NSF grant EF-0531670 to Feldmann and Schweitzer funded research at the USNM. J. Martin, Natural History Museum of Los Angeles County, California, U.S.A. provided a specimen of Homolodromia robertsi for comparative purposes. The type specimen of Latheticocarcinus ludvigseni was loaned by B. J. Dougherty, Geological Survey of Canada, Ottawa. Careful reviews of the manuscript were provided by J. W. M. Jagt, Natuurhistorisch Museum Maastricht, the Netherlands, and H. Karasawa, Mizunami Fossil Museum, Japan. We sincerely thank all of these individuals. LITERATURE CITED Alcock, A. 1899. Materials for a carcinological fauna of India, 5: The Brachyura Primigenia or Dromiacea. Journal of the Asiatic Society of Bengal, 68 (II:3):123–169. Bell, T. 1858. A Monograph of the Fossil Malacostracous Crustacea of Great Britain, Pt. I, Crustacea of the London Clay. Monograph of the Palaeontographical Society, London, 10(1856). viii + 9–44 pp., 11 pls. ———. 1863. A Monograph of the Fossil Malacostracous Crustacea of Great Britain. Part II. Crustacea of the Gault and Greensand. Palaeontographical Society Monograph, London, 14(1860). vii + 40 pp., 11 pls. Berglund, R.E., and R.M. Feldmann. 1989. A new crab, Rogueus orri n. gen. and sp. (Decapoda: Brachyura), from the Lookingglass Formation (Ulatisian Stage: Eocene) of southwestern Oregon. Journal of Paleontology, 63:69–73. Beurlen, K. 1930. Vergleichende Stammesgeschichte. Grundlagen, Methoden, Probleme unter besonderer Berücksichtigung der höheren Krebse. Fortschrifte in der Geologie und Paläontologie,

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