Necrocarcinus labeschii ( Deslongchamps, 1835 )

Guinot, Danièle, Vega, Francisco J. & Van Bakel, Barry W. M., 2008, Cenomanocarcinidae n. fam., a new Cretaceous podotreme family (Crustacea, Decapoda, Brachyura, Raninoidia), with comments on related families, Geodiversitas 30 (4), pp. 681-719 : 681-719

Necrocarcinus labeschii ( Deslongchamps, 1835) ( Fig. 8E, F )

Orithyia labeschii Deslongchamps, 1835: 40 , pl. 1, figs 7, 8.

MATERIAL EXAMINED. — Cenomanian, Le Mans, Sables du Perche Formation, 1 specimen (carapace length 19 mm; width 24 mm) ( MHN LM 3808, Guéranger Colln) (see Breton & Collins 2007). — Albian, Cambridge, 5 carapaces ( MNHN). — Lower Cenomanian, Mantelliceras dixoni Zone , France, Normandy, Seine- Maritime, Pétreval, at Annouville-Vilmesnil, 1 carapace, G. Breton coll. (G. Breton Colln). — Middle Albian, top of Dimorphoplites niobe Zone , France, Pas-de- Calais, Wissant, P. Destombes coll., J. S. H. Collins det. 10.IX. 1981, 2 carapaces (G. Breton Colln).

OCCURRENCE. — Cretaceous (Albian-Cenomanian) ( Glaessner 1929a, 1969; Wright & Collins 1972; Ilyin & Alekseev 1998; Breton & Collins 2007). The earliest known N. labeschii is from the lower Albian, basal zone, Cauville, Seine-Maritime, France (G.Breton pers. comm.).

Remarks

Necrocarcinus labeschii , from the Cenomanian of Normandy,has been described and figured by many authors including Bell (1858: 20, pl. 4, figs 4-8), Carter (1898: 27, pl. 1, fig. 9), Glaessner (1969: R495, fig. 306.3), Wright & Collins (1972: 63, pl. 11, figs 1-8, pl. 22, fig. 8a-c), Förster (1968: 173, fig. 3, pl. 13, figs 7, 8; 1970: 136, 142, figs 5b, 6b), Ilyin & Alekseev (1998: fig. 2a), Breton & Collins (2007: 18, fig. 6). No illustrations, however, show the sternum or legs. The male abdomen was described but not figured ( Carter 1898: 27, 28; Wright & Collins 1972: 64), there is however some doubt whether or not it was attached to a carapace. The female specimen with abdomen preserved, figured by Carter (1898: pl. 1, fig. 9), “has not been traced” according to Wright & Collins (1972: 63).

Necrocarcinus bodrakensis Levitski, 1974 , upper Albian of Crimea, has been synonymised with N. labeschii ( Ilyin & Alekseev 1998: 594) .

REMARKS ON THE STATUS OF SEVERAL SPECIES PREVIOUSLY ASSIGNED TO NECROCARCINUS

The status of several necrocarcinid species is reevalu-

ated herein, based generally on carapace characters

only and in comparison to N. labeschii (supposed to be a podotreme crab), and generally in the absence of any ventral characters.

Necrocarcinus woodwardii Bell, 1863 ( Bell 1863: 20, pl. 4, figs 1-3)

From the upper Albian to the lower Cenomanian of England, northern France and Germany ( Carter 1898: 29, pl. 2, fig. 1; Lőrenthey 1929: 300, fig. 22a; Förster 1968: 175; Wright & Collins, 1972: 65, pl. 12, figs 1, 2; Feldmann et al. 1993: 36; Fraaye 1994: 264, fig. 1; Schweitzer et al. 2003a: table 3), it can be assigned to Necrocarcinus .

Necrocarcinus undecimtuberculatus Takeda & Fujiyama, 1983 ( Takeda & Fujiyama 1983: 133, fig. 3, pl. 1, figs 1, 2)

From the upper Aptian of northern Japan ( Fraaye 1994: 264; Schweitzer et al. 2003a: 33, table 3), N. undecimtuberculatus differs from N. labeschii by lacking tubercles on the mesogastric and urogastric regions. This species, which would be better assigned to Paranecrocarcinus Van Straelen, 1936 , is the oldest known necrocarcinid.

Necrocarcinus texensis Rathbun, 1935 ( Rathbun 1935: 45, pl. 11, figs 20-22)

From the upper Albian of Texas ( Beurlen 1958: fig.1a; Förster 1968: 174; Bishop & Williams 1991: 458, fig. 6A; Schweitzer et al. 2003a: table 3), with three longitudinal ridges surrounded by a few large tubercles, it is a possible Necrocarcinus .

Necrocarcinus tauricus Ilyin & Alekseev, 1998 ( Ilyin & Alekseev 1998: 592, figs 1a, 2b)

From the upper Albian of southwest Crimea, N. tauricus is very close to N. labeschii , and its status is not in doubt (see also Ilyin 2005: 201).

Necrocarcinus olsonorum Bishop & Williams, 1991 ( Bishop & Williams 1991: 452, figs 1-5, 6B) From the Turonian of the Western Interior ( USA), N. olsonorum is distinguished by its highly arched carapace, dorsal surface with rounded boss-like spines and small granules, and lateral margins bearing five projecting bosses (see Fraaye 1994:

264, fig. 1; Schweitzer et al. 2003a: table 3). Its

configuration indicates necrocarcinid affinities as stated by Bishop & Williams (1991: 458).

Necrocarcinus carinatus Feldmann, Tshudy & Thomson, 1993 ( Feldmann et al. 1993: 36,

fig. 29.6)

From the lower Campanian of James Ross Basin, Antarctica ( Fraaye 1994: 264, fig. 1; Schweitzer et al. 2003a: table 3), with its well-developed axial keel and spinous lateral keels, N. carinatus is fairly close to N. tricarinatus according to Feldmann et al. (1993: 37) but differs by outline and ornamentation of the gastric and epibranchial regions. Schweitzer et al. (2003a: 39, table 3), by listing N. carinatus among the Necrocarcinus species, regarded it as a necrocarcinid, whereas they recognised Orithopsis tricarinata as a dorippoid. We suggest placement in Orithopsis (see under Orithopsis ).

Necrocarcinidae gen. et sp. indet ( Schweitzer et al.

2003b: 889, fig. 1, as indeterminate genus and species)

The unnamed mangrove-dwelling crab from the Cenomanian of Egypt assigned to the Necrocarcinidae , thus assumed to be a dorippoid by Schweitzer et al. (2003b), has two extremely elongate pereiopods (probably P2 and P3), a much shorter P1, with a stout chela. Despite the absence of other informations concerning P4 and P5 and the venter, we agree with the hypothesis that this fossil could belong to the Dorippoidea.

Necrocarcinus bispinosus Segerberg, 1900 ( Segerberg 1900: 372, pl. 9, fig. 7)

From the Danian of Denmark and Sweden, N. bispinosus was considered to belong to Campylostoma ( Fraaije 2002: 913) (see under Campylostoma ).

Necrocarcinus rathbunae Roberts, 1962 ( Roberts 1962: 181, pl. 85, fig. 12, pl. 87, figs 1, 2) From the lower Campanian of New Jersey ( Bishop& Williams1991:452, fig.6E; Schweitzer et al. 2003a:33, 39, table 3), N. rathbunae was considered of uncertain position according to Fraaye (1994: 264, fig. 1).

Necrocarcinus senonensis Schlüter, 1868 (in von

der Marck & Schlüter 1868: 297, pl. 44, fig. 3)

Cenomanocarcinidae n. fam. ( Crustacea, Decapoda ) from Cretaceous

(senior synonym of N. insignis Segerberg, 1900: 372, pl. 9, figs 1, 6)

From the Danian of Sweden, Denmark and central Poland ( Segerberg 1900:26, pl. 9, figs 2, 3, 5; Förster 1968: 175, pl. 13, fig. 4; Fraaye 1994: 262, fig. 1, pl. 1; Fraaije 2002: 913, 914; Schweitzer et al. 2003a: table 3; van Bakel et al. 2005: 286), N. senonensis is now better known thanks to a specimen from the middle Danian of Fakse, Denmark, figured by Collins & Jakobsen (1995: 39, pl. 10, fig. 11). It shows a strong epibranchial spine (partially broken), a pointed subdistal posterolateral tooth (well visible in Segerberg 1900: 372, pl. 9, fig. 6), and ornament of rather large tubercles. The same Danish specimen figured here ( Fig. 9A ), which is a reconstruction, shows on the lateral margins two long and subequal spines, a unique configuration which does not conform to Necrocarcinus . The species needs to be re-evaluated, and a re-examination of material of “ Necrocarcinus ” senonensis and “ N.” cf. senonensis of Mertin (1941: 239, fig. 27) from the Santonian- Campanian of Germany is called for.

Necrocarcinus angelicus Fraaije, 2002 ( Fraaije 2002: 914, figs 1.1, 2)

From the upper Maastrichtian of the Netherlands, N. angelicus is closely related to Orithopsis tricarinata , and we propose to include it in that genus as O. angelica (see under Orithopsis , below).

Necrocarcinus pierrensis Rathbun, 1917 ( Rathbun 1917: 389, pl. 33, figs 4, 5, as Campylostoma )

From the upper Campanian-lower Maastrichtian of the Western Interior ( USA) ( Rathbun 1935: 45, pl. 12, fig. 5; Bishop 1981: 387, 388, 391, fig.13.4B; 1984: figs 8, 13; 1985: 621, figs 11, 12.1; 1986: 135, fig. 9B; Bishop & Williams 1991: 452, 458, fig. 6D; Fraaye 1994: 263, fig. 1; Fraaije 2002: 914; Schweitzer et al. 2003a: 33, 34, 36, table 3).

Necrocarcinus davisi Bishop, 1985 ( Bishop 1985: 619, figs 3.6, 10-12)

Necrocarcinus davisi comes from the lower-middle Campanian of South Dakota ( Bishop 1986: 135, fig.9I; Bishop & Williams 1991:fig.6C; Fraaye 1994:

263, fig. 1; Schweitzer et al. 2003a: table 3).

Guinot D. et al.

Necrocarcinus pierrensis and N. davisi are close by having an elongate carapace, with spiniform lateral borders (very strong, complete spines in N. pierrensis ), a concave posterior border, and spinous dorsal ornamentation. It should be noted that N. pierrensis and N. davisi , known only by their dorsal carapaces, evoke the Recent Orithyia Fabricius, 1798 . This monotypical, primitive heterotreme genus (type species: Orithyia sinica (Linnaeus, 1771) ; see Hartnoll 1971: fig. 2A, pl. 1, fig. c), which deserves its own family, the Orithyiidae Dana, 1852 , has been the subject of a long debate, and is often referred to the Dorippoidea, or considered to be related (Guinot, Tavares & Castro in study).

With respect to its keeled spines on both the antero- and posterolateral margins, “ N. ” pierrensis (as well as N. davisi , probably) is distinct from Necrocarcinus emend. According to Schweitzer et al. (2003a: 33) the apparent similarity between “ N. ” pierrensis and Camarocarcinus Holland & Cvancara, 1958 (and Cristella Collins & Wienberg Rasmussen, 1992 ) has led to independent speculation that the later genera may be derived from “ N. ” pierrensis ( Bishop & Williams 1991: 458; Collins & Wienberg Rasmussen 1992: 38; Fraaye 1994: fig. 1; Fraaije 2002: 914) (see under Camarocarcinus ).

Necrocarcinus wrighti Feldmann, Tshudy & Thomson, 1993 ( Feldmann et al. 1993: 35,

figs 29.1-29.5, 30; see Fraaye 1994: 264, fig. 1; Schweitzer et al. 2003a: table 3)

From the lower Campanian-Maastrichtian of Antarctica, N. wrighti shows developed rostral spines, an elongate ischium and wide mxp3 exopod, which match Necrocarcinus emend. In N. wrighti , the rostrum which consists of three teeth (the median one with only a shallow sulcus), the longer and more slen- der postorbital spine, and smaller and more numerous spines on the gastric region are distinctive from N. labeschii . Fronto-orbital spines are reminiscent of Orithopsis (see under Orithopsis ). The generic status of “ N. ” wrighti needs to be reevaluated.

The partially preserved and slightly displaced thoracic sternum of “ N. ” wrighti shows anterior sternites (damaged) as a reversed V-shaped plate, sternite 4 as a long and wider plate, sternite 5 as a wider plate,

sternites 4 and partly 5 hollowed by a rather deep depression ( Feldmann et al. 1993: 35-36, figs 29.4, 29.5), the sutures being only lateral. This rather deep and undivided median portion corresponds to a podotreme sterno-abdominal depression (rather than to a eubrachyuran sterno-abdominal cavity). Anyway, the sternal configuration of “ N.” wrighti does not match any known podotreme (or eubrachyuran) condition and represents the structure of an extinct family (see under Camarocarcinus ).

Figures

FIG. 8. — A-D, “Necrocarcinus” renfroae Stenzel, 1945, Early Cretaceous, upper Albian, Texas, Pawpaw Formation; A, B, paratype (BEG-21094-A3), frontal and dorsal views; C, D, holotype (BEG-21093-A3), frontal and dorsal views; E, F, Necrocarcinus labeschii (Deslongchamps, 1835), Albian, Calais, Escalles, northwest France (B. van Bakel Colln), left chela and carapace. Scale

FIG. 9.— A, “Necrocarcinus” senonensis Schlüter in von der Marck & Schlüter 1868,middle Danian,Denmark,Fakse quarry,reconstructed photograph of the carapace (Collins & Jakobsen 1995: pl. 10, fig. 11), with two strong lateral spines; B, D, Camarocarcinus quinquetuberculatus Collins & Wienberg Rasmussen,1992, holotype, probably male (cast, MGUH 21609;courtesy of S. L. Jakobsen), dorsal and ventral views; C, Cretacoranina cf. dichrous (Stenzel, 1945) (Palaeocorystidae), middle Cenomanian, Karai Formation, between Odiyam and Kunnam,Tamil Nadu, India,leg.A.S.Gale, 1999 (OUM KY.2861), thoracic sternum, see individualised sternite 3, lateral expansions of sternite 4,and deep sutures 4/5; E, F, Orithopsis tricarinata (Bell,1863);E, upper Albian,Upper Greensand,England,Cowstones,holotype of O. bonneyi Carter, 1872, “unretouched photograph of specimen prepared since the original illustration” (Wright & Collins 1972: pl. 12, fig.3) (SM B58557);F, upper Albian,Cambridge Greensand (KBIN,Van Straelen Collection,unnumbered),thoracic sternum,with preserved