Pustulina suevica Quenstedt, 1857

(Figs 13; 14)

Pustulina suevica Quenstedt, 1857: 807, pl. 99, fig. 30. Van Straelen 1925: 289. Beurlen 1928: 200, fig. 24a (non 24b). Schweigert et al. 2000: 9, pl. 5, figs 1-3. Dietl & Schweigert 2001: 53, 54, fig. 87. Schweitzer et al. 2010: 26.

Eryma fraasi Oppel, 1861: 359; 1862: 39, pl. 9, fig. 1.

Enoploclytia perroni Étallon, 1861: 161, pl. 9, fig. 1. Van Straelen 1925: 279. Glaessner 1929: 148. Secrétan 1964: 70. n. syn.

Palaeastacus solitarius Oppel, 1862: 46, pl. 11, fig. 1. Schweitzer et al. 2010: 25.

Eryma pseudobabeaui Dollfus, 1863: 36, pl. 1, figs 1-2. Sauvage 1891: 94, pl. 4, fig. 5 (non 4). Glaessner 1929: 158. Carriol 1991: 225. n. syn.

Enoploclytia edwardsi Sauvage, 1891: 87, pl. 3, figs 1-4. Van Straelen 1925: 282. Glaessner 1929: 146. Schweitzer et al. 2010: 22. n. syn.

Enoploclytia dorsetensis Woods, 1930: 81, pl. 23, figs 1-3. n. syn.

Eryma perroni – Oppel 1861: 358; 1862: 33. Woodward 1900: 9. Whiteaves 1903: 322. Beurlen 1928: 157, 164, 278. Mertin 1941: 160. Charbonnier et al. 2012b: 558, fig. 18E-F.

Eryma suevica – Oppel 1861: 359; 1862: 38, pl. 8, fig. 9.

Enoploclytia pseudo-babeaui – Van Straelen 1925: 280.

Phlyctisoma perroni – Förster 1965: 140; 1966: 141, pl. 18, fig. 6. Taylor 1979: 24.

Phlyctisoma pseudobabeaui – Förster 1966: 141, pl. 18, fig. 6.

Phlyctisoma sp. Förster 1966: 142, pl. 18, fig. 7.

Pustulina edwardsi – Carriol 1991: 224.

Pustulina perroni – Schweitzer et al. 2010: 26.

Pustulina pseudobabeaui – Schweitzer et al. 2010: 26.

TYPE MATERIAL. — Holotype GPIT without number.

TYPE LOCALITY. — Nusplingen, Baden-Württemberg, Germany.

TYPE AGE. — Kimmeridgian.

DESCRIPTION

Carapace

Sub-cylindrical carapace; short, spiny rostrum; dorsal margin of cephalic region curved downward; fusiform intercalated plate; orbital notch slightly curved; inflated cardiac, hepatic and branchial regions; deep cervical groove, almost straight and sub-vertical, joined to dorsal margin and to antennal groove; deep, wide antennal groove; elongated gastro-orbital groove, joined to cervical groove at carapace mid-height, with two divergent branches delimiting two gastro-orbital lobes; deep postcervical groove, inflected at carapace mid-height, strongly inclined dorsally, not joined to dorsal margin; shallow branchiocardiac groove, almost straight, not joined to dorsal margin and interrupted in branchial region; shallow cardiac groove, straight, slightly inclined forward, joined to the postcervical groove and to dorsal margin.

Eryma pseudobabeaui Dollfus, 1863 (Kimmeridgian, Le Havre, France): original figures of Dollfus (1863: pl. 1, figs 1-2); J -K, syntypes of Enoploclytia edwardsi Sauvage,1891 (Kimmeridgian,Boulogne-sur-Mer, France):original figures of Sauvage (1891: pl. 3,figs 1-2). Abbreviations: a1, antennulae; a2, antenna; b, antennal groove; b 1 , hepatic groove; c, postcervical groove; cd, cardiac groove; d, gastro-orbital groove; e, eye; e 1 e , cervical groove; i, inferior groove; ip, intercalated plate; Mxp3, third maxilliped; ne, nephridiopore; sc, scaphocerite. Photographs and line drawing: J. Devillez. Scale bars: 1 cm.

Pleon and uropods

Somites with narrow subtriangular pleurites, with a rounded bulge on their basis.

Cephalic appendages

Last segment of antennular peduncles (basipodite) articulated with two flagella, flagella made of numerous and short cylindrical articles; antennas made of numerous and short cylindrical articles.

Thoracic appendages

Elongated Mxp3, made of cylindrical spineless segments; chelate P 1; P 1 propodus short, as long as wide; narrow, inflated dactylar bulge; short, wide P 1 fingers, straight dorsally, slightly curved downward, equal in length; occlusal margin without teeth; P 1 carpus short, subtriangular; elongated P 1 merus, with a short process at outer side of its ventral extremity; P 2- P 3 chelate.

Ornamentation

Carapace densely covered by rounded tubercles, smaller in ventral part of branchial region and in pterygostomial region; intercalated plate covered by tubercles; row of tubercles parallel to the intercalated plate in gastric region; oblique row of tubercles ended by an orbital spine in gastric region; antennal row of coarse tubercles; tergites of pleonal somites covered by small rounded depressions, mainly on their posterior part; pleurites of pleonal somites covered by small depressions; P 1 propodus densely covered by coarse tubercles organised in longitudinal rows on both ventral and dorsal surfaces; P 1 fingers covered by rounded depressions; P 1 carpus covered by coarse tubercles; P 1 merus covered by small tubercles on dorsal margin, and with a row of spines along the extern ventral margin; smooth P 2- P 5.

DISCUSSION

Because the species was described from a single isolated P 1 chela from the Kimmeridgian of Nusplingen (Germany), Pustulina suevica was recognized as an erymoid lobster only recently. Indeed, Schweigert et al. (2000) pointed out the strong similiraties between the holotype of P. suevica and the P 1 chelae of the holotypes of Eryma fraasi Oppel, 1861 (Fig. 13B) and Palaeastacus solitarius Oppel, 1862 (Fig. 13C), from the same locality. These species share a subrectangular P 1 propodus bearing wide and short fingers, ornamented by longitudinal rows of coarse tubercles. Considering these elements, Schweigert et al. (2000) concluded that E. fraasi and P. solitarius are junior synonyms of P. suevica. Moreover, the carapace of E. fraasi is mostly preserved. This allowed Schweigert et al. (2000) to point out the similarities of the carapace groove pattern of Pustulina and Phlyctisoma Bell, 1863. So, Phlyctisoma became a junior synonym of Pustulina, and we concur with the conclusions of Schweigert et al. (2000).

Étallon (1861) described Enoploclytia perroni from the Oxfordian of Frasne (France; Fig. 13 D-E). In the literature, this species was assigned to Eryma (Oppel 1861, 1862; Woodward 1900; Whiteaves 1903; Beurlen 1928; Mertin 1941; Charbonnier et al. 2012b), Phlyctisoma (Förster 1965; Förster 1966; Taylor 1979) or Pustulina (Schweitzer et al. 2010). Examination of the syntype of E. perroni shows a carapace groove pattern and P 1 chela typical of Pustulina: the elongated gastro-orbital groove with two divergent branches, the concave hepatic groove, the inflected postcervical groove joined to the posterior extremity of the hepatic groove, the presence of the cardiac groove, the short and subrectangular propodus with its short and wide fingers. The ornamentation of E. perroni made of coarse rounded tubercles, becoming smaller ventrally, is also identical to that of P. suevica. So, we consider E. perroni as another junior synonym of P. suevica. For the same reasons than E. perroni, Eryma pseudobabeaui Dollfus, 1863 (Kimmeridgian, Le Havre, France; Fig. 13H, I), Enoploclytia edwardsi, Sauvage, 1891 (Kimmeridgian, Boulogne-sur-Mer, France; Fig. 13J, K) and Enoploclytia dorsetensis Woods, 1930 (Oxfordian, Weymouth, United Kingdom; Fig. 13F, G) are considered as junior synonyms of P. suevica. We noticed that E. edwardsi has already been considered as a synonym of Phlyctisoma pseudobabeaui and E. dorsetensis as a synonym of P. perroni by Förster (1966).

Among the crustacean faunas of the Oxfordian-Kimmeridgian, Pustulina suevica is a typical species with a wide distribution in Western Europe. Fossils of this species are reported from many localities in France, Germany and United Kingdom. Moreover, it is the only erymoid species from the lithographic limestones of southern Germany to have been reported out of this country.

P. suevica is distinct from some other species of the genus by its ornamentation. Indeed, there are no depressions as in P. calloviensis (Förster, 1966) and P. elegans (Förster, 1966), the tubercles of P. suevica are coarser than those of P. trisulcata (Schweitzer & Feldmann, 2001), there are no small tubercles between the coarse ones as in P. tuberculata (Bell, 1863), and there is a strong antennal row in P. suevica contrary to P. calloviensis, P. cretacea (Roger, 1946), P. elegans, P. minuta, and Pustulina victori Devillez, Charbonnier, Hyžný & Leroy, 2016. Pustulina suevica is also the only species with P. elegans to have an oblique orbital row. Like P. minuta, the cervical groove of P. suevica is almost straight contrary to P. calloviensis, P. colossea Devillez, Charbonnier, Hyžný & Leroy, 2016, P. elegans, and P. occitana Devillez, Charbonnier, Hyžný & Leroy, 2016. The gastro-orbital groove is also sub-horizontal while that of P. calloviensis, P. colossea, P. elegans, P. occitana, P. spinulata (Secrétan, 1964), and P. tuberculata is oblique. Moreover, the P 1 fingers of P. suevica are sometimes ended by a hook contrary to P. minuta and P. tuberculata. Finally, the pleurites of the pleon are narrower than that of P. minuta and P. trisulcata.