Oncopareia bredai Bosquet, 1854

Tshudy, Dale M., Hyžný, Matúš, Veselská, Martina Kočová & Jagt, John W. M., 2022, Taxonomic revision of the extinct clawed lobster genus Oncopareia Bosquet, 1854 (Decapoda, Astacidea, Nephropidae), Palaeontologia Electronica (a 20) 25 (2), pp. 1-31 : 9-15

Oncopareia bredai Bosquet, 1854

Figures 1D , 2A , 3A–N , 4A–E , 5A–L , 6A–C , 7A–K partim 1854 Oncopareia bredai Bosquet , p. 128, pl. 10, figs. 5, 61 and 8 [non pl. 10, figs. 6, 7 and 71 a, b = Hoploparia beyrichi ].

1862 Nymphaeops coesfeldiensis Schlüter , p. 728, pl. 13, figs. 3, 6.

1868 Nymphaeops coesfeldiensis Schlüter ; Schlüter in von der Marck and Schlüter, p. 295.

1868 Enoploclytia paucispina Schlüter in von der Marck and Schlüter, p. 303, pl. 44, fig. 6.

1879 Nymphaeops coesfeldiensis Schlüt. ; Schlüter, p. 597, pl. 15, fig. 1, 2.

1886 Homarus bosqueti Pelseneer , p. 166 [6], fig. 4.

partim 1887 Hoploparia biserialis Fritsch in Fritsch and Kafka, p. 35, pl. 5, fig. 2

[non text-fig. 56; pl. 3, fig. 5, pl. 5, fig. 1 = Hoploparia biserialis ].

1929 Nymphaeops coesfeldiensis Schlüter ; Glaessner, p. 279.

1941 Oncopareia bredai Bosquet ; Mertin, p. 183, fig. 9 [cephalothorax, but not pleon, and necessarily rostrum]

?1941 Oncopareia coesfeldensis [sic] (Schlüter); Mertin, p. 182, text-figs.

9a,?9b, 10b-d, k-m; pl. 3, figs. 2-7.

?1941 Oncopareia cf. coesfeldensis [sic] (Schlüter); Mertin, p. 187, pl. 3, fig.

9.

1964 Ctenocheles coesfeldiensis (Schlüter) ; Secretan, p. 152.

1990 Oncopareia bredai Bosquet ; Feldmann et al., p. 26, text-fig. 4; pl. 1, figs. 4–6.

2000a Oncopareia bredai Bosquet ; Tshudy and Sorhannus, p. 484, figs. 3, 4.

2000b Oncopareia bredai Bosquet ; Tshudy and Sorhannus, fig. 2.2.

2010 Oncopareia bredai Bosquet ; Schweitzer et al., p. 31.

2010 Oncopareia coesfeldiensis (Schlüter) ; Schweitzer et al., p. 31.

2016 Oncopareia bredai Bosquet ; Feldmann et al., p. 23.

2016 Oncopareia coesfeldiensis (Schlüter) ; Feldmann et al., p. 23, fig.

20.3b.

Type material. IRScNB 90-33f (lectotype; specimen consisting of cephalothorax, partial pleon, left

[crusher] palm, and basis through merus of right claw), IRScNB 90-3b ( Tshudy, 1993, fig. 39.2;

incomplete cheliped); IRScNB 90-15a, b ( Tshudy,

1993, fig. 38.4; cephalothorax fragment), IRScNB

90-17 ( Tshudy, 1993, fig. 38.5; cephalothorax),

IRScNB 90-18, 90-19b, c, g, i, k, q ( Tshudy, 1993,

fig. 39.5-8; incomplete cheliped), IRScNB 90-22a,

c, 90-23b ( Tshudy, 1993, fig. 38.6; partial pleon),

IRScNB 90-26a, 90-27a, b ( Tshudy, 1993, fig. 39.3;

b: complete cheliped), IRScNB 90-28a, b ( Tshudy,

1993, fig. 39.1; incomplete pair of chelipeds),

IRScNB 90-32a, c, e (a: cheliped; c: cephalothorax

[ Tshudy, 1993, fig. 38.1]), IRScNB 90-34, 90-35a, c

( Tshudy, 1993, fig. 38.2-3; pleon) and IRScNB 90-

36a, 90-37 and 90-40.

Bosquet (1854, pl. 10, figs. 5–8) did not explicitly select a type when erecting Oncopareia bredai , but did illustrate five specimens, all unregis-

tered. However, he did cite collections by name. As Bosquet did not designate type specimens, his illustrated specimens were not identified by museum/specimen numbers and none of the specimens that we have examined could be confidently matched up with his illustrations, it seems best to designate a lectotype for Oncopareia bredai . Specimen IRScNB 90-33f ( Figure 3G here), from the “type series” of O. bredai (sensu Bosquet, 1854) , according to the label accompanying the specimen, is designated lectotype here. The specimen consists of a cephalothorax, partial pleon, left (crusher) palm, and basis through merus of the right claw. This specimen serves well as the lectotype because it includes a cephalothorax and its associated, distinctive, Thaumastocheles -like pleon.

Additional material. Four specimens from the upper lower Maastrichtian ( Gulpen Formation , Vijlen Member, interval 6) of Haccourt, north-east Belgium , originally described by Feldmann et al. (1990), i.e., a near-complete cephalothorax and pleon with fragments of appendages ( USNM 444296 ), an incomplete pleon with tailfan ( USNM 444297 ), an incomplete cephalothorax with fragments of pleon and appendages ( USNM 444298 ), as well as a fragmentary cephalothorax and chelipeds ( USNM 444299 ). From the same locality and stratigraphical level come seven additional specimens ( Figure 5 ), preserving partial cephalothoraxes, pleons, claws, and partial tailfans ( NHMM JJ 6741 , JJ 6764 , JJ 6839 , JJ 7194 , JJ 14618 , JJ 15534 , and JJ 15836). One claw ( NHMM JJ 5932 ) is from the Zeven Wegen Member ( Gulpen Formation , late Campanian) at the CBR-Lixhe quarry (Lixhe, north-east Belgium) . One specimen, a cephalothorax with associated pleonal segments 1 and 2 and both claws ( MNHN R03408 ; Figure 4 ) is from the upper Maastrichtian (Kunrade Formation) at Kunrade, the Netherlands . Two syntypes of Hoploparia biserialis Fritsch , in Fritsch and Kafka, 1887 ( NM O3470 and NM O6861 ) originated from the Turonian of Vinary , Czech Republic , and representing isolated cephalothoraxes ( Figure 6 ); NM O3470 was figured by Fritsch and Kafka (1887, pl. 5, fig. 2). These two specimens are removed from the type series of H. biserialis herein (see below for further details).

The only material referred to Nymphaeops coesfeldiensis ( Figure 7 ) that we have studied in person includes a few fragmentary specimens of pectinate and crusher claw ( NHMM HB 393 and HB 756; Figure 7J–K ) from the Vaals Formation (lower Campanian) of Haccourt, north-east Belgium, recorded by Jagt and Bongaerts (1986). In addition, there is a single, near-complete cephalothorax, with associated appendages ( NHMM JJ 9974), from the same locality and stratigraphical level ( Figure 7I ). In this respect, it is of note that Bosquet (1860, 1868) listed, but did not illustrate, O. bredai , from the so-called ‘Hervien’ in the Liège-Limburg Basin. This corresponds nowadays with the Vaals Formation of early Campanian age. We have attempted to borrow Schlüter’s 1862 type specimen and other individuals from the ‘Akademie zu Münster’, but these may have fallen victim to air raids during World War II. We are aware of larger suites of specimens of nephropid lobsters contained in the Karl-Heinz Hilpert Collection at the Ruhrmuseum (Essen, Germany; see Jagt et al., 2004; Van Bakel et al., 2005) that may include topotypical material of Nymphaeops coesfeldiensis . A detailed assessment of this material is deferred to another occasion.

Diagnosis. Oncopareia with pleura having prominent, rounded, marginal carina; intercervical groove extending toward below upper end of cervical groove; pectinate claw denticles perpendicular to longitudinal axis of fingers.

Emended description. Carapace laterally compressed, twice as long as high. Postcervical groove intersecting dorsal median line slightly posteriorly to mid-length on carapace. Postcervical groove long, well incised dorsally and laterally; extending ventrally over dorsal surface, then, as combined branchiocardiac-postcervical groove, angling anteroventrally to hepatic groove. Hepatic groove deeply incised, joining cervical and antennal grooves in semicircular loop. Cervical groove originating at level of base of orbit, paralleling postcervical groove except at upper end of cervical groove, which is deflected anteroventrally. Prominence “ω” distinct. Intercervical groove narrow, extending anteroventrally from postcervical groove, at level just above cervical groove origin, toward, but not reaching, lower portion of cervical groove; intercervical groove terminating at level of midheight on cervical groove. Cervical groove and antennal groove together forming a smooth, semicircular arc. Antennal groove vague anterior to prominence “ω”; curving toward antennal spine, effacing well before reaching spine. Branchiocardiac carina broad, subtle; extending from near posterior margin to postcervical groove, curving downward near groove. Antennal region flat, featureless. Very shallow branchiocardiac groove present; most evident near postcervical groove. Posteromarginal carina wide, smooth, with a few pits. Cephalic region granulose, with a row of setal pits extending from top of cervical groove to near antennal spine. Antennal spine strong; supraorbital and postorbital spines slightly smaller. Thoracic region pitted dorsally, with granules generally sparse laterally but dense ventrally.

Pleonal terga wide, convex. Each segment with lateral carina at tergum-pleuron boundary. Pleura short; at least pleura 2 and 3 with blunt ventral margin.

Tergum 2 rectangular. Anterior margin straight, posterior margin subtly convex backward, lateral margins subtly convex, locally indented near posterior margin. Anterior margin depressed laterally; depressed area progressively widening toward pleuron. Tergum coarsely and densely pitted. Lateral carina prominent, rounded, widest anteriorly. Pleuron trapezoidal, convex axially; upper surface coarsely pitted, lower surface smooth.

Terga 3–6 of similar width. Tergum 3 rectangular. Anterior margin slightly concave forward, lateral margins slightly convex, widest at mid-length, subtly indented locally near posterior margin. Anterior margin depressed laterally. Tergum coarsely, densely pitted except on smooth dorsomedian keel, depression along anterior margin, and in subtle furrow adjacent to upper limit of lateral carina. Summit of lateral carina well defined, extending obliquely ventrally posteriorly. Pleuron trapezoidal; ventral and posterior margins shorter than dorsal and anterior margins. Pleuron concave overall; margins convex. Surface mostly smooth, granulose ventrally. Surface locally swollen at anteroventral corner, maybe with spine.

Tergum 4 trapezoidal, wider anteriorly than posteriorly. Lateral margins convex along most of length; straight for a short distance posteriorly. Tergum coarsely and densely pitted except on smooth dorsomedian keel. Lateral carina extending from near anterior margin to posterior margin, curving posteroventrally over anterior half and extending on level over posterior half. Upper surface of carina very broad, inflated. Carina summit rounded, distinct. Pleuron margins convex, with sharply rounded, posteroventrally directed termination. Upper surface almost smooth, lower surface granulose. Small, dome-like swelling anteroventrally.

Tergum 5 rectangular. Lateral margins convex. Tergum smooth on dorsomedian keel, otherwise coarsely and densely pitted. Lateral carina as on segment 4. Pleuron round. Small, dome-like swelling anteroventrally. Pleuron upper surface granulose, lower surface densely granulose. Pleural margins 4–5 with very narrow, rounded, raised margin.

Sixth tergum trapezoidal, much wider anteriorly. Lateral margins slightly convex. Tergum subtly convex, densely and coarsely pitted, sparsely so dorsoventrally. Tergum dropping off sharply laterally. Posteriorly, tergum concave laterally between subtly convex dorsum and drop off. Pleuron triangular, very short, highest near anteroventral corner. Small, dome-like swelling anteroventrally. Pleuron granulose. Telson with round granules.

Claws strongly dissimilar in form; relative size of left and right claw varying. Crusher claw very Hoploparia -like; delicate pectinate claw with bulbous palm, long fingers and acicular dentition much like that of extant Thaumastocheles .

Pectinate claw with bulbous palm approximately twice longer than thick. Large process on at least one side of palm receiving spine at base of dactylus. Palm surface with minute granules. On at least one surface of palm, spine distally near base of fixed finger. Fingers long, slender, with acicular denticles of varying length, of formula ADCDBD- CDA (“A” being largest), in one plane. Fixed finger nearly straight, widest at base; width tapering very gradually distally. Dactylus slightly curved; widest over proximal portion near mid-length. Fingers proximally triangular in cross section; otherwise compressed. Tips of fingers not occludent; one slightly shorter than other. Most denticles straight, angling slightly distally. Some larger denticles slightly curving, angling distally at base, proximally at end. Surface of both fingers pitted.

Crusher palm shape varying between specimens. On most, left palm nearly twice as long as wide. Palm widening distally over proximal half; margins parallel over distal half. Palm compressed, slightly thicker proximally; upper and lower surfaces gently convex. Palm surface variously granulose, pitted. Upper and lower palm surfaces very subtly concave at base of fixed finger. Both surfaces with spine submedially, near base of fixed finger. Submedial process on upper and lower distal margin articulating with spines on dactylus base. Inner margin somewhat flattened, with a few alternating spines on upper and lower edges. Outer margin rounded; axis with band of setal pits; upper surface of outer margin with irregular row of spines directed slightly distally. Fingers moderately strong, ovate in cross section. Fixed finger occlusal surface with multi-cusped platform proximally; otherwise with blunt, conical denticles. Dactylus with upward and distally-directed spine proximally on corner of upper/non-occluding surfaces. Dactylus dentition conical; pattern varying between specimens. Merus (IRScNB 90-17 and 90-33f) large, strongly compressed laterally, widening distally; upper proximal corner terminating in long point. Upper palm surface variously pitted, granulose.

On two specimens (IRScNB 90-33f and 90-32a; latter with small, Thaumastocheles -like claw), crusher claw palm proportionately shorter than on other specimens, and widening distally over entire length, giving subtriangular outline; upper and lower surfaces more convex than on other specimens; palm thickens distally.

Remarks. Pelseneer (1886) was the first to note the Thaumastocheles -like morphology in Bosquet’s type series. He examined both the Bosquet and Ubaghs collections but did not recognize Bosquet’s combining of the two lobster species. He did, however, realize the uniqueness of one of the Thaumastocheles -like pleons and referred to this as a new species, Homarus bosqueti . Pelseneer (1886) correctly described the terga as being less convex than those of O. bredai , but his figures 3 and 4 show the opposite. Homarus bosqueti is considered herein to be a junior objective synonym of Oncopareia bredai .

A re-examination of the syntype series of Hoploparia biserialis has revealed that two isolated cephalothoraxes assigned to this species by Fritsch himself (one of them being figured in Fritsch and Kafka, 1887, pl. 5, fig. 2), belonged to a different genus. Indeed, there is an imminent difference in the groove pattern between the two cephalothoraxes assigned to H. biserialis shown in Fritsch and Kafka (1887, pl. 5, figs. 1 and 2), one of them belonging to H. biserialis and another being reassigned to O. bredai herein. We were not able to identify any morphological differences between the type material of O. bredai and the Czech material here attributed to that species.

The type specimen of Nymphaeops coesfeldiensis has not been located; it is here considered lost. Our examination of the original figures ( Schlüter, 1862, pl. 13, figs. 3, 6; Schlüter, 1879, pl. 15, figs. 1, 2, refigured herein in Figure 7 ) has not revealed any difference from the material of O. bredai leading us to recognize N. coesfeldiensis as a junior subjective synonym of O. bredai . In this respect, it is important to note that Mertin (1941) lumped material from several ‘Senonian’ localities into one species, i.e., O. coesfeldiensis . Whether or not all of the specimens Mertin (1941) assigned to O. coesfeldiensis are correctly referred to one species is uncertain. His unwhitened photographs are difficult to interpret and his line drawings do not seem to be consistent with the photographs. For example, he did not connect the branchiocardiac and cervical grooves ventrally (as hepatic groove) in his figure 9a. Cephalothoraxes and pleons in his plates strongly resemble those of O. bredai as revised herein. Mertin (1941, p. 187, pl. 3, fig. 9) also listed Oncopareia cf. coesfeldiensis from the lower ‘Senonian’ of north-central Germany and Oncopareia sp. ( Mertin, 1941, p. 187, text-fig. 10e, f and n) from the upper ‘Senonian’ of the same area. The latter shows heterochelous claws, the pectinate claw having acicular dentition.

Oncopareia bredai differs from its congeners by the intercervical groove extending toward below the top of the cervical groove. In O. bredai , the marginal carina on pleonal pleura is wider than that on O. esocina . The pectinate claw of O. bredai possesses denticles which are perpendicular to the longitudinal axis of the fingers, unlike those in O. klintebjergensis which are angled distally.

Range and occurrence. Oncopareia bredai was first described from what is now the Kunrade Formation, a lateral equivalent of part of the Maastricht Formation, of late Maastrichtian age, and exposed in the eastern part of southern Limburg, the Netherlands. Subsequent records are from the Vijlen Member, interval 6 (Gulpen Formation), of late early Maastrichtian age, as exposed at the former SA Ciments Portland Liegeois (CPL) quarry near Haccourt and the adjacent Ciments Belge Réunie (CBR) quarry, province of Liège, north-east Belgium. Czech occurrences, referred to Hoploparia biserialis by Fritsch, in Fritsch and Kafka (1887) originate from the uppermost part of the Jizera Formation (upper Turonian) at Vinary near Vysoké Mýto in the Bohemian Cretaceous Basin, Czech Republic. The stratigraphical level of N. coesfeldiensis was indicated as ‘Senonian’, both lower and upper, of north-central and northwest Germany by Mertin (1941); in current terminology, this would correspond to the lower and upper Campanian.

Figures

FIGURE 1. Morphology of cephalothorax and a general habitus of studied taxa. A, Cephalothorax of nephropid lobsters with selected morphological characters (modified from Tshudy and Sorhannus, 2003). B, Habitus of thaumastocheliform lobsters, exemplified by Thaumastocheles zaleucus (from Calman, 1911). C, Habitus of ctenochelid ghost shrimps, exemplified by Ctenocheles balssi (from Kishinouye, 1926). D, Groove pattern of Oncopareia bredai Bosquet, 1854 (redrawn from Tshudy and Sorhannus, 2000b, fig. 2.2). E, Groove pattern of Hoploparia beyrichi Schlüter, 1862 (redrawn from Tshudy and Sorhannus, 2000b, fig. 2.3).

FIGURE 2. Oncopareia bredai (sensu Bosquet, 1854), as a composite species. A, Cephalothorax of O. bredai (from Bosquet, 1854, pl. 10, fig. 5). B, Pleon of Hoploparia beyrichi Schlüter, 1862 (from Bosquet, 1854, pl. 10, fig. 6). C, Claw of H. beyrichi (from Bosquet, 1854, pl. 10, fig. 71b). D, Composite drawing from Mertin (1941, text-fig. 9e). E, Pleonal segments 1 and 2 of O. bredai (as Homarus bosqueti Pelseneer, 1886; from Pelseneer, 1886, fig. 4).

FIGURE 3. Oncopareia bredai Bosquet, 1854; upper Maastrichtian (Kunrade Formation) of the Kunrade area, southern Limburg, the Netherlands. A, Cephalothorax in lateral view (from Bosquet, 1854, pl. 10, fig. 5). B, Anterior portion of cephalothorax in lateral view (from Bosquet, 1854, pl. 10, fig. 8). C, Cephalothorax with pleon and tailfan in lateral view (from Bosquet, 1854, pl. 10, fig. 61). D, Cephalothorax in lateral view (IRScNB 90-32c). E, Cephalothorax in lateral view (IRScNB 90-17). F, Anterior portion of cephalothorax in lateral view (IRScNB 90-15b). G, Incomplete individual in lateral view (IRScNB 90-33f, designated lectotype herein). H, I, Pleon in lateral (H) and dorsal (I) views (IRScNB 90-35c). J, Pleonal segments 1–3 in lateral view (IRScNB 90-23b). K, Right (crusher) and left (pectinate) claws of the same individual (IRScNB 90-28). L, Fingers of pectinate claw in lateral view (IRScNB 90-3b). M, Crusher claw in lateral view (IRScNB 90-32a). N, Pectinate claw in lateral view (IRScNB 90-27b). Specimens in D–F and H–N were whitened with ammonium chloride prior to photography.

FIGURE 4. Oncopareia bredai Bosquet, 1854; cephalothorax with associated pleonal segments 1 and 2 and both claws (MNHN R03408) from the upper Maastrichtian (Kunrade Formation) at Kunrade, the Netherlands. A, Cephalothorax in left lateral view. B, Cephalothorax in dorsal view. C, Cephalothorax in right lateral view. D, Both claws. E, Crusher claw in lateral view.

FIGURE 5. Oncopareia bredai Bosquet, 1854, cephalothoraxes, partial pleons and isolated claws; with exception of H (which is from the upper Campanian Zeven Wegen Member), all are from the Vijlen Member, interval 6 (upper lower Maastrichtian) at the CPL/CBR quarry complex near Haccourt and Lixhe, north-east Belgium. A, Cephalothorax in right lateral view (NHMM JJ 15836). B, C, Cephalothorax in right lateral view, and counterpart (NHMM JJ 6741). D, E, Cephalothorax in right lateral view, and counterpart (NHMM JJ 14618). F, Fragmentary cephalothorax and pleon in right lateral view (NHMM JJ 6764). G, Isolated pleura (NHMM JJ 6741). I, Fragment of cephalothorax and pectinate claw (NHMM JJ 14618). J, pectinate claw in lateral view (NHMM JJ 15534). K, crusher claw in lateral view (NHMM JJ 7194). L, fragmentary pectinate claw in lateral view (NHMM JJ 6839).

FIGURE 6. Oncopareia bredai Bosquet, 1854; isolated cephalothoraxes from the upper Turonian (uppermost part of the Jizera Formation) at Vinary near Vysoké Mýto, Czech Republic. A, Isolated cephalothorax in lateral view (NM O3470). B, Digital copy of Fritsch and Kafka (1887, pl. 5, fig. 2). C, Isolated cephalothorax in lateral view (NM O6861). Both specimens were whitened with ammonium chloride prior to photography.

FIGURE 7. Nymphaeops coesfeldiensis Schlüter, 1862 (= Oncopareia bredai Bosquet, 1854, as here understood). A, Incomplete cephalothorax with associated pleon in lateral view (from Schlüter, 1862, pl. 13, fig. 3). B, Cephalothorax and pleon in dorsal view (from Schlüter, 1862, pl. 13, fig. 6). C, Reconstruction of cephalothorax in dorsal view (from Schlüter, 1879, pl. 15, fig. 2). D, Near-complete individual in lateral view (from Schlüter, 1879, pl. 15, fig. 1). E, Cephalothorax and pleon in lateral view (from Mertin, 1941, text-fig. 9a). F, Pectinate claw (from Mertin, 1941, text-fig. 10k). G, Crusher claw (from Mertin, 1941, text-fig. 10b). H, Crusher claw (from Mertin, 1941, text-fig. 10d). Nymphaeops coesfeldiensis Schlüter, 1862 (= Oncopareia bredai Bosquet, 1854, as here understood) from the Vaals Formation (lower Campanian) at the former CPL SA quarry, Haccourt, north-east Belgium (I–K). I, Near-complete cephalothorax with associated appendages in lateral view (NHMM JJ 9974). J, Crusher claw (NHMM HB 393). K, Fingers of pectinate claw (NHMM HB 756). Figure parts A–H not to scale.