(Fig. 7 D-F)
Glyphea veltheimii Münster, 1839: 22, pl. 10, fig. 1. — Fraas 1855: 94. — Glaessner 1929: 159. — Woods 1930: 80, pl. 22, figs 5-6. — Van Straelen 1936: 9. — Förster 1966: 124, fig. 23, pl. 17, figs 2, 4. — Taylor 1979: 34. — Feldmann & Titus 2006: 64. — Karasawa et al. 2013: table 1.
Eryma veltheimii – Bronn 1849: 579. — Oppel 1861: 358; 1862: 36, pl. 7, fig. 5. — Beurlen 1928: 156. — Schweigert et al. 2000: 7, fig. 1c, pl. 4, figs 1-3. — Garassino & Schweigert 2006: 8, fig. 8, pl. 1, fig. 2, pl. 11, figs 3-4. — Feldmann & Titus 2006: 64. — Fürsich et al. 2007a: table 2. — Odin et al. 2019: 652.
Clytia veltheimii – Beurlen 1928: 170.
Galicia veltheimii – Schweitzer et al. 2010: 25. — Charbonnier & Garassino 2012: 864. — Audo et al. 2014: 462. — Schweigert 2015: fig. 544.
TYPE MATERIAL. — Holotype BSPG AS VII 186.
TYPE LOCALITY. — Kapfelberg quarry near Kelheim, Bavaria, Germany.
TYPE AGE. — Kimmeridgian.
DESCRIPTION
Carapace
Short, spiny rostrum; fusiform intercalated plate; elongated cephalic region, representing almost half of the length of the carapace; wide, deep cervical groove, strongly inclined, joined to dorsal margin; deep gastro-orbital groove, elongated and oblique; postcervical groove not identified; wide, deep branchiocardiac groove, strongly inclined.
Somites with subtriangular pleurites; telson with a median line and two longitudinal crests along its lateral margins, crests ended by a small spine; uropods as long as telson, with a small spine on the external margin; uropodal endopods with a longitudinal carina; uropodal exopods with a diaeresis, and a longitudinal carina.
Chelate P 1; P 1 propodus subrectangular, strongly elongated, dorso-ventrally compressed; presence of a strong spine at the inner distal extremity of the propodus, above the dactylus; slender fingers, progressively narrowing to their distal extremity, slightly curved inward; occlusal margin without teeth; P 1 carpus short, subtriangular; thin P 2- P 5.
Carapace covered by small tubercles; intercalated plate covered by small tubercles; oblique orbital row of tubercles in cephalic region; presence of a strong antennal spine; smooth P 1 propodus and fingers.
DISCUSSION
This species is known by some specimens, mainly from the Kimmeridgian plattenkalks of Wattendorf (Audo et al. 2014). Because of the compression of the holotype, it is difficult to clearly identify the grooves of the carapace. Indeed, only the cervical, gastro-orbital and branchiocardiac grooves are wellmarked. However, the assignation to Eryma is supported by the gastro-orbital groove wich is not divided in two branches and by the shape of the P 1 chelae (elongated subrectangular propodus; elongated fingers, curved inward and narrowing to their distal extremity).
The difficulty to identify the carapace grooves does not allow the comparison with that of other species. The presence of a spine at the distal extremity of the inner margin of the propodus (Fig. 7D) is characteristic of this species. Moreover, the lack of ornamentation on the P 1 chelae is a characteristic only found in E. veltheimii and E. westphali. A spiny rostrum like in E. veltheimii is not found in E. mandelslohi, E. modestiforme, and E. ventrosum. Garassino & Schweigert (2006) pointed out the absence of spines on the margin of the telson, contrary to E. modestiforme. However, the careful examination of the type specimen shows that two marginal spines are present (Fig. 7F). Finally, the carapace of E. veltheimii is only covered by tubercles, contrary to E. georgeii, E. mandelslohi, E. quadriverrucatum, E. ventrosum, and E. westphali.
Eryma westphali Schweigert, Dietl & Röper, 2000 (Fig. 7 G-J)
Eryma westphali Schweigert, Dietl & Röper, 2000: 6, pl. 3, figs 1-3. — Dietl & Schweigert 2001: 53, 110, fig. 85. — Garassino & Schweigert 2006: 8. — Feldmann & Titus 2006: 64.
Galicia westphali – Schweitzer et al. 2010: 25. — Hyžný et al. 2015: 375.
TYPE MATERIAL. — Holotype SMNS 24227.
TYPE LOCALITY. — Nusplingen, Baden-Württemberg, Germany.
TYPE AGE. — Kimmeridgian.
DESCRIPTION
Carapace
Short, spiny rostrum; fusiform intercalated plate; deep, wide cervical groove, joined to dorsal margin and to antennal groove; deep antennal groove; short, sub-horizontal gastroorbital groove; postcervical groove slightly curved forward, not joined to dorsal margin, joined to branchiocardiac groove at carapace mid-height, with a short ventral extension; branchiocardiac groove dorsally deep, narrowing in its ventral part, not joined to dorsal margin, joined to hepatic groove; hepatic groove concavo-convex.
Somites with subtriangular pleurites, with a bulge on their basis; wide rounded uropods; uropodal endopods with a longitudinal carina; uropodal exopods with a small spine on external margin.
Chelate P 1; P 1 propodus subrectangular, narrow and strongly elongated; P 1 fingers shorter than propodus, slender, almost
straight or slightly curved inward, with a terminal hook; occlusal margin without teeth; P 1 carpus short, subtriangular; thin P 2- P 5; P 2- P 3 chelate.
Carapace with a dense, heterogeneous ornamentation; gastric and cardiac regions covered by tubercles; antennal, hepatic, and branchial regions covered by small rounded depressions; presence of an orbital spine; tergites and pleurites densely covered by small rounded depressions; presence of a strong antennal spine; smooth P 1- P 5.
DISCUSSION
This species is only known by three specimens. It is assigned to Eryma because of its typical carapace groove pattern (short gastro-orbital groove, postcervical and branchiocardiac grooves joined at carapace mid-height, hepatic groove concavo-convex) and the shape of the P 1 chelae (elongated propodus, subrectangular, with slender fingers progressively narrowing to their distal extremity).
Among the erymid lobsters, Eryma westphali is the only species with heterochelous P 1 chelae on a same specimen. Left propodus is more elongated and narrower than the right one, and the left fingers are straight and short while the right ones are curved and slightly longer (Fig. 7I, J). The strongly elongated and narrow P 1 propodus with relatively short fingers of E. westphali is characteristic of the species. Only Eryma veltheimii (Münster, 1839) have P 1 with a very long propodus, but it is clearly wider.
In E. westphali the branchial region is dorsally shorter than in E. georgeii, E. jungostrix, E. lerasi, E. modestiforme, and E. ventrosum. Contrary to E. lerasi and E. quadriverrucatum, E. westphali exhibits a ventral extension of the postcervical groove. Moreover, the dichotomy of the ornamentation of the carapace between the gastric and cardiac regions, covered by tubercles, and the remaining regions of E. westphali, covered by small depressions, is unique among the genus.
Eryma modestiforme (Schlotheim, 1822) (Figs 8; 9)
Macrourites modestiformis Schlotheim, 1822: 29, pl. 2, fig. 3.
Astacus leptodactylus Germar, 1827: 100, pl. 1, fig. 4.
Glyphea crassula Münster, 1839: 17, pl. 8, fig. 5.
Glyphea elongata Münster, 1839: 18, pl. 8, fig. 8, 11-12.
Glyphea laevigata Münster, 1839: 20, pl. 9, figs 5-7.
Palaeastacus poeschli Schweigert & Röper, 2001: 5, figs 3-4. — Garassino & Schweigert 2006: 11. — Schweitzer et al. 2010: 25. n. syn.
Glyphea modestiformis – Münster 1839: 17, pl. 8, fig. 9, pl. 9, figs 1-3. Eryma modestiformis – Bronn 1849: 579. — Oppel 1861: 358; 1862: 33, pl. 6, figs 5-8. — Van Straelen 1925: 271. — Beurlen 1928: 156, 157, 163, 164. — Glaessner 1929: 156. — Roger 1946: 42. — Kuhn 1961: 22. — Secrétan 1964: 69. — Förster 1965: 138, fig. 1, pl. 2, fig. 5. — Förster 1966: 118, fig. 21, pl. 16, figs 7-8 (non 9), pl. 17, fig. 1. — Schweigert & Garassino 2003: 178. — Fürsich et al. 2007b: 57. — De Grave et al. 2009: fig. 1B. — Feldmann & Schweitzer 2017: fig. 1c. — Odin et al. 2019: 652, fig. 6, ap. 1.
Astacus modestiformis – Quenstedt 1852: 268; 1867: 320; 1885: 409. — Fraas 1855: 94.
Eryma leptodactylina – Oppel 1861: 358; 1862: 35, pl. 7, figs 1-4. — Zittel 1885: fig. 873. — Secrétan 1964: 69. — Garassino & Schweigert 2006: 8.
Eryma leptodactylus – Van Straelen 1925: 273.
Clytia leptodactylina – Beurlen 1928: 170. — Vialle 1948: 65.
Clytia leptodactylus – Glaessner 1929: 116. — Kuhn 1961: 22.
Eryma elongata – Frickhinger 1994: 118, figs 201-202.
Eryma modestiforme – Schweigert et al. 2000: 4, fig. 1a, pl. 1, figs 1-5. — Dietl & Schweigert 2001: 53, 110, fig.86. — Schweigert & Röper 2001: 2, 7. — Garassino & Schweigert 2006: 6, fig. 8, pl. 1, fig. 1, pl. 11, figs 1-2. — Schweitzer et al. 2010: 24. — Charbonnier & Garassino 2012: 864, fig. 3B-C. — Karasawa et al. 2013: table 1, fig. 9 A. — Hyžný et al. 2015: 375, fig. 3 A. — Schweigert 2015: fig. 541. — Devillez et al. 2016: 518, 524. — Devillez & Charbonnier 2017: table 1, fig. 2a.
TYPE MATERIAL. — Holotype MFN 2236 P 1383/2 MB. A.0252.
TYPE LOCALITY. — Eichstätt, Bavaria, Germany.
TYPE AGE. — Kimmeridgian-Tithonian.
DESCRIPTION
Carapace
Sub-cylindrical carapace; slightly elongated, spineless rostrum; fusiform intercalated plate; orbital notch widely curved; narrow post-orbital area; elongated cephalic region, representing almost half of the length of the carapace; deep and wide cervical groove, almost straight dorsally, strongly inflected at carapace mid-height, joined to dorsal margin and to antennal groove; deep antennal groove; short, narrow gastro-orbital groove, oblique, originating as median inflexion of cervical groove; postcervical and branchiocardiac grooves subparallel, slightly curved, slightly inclined, not joined to dorsal margin; shallow postcervical groove, joined to branchiocardiac groove, with a short ventral extension; narrow, shallow branchiocardiac groove, joined to the posterior extremity of hepatic groove; hepatic groove concavo-convex, joined to cervical groove; inferior groove joined to hepatic groove.
Somites with wide, short subtriangular pleurites, with a bulge on their basis; s2 pleurites wider than others; telson with a longitudinal median groove and two longitudinal crests along
Solnhofen: specimen BSPG AS VII 198 (C), AS VII 194 (D), AS VII 197 (E); G, holotype BSPG AS VII 193 of Glyphea crassula Münster, 1839 from Solnhofen; H, holotype SMNS 64520 of Palaeastacus poeschli Schweigert & Röper, 2001 from Mülheim. Abbreviations: a, branchiocardiac groove; b, antennal groove; b 1 , hepatic groove; c, postcervical groove; d, gastro-orbital groove; e, eye; e 1 e , cervical groove; i, inferior groove; ip, intercalated plate; PoA, postorbital area; s1-s6, pleonal somites. Photographs and line drawing: J. Devillez. Scale bars: 1 cm.
lateral margins; uropods as long as telson; uropodal endopods with a longitudinal carina; uropodal exopods with a diaeresis, and a longitudinal carina.
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; short, triangular scaphocerite; wide epistome, with a linear contact with the carapace; wide, subrectangular mandibles (Fig. 9 A-C).
Elongated Mxp3; chelate P 1; P 1 propodus subrectangular; slender, straight P 1 fingers, longer than propodus, equal in length, sometimes with a small distal hook; occlusal margin without teeth; P 1 carpus short, subtriangular; elongated P 1 merus; P 2- P 3 chelate.
Carapace with a homogeneous ornamentation; carapace densely covered by small tubercles and depressions; gastric region with an orbital spine; antennal region with an antennal spine; pleonal tergites and pleurites densely covered by small rounded depressions; telson covered by small depressions; P 1 covered by small depressions; smooth P 2- P 5.
DISCUSSION
Eryma modestiforme is known by numerous specimens, almost complete. This is the most common erymoid of the lithographic limestones of southern Germany.
The type materials of Astacus leptodactylus Germar, 1827 (Fig. 8C), Glyphea crassula Münster, 1839 (Fig. 8H), Glyphea elongata Münster, 1839 (Fig. 8D), Glyphea laevigata Münster, 1839 (Fig. 8 E-G) and Palaeastacus poeschli Schweigert & Röper, 2001 (Fig. 8I), from the same formations than E. modestiforme, share the same characteristics: a subrectangular propodus bearing longer fingers, these fingers are slender and straight, without teeth on their occlusal margins, the carapace ornamentation is dense, made of small tubercles and depressions. These characteristics support the integration of A. leptodactylus, G. crassula, G. elongata, G. laevigata and P. poeschli into the synonymy of E. modestiforme.
The ventral extension of the cervical groove seen on E. modestiforme is absent in E. lerasi and E. quadriverrucatum. The morphology of the P 1 is also distinct from some other representatives of the genus. Indeed, the propodus is rectangular in E. modestiforme while it is trapezoidal in E. georgeii and E. mandelslohi. It is also relatively narrower than the propodus of E. georgeii, E. mandelslohi and E. quadriverrucatum. The almost straight P 1 fingers of E. modestiforme contrast with the curved ones of E. georgeii, E. lerasi, E. veltheimii, E. ventrosum, and E. westphali. Finally, the fine ornamentation of E. modestiforme, made of tubercles and depressions, is clearly distinct from the ornamentation of E. georgeii, E. jungostrix, E. mandelslohi, E. veltheimii, and E. westphali.
Eryma westphali Schweigert, Dietl & Röper, 2000 (Fig. 7 G-J)
Eryma westphali Schweigert, Dietl & Röper, 2000: 6, pl. 3, figs 1-3. — Dietl & Schweigert 2001: 53, 110, fig. 85. — Garassino & Schweigert 2006: 8. — Feldmann & Titus 2006: 64.
Galicia westphali – Schweitzer et al. 2010: 25. — Hyžný et al. 2015: 375.
TYPE MATERIAL. — Holotype SMNS 24227.
TYPE LOCALITY. — Nusplingen, Baden-Württemberg, Germany.
TYPE AGE. — Kimmeridgian.
DESCRIPTION
Carapace
Short, spiny rostrum; fusiform intercalated plate; deep, wide cervical groove, joined to dorsal margin and to antennal groove; deep antennal groove; short, sub-horizontal gastroorbital groove; postcervical groove slightly curved forward, not joined to dorsal margin, joined to branchiocardiac groove at carapace mid-height, with a short ventral extension; branchiocardiac groove dorsally deep, narrowing in its ventral part, not joined to dorsal margin, joined to hepatic groove; hepatic groove concavo-convex.
Somites with subtriangular pleurites, with a bulge on their basis; wide rounded uropods; uropodal endopods with a longitudinal carina; uropodal exopods with a small spine on external margin.
Chelate P 1; P 1 propodus subrectangular, narrow and strongly elongated; P 1 fingers shorter than propodus, slender, almost
straight or slightly curved inward, with a terminal hook; occlusal margin without teeth; P 1 carpus short, subtriangular; thin P 2- P 5; P 2- P 3 chelate.
Carapace with a dense, heterogeneous ornamentation; gastric and cardiac regions covered by tubercles; antennal, hepatic, and branchial regions covered by small rounded depressions; presence of an orbital spine; tergites and pleurites densely covered by small rounded depressions; presence of a strong antennal spine; smooth P 1- P 5.
DISCUSSION
This species is only known by three specimens. It is assigned to Eryma because of its typical carapace groove pattern (short gastro-orbital groove, postcervical and branchiocardiac grooves joined at carapace mid-height, hepatic groove concavo-convex) and the shape of the P 1 chelae (elongated propodus, subrectangular, with slender fingers progressively narrowing to their distal extremity).
Among the erymid lobsters, Eryma westphali is the only species with heterochelous P 1 chelae on a same specimen. Left propodus is more elongated and narrower than the right one, and the left fingers are straight and short while the right ones are curved and slightly longer (Fig. 7I, J). The strongly elongated and narrow P 1 propodus with relatively short fingers of E. westphali is characteristic of the species. Only Eryma veltheimii (Münster, 1839) have P 1 with a very long propodus, but it is clearly wider.
In E. westphali the branchial region is dorsally shorter than in E. georgeii, E. jungostrix, E. lerasi, E. modestiforme, and E. ventrosum. Contrary to E. lerasi and E. quadriverrucatum, E. westphali exhibits a ventral extension of the postcervical groove. Moreover, the dichotomy of the ornamentation of the carapace between the gastric and cardiac regions, covered by tubercles, and the remaining regions of E. westphali, covered by small depressions, is unique among the genus.
Eryma modestiforme (Schlotheim, 1822) (Figs 8; 9)
Macrourites modestiformis Schlotheim, 1822: 29, pl. 2, fig. 3.
Astacus leptodactylus Germar, 1827: 100, pl. 1, fig. 4.
Glyphea crassula Münster, 1839: 17, pl. 8, fig. 5.
Glyphea elongata Münster, 1839: 18, pl. 8, fig. 8, 11-12.
Glyphea laevigata Münster, 1839: 20, pl. 9, figs 5-7.
Palaeastacus poeschli Schweigert & Röper, 2001: 5, figs 3-4. — Garassino & Schweigert 2006: 11. — Schweitzer et al. 2010: 25. n. syn.
Glyphea modestiformis – Münster 1839: 17, pl. 8, fig. 9, pl. 9, figs 1-3. Eryma modestiformis – Bronn 1849: 579. — Oppel 1861: 358; 1862: 33, pl. 6, figs 5-8. — Van Straelen 1925: 271. — Beurlen 1928: 156, 157, 163, 164. — Glaessner 1929: 156. — Roger 1946: 42. — Kuhn 1961: 22. — Secrétan 1964: 69. — Förster 1965: 138, fig. 1, pl. 2, fig. 5. — Förster 1966: 118, fig. 21, pl. 16, figs 7-8 (non 9), pl. 17, fig. 1. — Schweigert & Garassino 2003: 178. — Fürsich et al. 2007b: 57. — De Grave et al. 2009: fig. 1B. — Feldmann & Schweitzer 2017: fig. 1c. — Odin et al. 2019: 652, fig. 6, ap. 1.
Astacus modestiformis – Quenstedt 1852: 268; 1867: 320; 1885: 409. — Fraas 1855: 94.
Eryma leptodactylina – Oppel 1861: 358; 1862: 35, pl. 7, figs 1-4. — Zittel 1885: fig. 873. — Secrétan 1964: 69. — Garassino & Schweigert 2006: 8.
Eryma leptodactylus – Van Straelen 1925: 273.
Clytia leptodactylina – Beurlen 1928: 170. — Vialle 1948: 65.
Clytia leptodactylus – Glaessner 1929: 116. — Kuhn 1961: 22.
Eryma elongata – Frickhinger 1994: 118, figs 201-202.
Eryma modestiforme – Schweigert et al. 2000: 4, fig. 1a, pl. 1, figs 1-5. — Dietl & Schweigert 2001: 53, 110, fig.86. — Schweigert & Röper 2001: 2, 7. — Garassino & Schweigert 2006: 6, fig. 8, pl. 1, fig. 1, pl. 11, figs 1-2. — Schweitzer et al. 2010: 24. — Charbonnier & Garassino 2012: 864, fig. 3B-C. — Karasawa et al. 2013: table 1, fig. 9 A. — Hyžný et al. 2015: 375, fig. 3 A. — Schweigert 2015: fig. 541. — Devillez et al. 2016: 518, 524. — Devillez & Charbonnier 2017: table 1, fig. 2a.
TYPE MATERIAL. — Holotype MFN 2236 P 1383/2 MB. A.0252.
TYPE LOCALITY. — Eichstätt, Bavaria, Germany.
TYPE AGE. — Kimmeridgian-Tithonian.
DESCRIPTION
Carapace
Sub-cylindrical carapace; slightly elongated, spineless rostrum; fusiform intercalated plate; orbital notch widely curved; narrow post-orbital area; elongated cephalic region, representing almost half of the length of the carapace; deep and wide cervical groove, almost straight dorsally, strongly inflected at carapace mid-height, joined to dorsal margin and to antennal groove; deep antennal groove; short, narrow gastro-orbital groove, oblique, originating as median inflexion of cervical groove; postcervical and branchiocardiac grooves subparallel, slightly curved, slightly inclined, not joined to dorsal margin; shallow postcervical groove, joined to branchiocardiac groove, with a short ventral extension; narrow, shallow branchiocardiac groove, joined to the posterior extremity of hepatic groove; hepatic groove concavo-convex, joined to cervical groove; inferior groove joined to hepatic groove.
Somites with wide, short subtriangular pleurites, with a bulge on their basis; s2 pleurites wider than others; telson with a longitudinal median groove and two longitudinal crests along
Solnhofen: specimen BSPG AS VII 198 (C), AS VII 194 (D), AS VII 197 (E); G, holotype BSPG AS VII 193 of Glyphea crassula Münster, 1839 from Solnhofen; H, holotype SMNS 64520 of Palaeastacus poeschli Schweigert & Röper, 2001 from Mülheim. Abbreviations: a, branchiocardiac groove; b, antennal groove; b 1 , hepatic groove; c, postcervical groove; d, gastro-orbital groove; e, eye; e 1 e , cervical groove; i, inferior groove; ip, intercalated plate; PoA, postorbital area; s1-s6, pleonal somites. Photographs and line drawing: J. Devillez. Scale bars: 1 cm.
lateral margins; uropods as long as telson; uropodal endopods with a longitudinal carina; uropodal exopods with a diaeresis, and a longitudinal carina.
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; short, triangular scaphocerite; wide epistome, with a linear contact with the carapace; wide, subrectangular mandibles (Fig. 9 A-C).
Elongated Mxp3; chelate P 1; P 1 propodus subrectangular; slender, straight P 1 fingers, longer than propodus, equal in length, sometimes with a small distal hook; occlusal margin without teeth; P 1 carpus short, subtriangular; elongated P 1 merus; P 2- P 3 chelate.
Carapace with a homogeneous ornamentation; carapace densely covered by small tubercles and depressions; gastric region with an orbital spine; antennal region with an antennal spine; pleonal tergites and pleurites densely covered by small rounded depressions; telson covered by small depressions; P 1 covered by small depressions; smooth P 2- P 5.
DISCUSSION
Eryma modestiforme is known by numerous specimens, almost complete. This is the most common erymoid of the lithographic limestones of southern Germany.
The type materials of Astacus leptodactylus Germar, 1827 (Fig. 8C), Glyphea crassula Münster, 1839 (Fig. 8H), Glyphea elongata Münster, 1839 (Fig. 8D), Glyphea laevigata Münster, 1839 (Fig. 8 E-G) and Palaeastacus poeschli Schweigert & Röper, 2001 (Fig. 8I), from the same formations than E. modestiforme, share the same characteristics: a subrectangular propodus bearing longer fingers, these fingers are slender and straight, without teeth on their occlusal margins, the carapace ornamentation is dense, made of small tubercles and depressions. These characteristics support the integration of A. leptodactylus, G. crassula, G. elongata, G. laevigata and P. poeschli into the synonymy of E. modestiforme.
The ventral extension of the cervical groove seen on E. modestiforme is absent in E. lerasi and E. quadriverrucatum. The morphology of the P 1 is also distinct from some other representatives of the genus. Indeed, the propodus is rectangular in E. modestiforme while it is trapezoidal in E. georgeii and E. mandelslohi. It is also relatively narrower than the propodus of E. georgeii, E. mandelslohi and E. quadriverrucatum. The almost straight P 1 fingers of E. modestiforme contrast with the curved ones of E. georgeii, E. lerasi, E. veltheimii, E. ventrosum, and E. westphali. Finally, the fine ornamentation of E. modestiforme, made of tubercles and depressions, is clearly distinct from the ornamentation of E. georgeii, E. jungostrix, E. mandelslohi, E. veltheimii, and E. westphali.
Eryma modestiforme (Schlotheim, 1822) (Figs 8; 9)
Macrourites modestiformis Schlotheim, 1822: 29, pl. 2, fig. 3.
Astacus leptodactylus Germar, 1827: 100, pl. 1, fig. 4.
Glyphea crassula Münster, 1839: 17, pl. 8, fig. 5.
Glyphea elongata Münster, 1839: 18, pl. 8, fig. 8, 11-12.
Glyphea laevigata Münster, 1839: 20, pl. 9, figs 5-7.
Palaeastacus poeschli Schweigert & Röper, 2001: 5, figs 3-4. — Garassino & Schweigert 2006: 11. — Schweitzer et al. 2010: 25. n. syn.
Glyphea modestiformis – Münster 1839: 17, pl. 8, fig. 9, pl. 9, figs 1-3. Eryma modestiformis – Bronn 1849: 579. — Oppel 1861: 358; 1862: 33, pl. 6, figs 5-8. — Van Straelen 1925: 271. — Beurlen 1928: 156, 157, 163, 164. — Glaessner 1929: 156. — Roger 1946: 42. — Kuhn 1961: 22. — Secrétan 1964: 69. — Förster 1965: 138, fig. 1, pl. 2, fig. 5. — Förster 1966: 118, fig. 21, pl. 16, figs 7-8 (non 9), pl. 17, fig. 1. — Schweigert & Garassino 2003: 178. — Fürsich et al. 2007b: 57. — De Grave et al. 2009: fig. 1B. — Feldmann & Schweitzer 2017: fig. 1c. — Odin et al. 2019: 652, fig. 6, ap. 1.
Astacus modestiformis – Quenstedt 1852: 268; 1867: 320; 1885: 409. — Fraas 1855: 94.
Eryma leptodactylina – Oppel 1861: 358; 1862: 35, pl. 7, figs 1-4. — Zittel 1885: fig. 873. — Secrétan 1964: 69. — Garassino & Schweigert 2006: 8.
Eryma leptodactylus – Van Straelen 1925: 273.
Clytia leptodactylina – Beurlen 1928: 170. — Vialle 1948: 65.
Clytia leptodactylus – Glaessner 1929: 116. — Kuhn 1961: 22.
Eryma elongata – Frickhinger 1994: 118, figs 201-202.
Eryma modestiforme – Schweigert et al. 2000: 4, fig. 1a, pl. 1, figs 1-5. — Dietl & Schweigert 2001: 53, 110, fig.86. — Schweigert & Röper 2001: 2, 7. — Garassino & Schweigert 2006: 6, fig. 8, pl. 1, fig. 1, pl. 11, figs 1-2. — Schweitzer et al. 2010: 24. — Charbonnier & Garassino 2012: 864, fig. 3B-C. — Karasawa et al. 2013: table 1, fig. 9 A. — Hyžný et al. 2015: 375, fig. 3 A. — Schweigert 2015: fig. 541. — Devillez et al. 2016: 518, 524. — Devillez & Charbonnier 2017: table 1, fig. 2a.
TYPE MATERIAL. — Holotype MFN 2236 P 1383/2 MB. A.0252.
TYPE LOCALITY. — Eichstätt, Bavaria, Germany.
TYPE AGE. — Kimmeridgian-Tithonian.
DESCRIPTION
Carapace
Sub-cylindrical carapace; slightly elongated, spineless rostrum; fusiform intercalated plate; orbital notch widely curved; narrow post-orbital area; elongated cephalic region, representing almost half of the length of the carapace; deep and wide cervical groove, almost straight dorsally, strongly inflected at carapace mid-height, joined to dorsal margin and to antennal groove; deep antennal groove; short, narrow gastro-orbital groove, oblique, originating as median inflexion of cervical groove; postcervical and branchiocardiac grooves subparallel, slightly curved, slightly inclined, not joined to dorsal margin; shallow postcervical groove, joined to branchiocardiac groove, with a short ventral extension; narrow, shallow branchiocardiac groove, joined to the posterior extremity of hepatic groove; hepatic groove concavo-convex, joined to cervical groove; inferior groove joined to hepatic groove.
Somites with wide, short subtriangular pleurites, with a bulge on their basis; s2 pleurites wider than others; telson with a longitudinal median groove and two longitudinal crests along
Solnhofen: specimen BSPG AS VII 198 (C), AS VII 194 (D), AS VII 197 (E); G, holotype BSPG AS VII 193 of Glyphea crassula Münster, 1839 from Solnhofen; H, holotype SMNS 64520 of Palaeastacus poeschli Schweigert & Röper, 2001 from Mülheim. Abbreviations: a, branchiocardiac groove; b, antennal groove; b 1 , hepatic groove; c, postcervical groove; d, gastro-orbital groove; e, eye; e 1 e , cervical groove; i, inferior groove; ip, intercalated plate; PoA, postorbital area; s1-s6, pleonal somites. Photographs and line drawing: J. Devillez. Scale bars: 1 cm.
lateral margins; uropods as long as telson; uropodal endopods with a longitudinal carina; uropodal exopods with a diaeresis, and a longitudinal carina.
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; short, triangular scaphocerite; wide epistome, with a linear contact with the carapace; wide, subrectangular mandibles (Fig. 9 A-C).
Elongated Mxp3; chelate P 1; P 1 propodus subrectangular; slender, straight P 1 fingers, longer than propodus, equal in length, sometimes with a small distal hook; occlusal margin without teeth; P 1 carpus short, subtriangular; elongated P 1 merus; P 2- P 3 chelate.
Carapace with a homogeneous ornamentation; carapace densely covered by small tubercles and depressions; gastric region with an orbital spine; antennal region with an antennal spine; pleonal tergites and pleurites densely covered by small rounded depressions; telson covered by small depressions; P 1 covered by small depressions; smooth P 2- P 5.
DISCUSSION
Eryma modestiforme is known by numerous specimens, almost complete. This is the most common erymoid of the lithographic limestones of southern Germany.
The type materials of Astacus leptodactylus Germar, 1827 (Fig. 8C), Glyphea crassula Münster, 1839 (Fig. 8H), Glyphea elongata Münster, 1839 (Fig. 8D), Glyphea laevigata Münster, 1839 (Fig. 8 E-G) and Palaeastacus poeschli Schweigert & Röper, 2001 (Fig. 8I), from the same formations than E. modestiforme, share the same characteristics: a subrectangular propodus bearing longer fingers, these fingers are slender and straight, without teeth on their occlusal margins, the carapace ornamentation is dense, made of small tubercles and depressions. These characteristics support the integration of A. leptodactylus, G. crassula, G. elongata, G. laevigata and P. poeschli into the synonymy of E. modestiforme.
The ventral extension of the cervical groove seen on E. modestiforme is absent in E. lerasi and E. quadriverrucatum. The morphology of the P 1 is also distinct from some other representatives of the genus. Indeed, the propodus is rectangular in E. modestiforme while it is trapezoidal in E. georgeii and E. mandelslohi. It is also relatively narrower than the propodus of E. georgeii, E. mandelslohi and E. quadriverrucatum. The almost straight P 1 fingers of E. modestiforme contrast with the curved ones of E. georgeii, E. lerasi, E. veltheimii, E. ventrosum, and E. westphali. Finally, the fine ornamentation of E. modestiforme, made of tubercles and depressions, is clearly distinct from the ornamentation of E. georgeii, E. jungostrix, E. mandelslohi, E. veltheimii, and E. westphali.