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. 2023 Mar 8;15(3):386.
doi: 10.3390/d15030386.

A Synoptic Review of the Cartilaginous Fishes (Chondrichthyes: Holocephali, Elasmobranchii) from the Upper Jurassic Konservat-Lagerstätten of Southern Germany: Taxonomy, Diversity, and Faunal Relationships

Eduardo Villalobos-Segura  1 , Sebastian Stumpf  1 , Julia Türtscher  1   2 , Patrick L Jambura  1   2 , Arnaud Begat  1   2 , Faviel A López-Romero  1 , Jan Fischer  3 , Jürgen Kriwet  1   2
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A Synoptic Review of the Cartilaginous Fishes (Chondrichthyes: Holocephali, Elasmobranchii) from the Upper Jurassic Konservat-Lagerstätten of Southern Germany: Taxonomy, Diversity, and Faunal Relationships

Eduardo Villalobos-Segura et al. Diversity (Basel). .

Abstract

The Late Jurassic-Early Cretaceous (164-100 Ma) represents one of the main transitional periods in life history. Recent studies unveiled a complex scenario in which abiotic and biotic factors and drivers on regional and global scales due to the fragmentation of Pangaea resulted in dramatic faunal and ecological turnovers in terrestrial and marine environments. However, chondrichthyan faunas from this interval have received surprisingly little recognition. The presence of numerous entire skeletons of chondrichthyans preserved in several localities in southern Germany, often referred to as Konservat-Lagerstätten (e.g., Nusplingen and the Solnhofen Archipelago), provides a unique opportunity of to study the taxonomic composition of these assemblages, their ecological distributions and adaptations, and evolutionary histories in detail. However, even after 160 years of study, the current knowledge of southern Germany's Late Jurassic chondrichthyan diversity remains incomplete. Over the last 20 years, the systematic study and bulk sampling of southern Germany's Late Jurassic deposits significantly increased the number of known fossil chondrichthyan genera from the region (32 in the present study). In the present work, the fossil record, and the taxonomic composition of Late Jurassic chondrichthyans from southern Germany are reviewed and compared with several contemporaneous assemblages from other sites in Europe. Our results suggest, inter alia, that the Late Jurassic chondrichthyans displayed extended distributions within Europe. However, it nevertheless also is evident that the taxonomy of Late Jurassic chondrichthyans is in urgent need of revision.

Keywords: Chondrichthyes; Germany; Kimmeridgian; Late Jurassic; Nusplingen; Solnhofen Archipelago; Tithonian; Wattendorf; biogeography; diversity.

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Conflict of interest statement

Conflicts of Interest: The authors declare no conflict of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, or in the decision to publish the results.

Figures

Figure 1
Figure 1. (A) Geographic and (B) palaeogeographic location maps (modified from [34,72]; biostratigraphic information based on [44,73]).
Figure 2
Figure 2. Dorsal fin spine of the myriacanthoid, †Chimaeropsis? franconicus (BT 5160.00) from the early Tithonian? (Upper Jurassic) of Streitberg near the castle ruin of Rabenstein.
Figure 3
Figure 3. †Chimaeropsis paradoxa von Zittel, 1887 [89].
(A) Fragmentary cranial remains (SNSB-BSPG AS I 1330) consisting mainly of dental parts from the early Tithonian of Solnhofen. This specimen was figured by von Meyer in Münster [50] (pl. 8, figure 1) and Riess [93] (pl. 3, figure 10). Abbreviations: md, mandibular tooth plates; sym, symphyseal tooth plates; pal, palatinal tooth plates; vo, vomerine tooth plate. (B) Holomorphic specimen (LF 2317) from the early Tithonian (Upper Jurassic) of Eichstätt. This specimen also was figured by Duffin et al. 98 (figure 9C). Photo provided and reproduced with permission by B. Lauer (Lauer Foundation for Paleontology).
Figure 4
Figure 4. †Ischyodus egertoni (Buckland, 1936 [101]).
(A) Holomorphic specimen (SNSB-BSPG 1954 I 366) from the early Tithonian Altmühltal Formation of Eichstätt, labelled as †I. quenstedti. This specimen was figured by Kriwet and Klug [19] (figure 675). (B) Right palatinal tooth plate (SMNS 96078/28) from the late Kimmeridgian Nusplingen Plattenkalk of the quarry of the Gesellschaft für Naturkunde Württemberg), labelled as †I. quenstedti. (C) Right palatinal tooth plate (SMNS 51566) from the upper Kimmeridgian of Nusplingen, labelled as †I. schuebleri. This specimen was collected by Fraas in 1855 and figured by Schweizer [66] (pl. 8, figures 5 and 6). (D) Isolated fin spine (SMNS 51565) from the upper Kimmeridgian of Nusplingen, labelled as †I. schuebleri.
Figure 5
Figure 5. Still undescribed, juvenile specimen of †Ischyodus egertoni (Buckland, 1936 [101]) (NMB without number) from the late Kimmeridgian Torleite Formation of Wattendorf in the northern part of the Franconian Alb.
Figure 6
Figure 6. Callorhynchidae indet.
(A) Cranial remains and basis of fin spine (SMNS 80144/22) from the late Kimmeridgian Nusplingen Plattenkalk of the quarry of the Gesellschaft für Naturkunde Württemberg, labelled as †I. schuebleri. (B) Cranial remains including the frontal clasper and complete dentition (SMNS 95823/4) from the late Kimmeridgian Nusplingen Plattenkalk of the quarry of the Gesellschaft für Naturkunde Württemberg, labelled as †Elasmodectes avitus. It was collected in 2011 and figured by Schweigert et al. [118] (pl. 4).
Figure 7
Figure 7. Specimens of †Elasmodectes avitus (von Meyer, 1862 [120]) from the Solnhofen Archipelago.
(A) Holotype (SNSB-BSPG 1908 I 39a) from the early Tithonian Altmühltal Formation of Blumenberg near Eichstätt. (B) Counterpart of the holotype (SNSB-BSPG 1908 I 39b). (C) Male specimen (JME SOS 3149a) from the early Tithonian Altmühltal Formation of Blumenberg near Eichstätt displaying the pelvic claspers. (D) Holomorphic specimen (SMNK without number) displaying the well-preserved caudal fin from the early Tithonian Altmühltal Formation of Solnhofen. (E) Holomorphic specimen (JME SOS 4003) from the early Tithonian Altmühltal Formation of Wintershof.
Figure 8
Figure 8. Specimens of †Elasmodectes avitus (von Meyer, 1862 [120]) from the upper Kimmeridgian (Upper Jurassic) of Nusplingen.
(A) Disarticulated male specimen (SMNS 80142/16) of the quarry of Egesheim auf dem Westerberg displaying cranial remains including the frontal clasper and dorsal fin spine. This specimen was recovered in 1993 and figured by Duffin [78] (figure 4). (B) Partially preserved skeleton (SMNS 51564) displaying part of the vertebral column, the dorsal fins including the fin spine and parts of the pelvic girdle. The specimen was figured by Schweizer [66] (figure 14, pl. 12, figures 1–4). (C) Strongly disarticulated specimen (SMNS 86901/35) of the quarry of the Gesellschaft für Naturkunde Württemberg) displaying the left mandibular tooth plate.
Figure 9
Figure 9. †Chimaerotheca schernfeldensis Duffin et al., 2021 [99], also see [131] (NHMUK PV Z.183) egg capsule morphotypes from the Solnhofen Archipelago.
Figure 10
Figure 10. Articulated skeleton of †Hybodus fraasi Brown, 1900 [172] from the early Tithonian of Solnhofen.
(A) Complete specimen (SNSB-BSPG 1899 I 2, holotype). (B) Close-up view of head. (C) Ambient occlusion photogrammetric model of head (courtesy of Christoph Kettler).
Figure 11
Figure 11. Specimen of †Asteracanthus ornatissimus Agassiz, 1837 [171] from the lower Tithonian of Solnhofen.
(A) Complete specimen (PBP-SOL-8003). Close-up view of (B) anterior and (C) posterior dorsal fin spine. (D) Dentition.
Figure 12
Figure 12. Jaws and associated teeth of †Strophodus sp. from the early Tithonian of Mörnsheim.
(A,B) Complete specimen (SNSB-BSPG 2010 I 91). (C,D) ambient occlusion photogrammetric model (arrows point to symphysis connecting palatoquadrates; photogrammetric model courtesy of Christoph Kettler).
Figure 13
Figure 13. Specimens of †Sphenodus Agassiz, 1843 [171] found in the Solnhofen Archipelago.
(A) Holotype specimen of †S. macer (Quenstedt, 1851 [106]) (SMNS 80142-44) from the upper Kimmeridgian of Nusplingen. (B) Fragmentary specimen of †S. macer (MCZ 13389) from the Upper Jurassic of Solnhofen. (C) Holotype specimen of †S. nitidus Wagner, 1862 [105] (BSGP-AS VII 647) from the Tithonian of Solnhofen. (D) †S. nitidus (SMNS 96844-7) from the Kimmeridgian of Nusplingen.
Figure 14
Figure 14. †Synechodus ungeri Klug, 2009 [70].
(A) Holotype specimen (SMNS 85975-1) from the Kimmeridgian of Nusplingen. (B) Holomorphic specimen (JME SOS 3152 B) from the Tithonian of Solnhofen.
Figure 15
Figure 15. †Paraorthacodus jurensis (Schweizer, 1964 [66]).
(A) Holotype specimen of †P. jurensis (GPIT 1210/1,) found in the Tithonian of Nusplingen. (B) Holomorphic specimen (SMNS 88987-1) found in the Tithonian of Nusplingen. (C) Holomorphic specimen (SNSB-BSPG 1964 XXIII 157) from the lower Tithonian of Eichstätt. (D) Holomorphic specimen (SNSB-BSPG 1996 I 31) from the lower Tithonian of Eichstätt.
Figure 16
Figure 16. †Macrourogaleus hassei Fowler, 1947 [211]
(A) Holotype specimen (SNSB-BSPG AS I 1363) from the lower Tithonian of Eichstätt. (B) Close up to the enlarged denticles in the upper lobe of caudal fin. (C) Holomorphic specimen (SNSB-BSPG AS I 1362) from the lower Tithonian of Eichstätt. (D) Holomorphic specimen (JME SOS 2209) from the lower Tithonian of Eichstätt. (E) Holomorphic specimen (AMNH 7498) from the lower Tithonian of Eichstätt.
Figure 17
Figure 17. †Paracestracion falcifer Wagner, 1857 [100].
(A) Holotype specimen (SNSB-BSPGM AS VI 505) from the lower Tithonian of Solnhofen. (B) Holomorphic specimen (NHMUK P 8657) from the Kimmeridgian of Eichstätt. (C) Almost complete specimen (JME SOS 2215) from the Kimmeridgian of Eichstätt.
Figure 18
Figure 18. Holotype specimen of †Paracestracion viohli Kriwet, 2008 [57] (JME Scha 728) from the upper Kimmeridgian of Schamhaupten.
Figure 19
Figure 19. Holotype specimen of †Paracestracion danieli Slater, 2016 [221] (JPBP-SOL-0005) from the Kimmeridgian of Eichstätt.
Figure 20
Figure 20. †Heterodontus zitteli Eastman, 1911 [219].
(A) Holotype specimen (NHMUK P 6938) from the Kimmeridgian of Eichstätt. (B) Holomorphic specimen (SNSB-BSPG 1885 I 12a) from the Tithonian of Eichstätt. (C) Holomorphic specimen (JME SOS 3577) from the lower Tithonian between Wintershof and Workerszell. (D) Holomorphic specimen (JME SOS 3153) from the lower Tithonian from Blumenberg.
Figure 21
Figure 21. †Phorcynis catulina Thiollière, 1852 [229].
(A) Holotype specimen (NHMUK P 5541) from the lower Tithonian of Eichstätt. (B) Almost complete specimen (SNSB-BSPG 1990 XVIII 51) from the lower Tithonian of Zandt. (C) Close up to the cephalic region. (D) Holomorphic specimen (JME SOS 3150) from the lower Tithonian of Blumenberg.
Figure 22
Figure 22. Holotype of the species †Palaeorectolobus agomphius Kriwet, 2008 [58] (MB.f. 11191) from the lower Tithonian of Kelheim.
Figure 23
Figure 23. †Bavariscyllium tischlingeri Thies, 2005 [55]
(A) Holotype specimen (JME SOS 4124) from the lower Tithonian of Eichstätt. (B) Holomorphic specimen (SMF P272) from the lower Tithonian of Eichstätt. (C) Holomorphic specimen (SMNS 96086) from the lower Tithonian of Eichstätt.
Figure 24
Figure 24. Specimens of †Palaeoscyllium formosum Wagner, 1857 [102].
(A) Holotype specimen (SNSB-BSPG AS I 1365) from the lower Tithonian of Solnhofen. (B) Holomorphic specimen (SNSB-BSPG AS I 589 A) from the lower Tithonian of Solnhofen. (C) Holomorphic specimen (JME SOS 3151) from the lower Tithonian of Solnhofen. (D) Holomorphic specimen (SMNK, without number).
Figure 25
Figure 25. Scyliorhinid type egg capsule. †Scyliorhinotheca sp. Kiel et al., 2013 [251] from the Solnhofen Archipelago.
Figure 26
Figure 26. †Palaeocarcharias stromeri de Beaumont, 1960 [51]
(A) Holotype specimen (JME SOS 2294) from the lower Tithonian of Eichstätt. (B) Close up of the cephalic regions. (C) Close up of the anterior portion of the jaws.
Figure 27
Figure 27. †Crassodontidanus serranus Kriwet and Klug, 2011 [267].
(A) Holotype specimen labial view (SMNS 3695/10) from the late Kimmeridgian of Nusplingen. (B) Lingual view. (C) Disarticulated teeth (GPIT 81512) from the Kimmeridgian of Nusplingen. (D) Close up of a tooth in labial view. (E) Close up of a tooth in lingual view. †Notidanoides muensteri (Agassiz, 1843 [171]). (F) Teeth of labial view (JME SOS 2213) from the Tithonian of Eichstätt. (G) Teeth in lingual view (SNSB-BSPG 1989 X 12) from the Kimmeridgian of Daiting.
Figure 28
Figure 28. †Notidanoides muensteri (Agassiz, 1843 [170]).
(A) Incomplete specimen (GPIT Pi 1210/3) from the Kimmeridgian of Nusplingen. (B) Close up of the cephalic region.
Figure 29
Figure 29. Specimens of †Pseudorhina Jaekel, 1989 [276].
(A) Holotype specimen of †P. alifera (Münster, 1842 [50]) (SNSB-BSPG AS VII 3) from the lower Tithonian of Eichstätt. (B) †Pseudorhina acanthoderma (Fraas, 1854 [64]) (SMNS 80431/20) from the upper Kimmeridgian of Nusplingen. (C) Close up of the cephalic region in dorsal view of †P. acanthoderma (SMNS 86214/41) from the Kimmeridgian from Nusplingen. (D) Close up of the cephalic region in ventral view of †P. acanthoderma (SMNS 80431/20).
Figure 30
Figure 30. †Protospinax annectans Woodward, 1918 [232].
(A) Holotype specimen (NHMUK PV P 8775) from the lower Tithonian from Solnhofen. (B) Holomorphic specimen preserved in ventral view (SNSB-BSPG 1963 I 19) from the lower Tithonian of Eichstätt.
Figure 31
Figure 31. †Asterodermus platypterus Agassiz, 1936 [171].
(A) Holotype specimen (NHMUK P 12067) from the lower Tithonian of Kelheim. (B) Holomorphic specimen (JME SOS 2212 A) from the lower Tithonian of Blumenberg. (C) Partial specimen (SNSB-BSPG AS I 1378) from the Tithonian of Kelheim. (D) Partial specimen (JME SOS 3647) from the Tithonian of Birkhof. (E) Holomorphic specimen (SNSB-BSPG 1964 XXIII 577) from the Tithonian of Solnhofen.
Figure 32
Figure 32. †Spathobatis bugesiacus Thiollière, 1852 [229].
(A) Holomorphic specimen (BMMS BK 3-1) from the lower Tithonian of Eichstätt. (B) Holomorphic specimen (SNSB-BSPG AS I 505) from the lower Tithonian of Eichstätt. (C) Holomorphic specimen (NHMUK PV-P-6010) from the Tithonian of Solnhofen.
Figure 33
Figure 33. Putative batomorph type egg capsule from the Solnhofen Archipelago. †Rajitheca sp. Steininger, 1966 [322] (private collection).
Figure 34
Figure 34. Multidimensional scaling dispersion plot, showing the dissimilarity relation between localities and geological times.
Figure 35
Figure 35
(A) Rough reconstruction of Europe during the Late Jurassic (150 Ma), depicting the position of the localities studied. Paleogeography of Europe Series © 2020 Colorado Plateau Geosystems Inc. Scottsdale Arizona. (B) Heat map and dendrogram showing the dissimilarity relations between geological areas based on the presence and absence of taxa. Colours in dendrogram indicate the assigned clusters in the analysis. Heat map colours: White (presence), black (absence). Numbers in the map refer to the localities in the heat map and dendrogram.
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