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Volume 27.1
January–April 2024
Full table of contents
ISSN: 1094-8074, web version;
1935-3952, print version
Recent Research Articles
See all articles in 27.1 January-April 2024
See all articles in 26.3 September-December 2023
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Christoph Gierl. Department of Earth and Environmental Sciences, Ludwig-Maximilians-Universität München, Richard-Wagner Straße 10, 80333 Munich, Bavaria, Germany; c.gierl@lrz.uni-muenchen.de
Christoph Gierl is a palaeontologist and a scientific assistant at the Ludwig-Maximilians Universität (LMU) Munich. There he got his Master degree in Geological Sciences. His interests lie in taxonomy, systematics and palaeogeography of fossil gobiiforms. He is currently working on a project on their fossil record and phylogeny.
Bettina Reichenbacher. Department of Earth and Environmental Sciences, Ludwig-Maximilians-Universität München, Richard-Wagner Straße 10, 80333 Munich, Bavaria, Germany; b.reichenbacher@lrz.uni-muenchen.de
Bettina Reichenbacher is professor for palaeontology at LMU Munich. She is interested in the evolution of Miocene teleost groups in the context of the geological history. Current projects comprise the evolutionary history of the European Gobiiformes, the cichlid fossils from Miocene palaeolakes in the Central Kenya Rift and the Iranian killifish genus Aphanius.
Further information on the two latter projects can be found here:
http://www.en.palaeontologie.geowissenschaften.uni-muenchen.de/forschung/forsch_gebiete/kenya/index.html
http://www.killi-data.org/infoweb18.php
FIGURE 1. Phylogeny of the Gobiiformes according to Thacker et al. (2015) and Nelson et al. (2016). 6brG = gobiiforms with six branchiostegal rays, 5brG = gobiiforms with five branchiostegal rays.
FIGURE 2. †Paralates bleicheri Sauvage, 1883. 1. NMB Ruf. 13_1 skull. 2. NMB Ruf. 15b (Neotype, designated by Gaudant, 1979). 3. NMB Ruf. 15a. Abbreviations: chy = ceratohyal, d = dentary, f = frontal, mx = maxilla, meth = mesethmoid, op = opercle, pmx = premaxilla, pop = preopercle, q = quadrate, sop = subopercle. (Photos by M. Schellenberger.)
FIGURE 3. Comparison of frontals (highlighted by the black lines). 1. †Paralates bleicheri Sauvage, 1883, NMB Ruf. 13_1 skull mirrored. (Photo by M. Schellenberger.) 2. †Paralates chapelcorneri n. sp., NHMUK PV P 59786. Same scale for both skulls.
FIGURE 4. †Paralates bleicheri Sauvage, 1883, NMB Ruf. 18. 1. Viewed under normal light. (Photo by M. Schellenberger.) 2. Imaged with UV light (Photo by H. Tischlinger).
FIGURE 5. †Paralates chapelcorneri n. sp., NHMUK PV P 59786. 1. Skull. 2. Complete specimen. 3. Detail of opercle with cycloid scales. Abbreviations: art = anguloarticular, d = dentary, ect = ectopterygoid, f = frontal, meth = mesethmoid, mx = maxilla, op = opercle, pmx = premaxilla, pop = preopercle, q = quadrate, scl = supracleithrum.
FIGURE 6. †Paralates chapelcorneri n. sp., NHMUK PV P 59785. 1. Complete specimen. 2. Region of the first dorsal fin. Arrows point from the pterygiophores to the interneural spaces into which each inserts. Pterygiophore formula = 3-2121100 The spinous rays of the first dorsal fin are marked with roman numerals. Abbreviations: 1-7 = pterygiophores, ptm = post-temporal, ns-v3-10 = neural spines of vertebrae 3-10, v2-4 = vertebral centra 2-4.
FIGURE 7. †Paralates chapelcorneri n. sp., NHMUK PV P59785_counterpart. 1. Complete specimen. 2. Caudal skeleton and fin. Small Roman numerals in parentheses label unbranched, unsegmented rays, small Roman numerals indicate segmented unbranched rays, and Arabic numerals designate segmented, branched rays (after Fricke, 1983). Abbreviations: epu = epural, hpu = hemal spine of preural centrum, hy = hypural, npu = neural spine of preural centrum, phy = parhypural, pu = preural centrum, us = urostyle.
FIGURE 8. †Paralates chapelcorneri n. sp., NHMUK PV P 59784. 1. Complete specimen. 2. Skull showing possible symplectic and six branchiostegals. Abbreviations: 1-6 = branchiostegals, ect = ectopterygoid, mpt = metapterygoid, sf = suspensorium fenestra, sym = symplectic. (Photos by K. Webb; © The Trustees of the Natural History Museum, London)
TABLE 1. Meristic counts of all studied specimens of †Paralates . Numbers not in bold are subject to some uncertainty.
Oligocene of Rouffach, no. Ruf | Upper Eocene of Isle of Wight (Chapelcorner Fish Bed) |
||||||||||
Specimen # | 6a/b | 9_1 | 9_2 | 11/13_2 | 15a/b | 18 | P 59784 | P 59785 | P 59786 | C 23632 | C 23633 |
First dorsal fin (D1) | - | - | >V | - | VII | VII | VII | VII | VII | - | - |
Second dorsal fin (D2) | I,10 | I,9 | I,9 | I,9 | I,8 | I,10 | I,9 | I,10 | >7 | - | - |
Anal fin (A) | 10 | >8 | 10 | >7 | I,8 | I,8 | I,8 | I,9 | - | - | - |
C (branched, segmented) | 12 | 13 | ? | 13 | 12 | 12 | 12 | 13 | - | - | - |
C (unbranched, segmented) | 2 + 1 or 2 | 2 + 2 | |||||||||
C (unbranched, unsegmented) | 7 | 10 + ? | 10 + 12 | ||||||||
Ventral fin (V) | - | - | - | - | I,5 | - | I,5 | I,5 | - | - | - |
Pectoral fin (P) | - | >11 | 14 | >9 | 12 | - | >14 | 16 | >11 | >13 | >9 |
Vertebrae [postabdominal vert.] | >23 [17] | >21 [16] | >27 | >23 [17] | 29 [17] | 30 [18] | 30 [18] | 31 [19] |
>23 | >11 | 29 [17] |
Pterygiophore formula D1 | - | - | - | - | - | - | - | 3-2121100 | - | - | - |
TABLE 2. Morphometrics of the genus †Paralates in mm. Values standardized to standard length are given in brackets. Abbreviations: A = anal fin, C = caudal fin, D1 = first dorsal fin, D2 = second dorsal fin, D2C = distance between end of D2 and first procurrent ray of C, P = pectoral fin, V = pelvic fin.
# | Ruf. 11/13_2 | Ruf. 15a/b | Ruf. 18 | P 59784 | P 59785 | P 59786 |
Total length | 29.1 (115%) | 29.5 (118%) | 34.5 (119%) | 35.3 (116%) | 22.5 (116%) | >22.3 |
Standard length | 25.2 | 25.1 | 29.1 | 30.4 | 19.4 | >22.3 |
Max body height | 5.4 (21.4%) | 5.4 (21.5%) | 5.5 (18.9%) | 5.9 (19.4%) | 4.4 (22.7%) | - |
Head length | 9.4 (37.3%) | 6.9 (27.5%) | 7.1 (24.4%) | 9.5 (31.3%) | 5.6 (28.9%) | 6.6 |
Head height | 6.6 (26.1%) | 5.4 (21.5%) | 5.2 (17.9%) | 6.9 (22.7%) | 4.9 (25.3%) | - |
Horizontal eye diameter | 1.5 (5.9%) | 1.2 (4.8%) | 1.3 (4.5%) | - | 1.7 (8.8%) | 1.4 |
Dist snout to D1 | 11.7 (46.4%) | 9.2 (36.7%) | 10.8 (37.1%) | 11.7 (38.5%) | 7.8 (40.2%) | 7.9 |
Dist snout to D2 | 14.6 (57.9%) | 14.5 (57.8%) | 16.2 (55.7%) | 17.4 (57.2%) | 11.2 (57.7%) | 12.8 |
Dist snout to A | 15.8 (62.7%) | 15.7 (62.5%) | 16.5 (56.7%) | 18.1 (59.5%) | 11.5 (59,3%) | 13.3 |
Dist snout to P | 9.4 (37,3%) | 8.7 (34.7%) | - | 10.7 (35.2%) | 6.8 (35.1%) | 7.4 |
Dist snout to V | 9.2 (36.5%) | 8.2 (32.7%) | - | 10.6 (34.9%) | 6.6 (34.0%) | - |
Length D1 | >2.1 | >2.4 | >2.4 | >3.4 | >1.7 | - |
Length D2 | 2.8 (11.1%) | >2.4 | 4.4 (15.1%) | >4.3 | >3.1 | - |
Length A | 2.6 (10.3%) | >2.4 | >2.3 | >3.5 | >2.9 | - |
Length P | >0.8 | >1.3 | - | >1.9 | >0.8 | >2.8 |
Length V | >1.2 | >1.5 | - | >2.7 | 2.7 (13.9%) | - |
Base length D1 | >0.6 | 2.3 (9.2%) | 2.3 (7.9%) | 2.6 (8.6%) | 1.6 (8.2%) | - |
Base length D2 | 3.1 (12.3%) | 3.0 (12.0%) | 4.0 (13.7%) | >3.5 | 2.4 (12.4%) | - |
Base length A | 3.0 (11.9%) | 2.5 (19.0%) | 3.7 (12.7%) | 3.6 (11.8%) | 1.9 (9.8%) | - |
Length caudal peduncle | 6.3 (25.0%) | 6.8 (27.1%) | 8.9 (30.6%) | 8.3 (27.3%) | 5.7 (29.4%) | - |
D2C | 6.7 (26.6%) | 7.3 (29.1%) | 8.4 (28.6%) | 7.5 (24.7%) | 5.0 (25.8%) | - |
Min height of caud ped | 2.0 (7.9%) | 1.7 (6.8%) | 2.6 (8.9%) | 3.0( 9.7%) | 2.2 (11.3%) | - |
Length of caudal fin | 3.8 (15.1%) | >3.8 | >5.0 | 5.4 (17.8%) | 3.6 (18.6%) | - |
Length of vertebrae | 0.5-0.6 (1.9-2.3%) | 0.6-0.7 (2.4-2.8%) | 0.6-0.8 (2.1-2.7%) | 0.5-0.7 (1.6-2.3%) | 0.4-0.5 (2.1-2.6%) | 0.4-0.5 |
TABLE 3. Meristic characters of recent gobioid taxa for which the pterygiophore formula 3-212110(0) has been observed in at least one species of the genus. The number of caudal fin rays refers to all segmented rays (branched and unbranched). Abbreviations: A = anal fin, C = caudal fin, D1 = first dorsal fin, D2 = second dorsal fin, P = pectoral fin, V = pelvic fin.
Taxon | D1 | D2 | A | C | V | P | Vertebrae [postabd] | Source | Family |
†Paralates bleicheri Sauvage, 1883 | VII | I,10 | I,9 | >12-13 | I,5 | 14 | 29 [17-18?] |
this study | inc. sed. |
†Paralates chapelcorneri n. sp. | VII | I,10 | I,8-9 | 14-17 | I,5 | 14-16 | 30?-31 [18?-19] | this study | inc. sed. |
Gobiodon okinawae Sawada, Arai & Abe, 1972 | VI | I,10 | I,9 | 15 | - | - | 27 [17] |
unpublished data | Gobiidae |
Gobiosoma hemigymnum (Eigenmann and Eigenmann 1888) | VII | I,11 | I,9 | 17 | I,5 | 17-19 | 27 [16] |
van Tassell et al. 2015 | Gobiidae |
Gobiosoma spp. | - | - | - | - | - | - | 27-28 [15-17] |
Birdsong et al. 1988 | Gobiidae |
Gobulus spp. | - | - | - | - | - | - | 27-28 [16-17] |
Birdsong et al. 1988 | Gobiidae |
Gobulus spp. | VII-VIII | I,9-13 | I,7-13 | 15-17 | I,5 | 15-18 | 27-28 [16-17] |
Hoese and Reader 2001 | Gobiidae |
Gobiomorphus alpinus Stokell, 1962 | III-VII | I,7-9 | I,8-10 | - | - | 16-19 | 29-31 | McDowall 1994 | Eleotridae |
Gobiomorphus spp. | III-VIII | I,7-10 | I,7-10 | - | - | 13-21 | 27-31 | McDowall 1975 | Eleotridae |
Gobiomorphus spp. | - | - | - | - | - | - | 28-31 [16-19] |
Birdsong et al. 1988 | Eleotridae |
Thalasseleotris | 16 | 27 [17] |
Hoese and Larson 1987 | Thalasseleotrididae |
Revision of so-called Pomatoschistus (Gobiiformes, Teleostei) from the late Eocene and early Oligocene
Plain Language Abstract
Gobiiformes (or 'gobies') belong to the most diverse groups among vertebrates, and are found in both freshwater and marine habitats. However, it is often difficult to distinguish between morphologically similar forms. Moreover, species identification is based mostly on soft-tissue features, such as the pattern of sensory pores on the head, which makes fossil classification extremely challenging. Pomatoschistus bleicheri (Sauvage) from the lower Oligocene (~30 Ma old) of Rouffach (France) and Pomatoschistus(?) cf. bleicheri (Sauvage) from the upper Eocene (~35 Ma old) of the Isle of Wight (England) epitomize this problem. They are of special interest because they are regarded as the oldest fossil representatives of the more advanced gobies (Gobiidae + Oxudercidae). Here, we show that these fossils do not belong to the genus Pomatoschistus or indeed to the Gobiidae + Oxudercidae, nor can they be accommodated in any existing family of the "basal gobiiforms." Instead, the fossils from Rouffach and the Isle of Wight can both be placed in the same extinct genus, †Paralates Sauvage. Furthermore, differences in the shape of the frontal bones (on the forehead) and numbers of caudal fin rays lead to the conclusion that the material from Rouffach corresponds to †Paralates bleicheri Sauvage, while that from the Isle of Wight is assigned to a new species, which we name †Paralates chapelcorneri n. sp. Therefore, the oldest known fossil representative of the modern gobies remains †Gobius jarosi Přikryl and Reichenbacher, 2017 from the lower Miocene (20.4 Ma-19.1 Ma old) of Moravia (Czech Republic).
Resumen en Español
Revisión de los llamados Pomatoschistus (Gobiiformes, Teleostei) del Eoceno tardío y Oligoceno temprano
Los Gobiiformes (Teleostei) están entre los grupos más ricos en especies entre los vertebrados, y se encuentran en todos los hábitats acuáticos. La identificación de las especies actuales de Gobiiformes se basa principalmente en caracteres de los tejidos blandos, y su diversidad y variabilidad fenotípica a menudo hace que la determinación de las especies sea difícil. Los fósiles carecen normalmente de características de los tejidos blandos y la asignación segura de Gobiiformes fósiles a nivel de familia y género puede ser extremadamente problemática. "Pomatoschistus bleicheri (Sauvage)" del Oligoceno inferior de Rouffach (Francia) y "Pomatoschistus (?) cf. bleicheri (Sauvage)" del Eoceno superior de la Isla de Wight (Inglaterra) ejemplifican estas dificultades. Estos hallazgos son de especial interés, ya que pueden representar los miembros más antiguos, basados en el esqueleto fósil, de los Gobiidae + Oxudercidae (Gobiiformes con cinco branquiostegales, en adelante 5brG). El nuevo examen del material tipo de estas dos especies revela ahora la presencia de una premaxila con un proceso postmaxilar, lo que impide la asignación de estos fósiles al género Pomatoschistus. De hecho, ni siquiera pertenecen al grupo 5brG porque muestran seis branquiostegales. Concluimos que tanto los fósiles de Francia como los de Inglaterra pertenecen a †Paralates Sauvage. Las diferencias en la forma de los huesos frontales y el número de radios de la aleta caudal nos permiten asignar el material francés a †Pa. bleicheri Sauvage y los ejemplares ingleses a †Pa. chapelcorneri n. sp. Así, la especie 5brG más antigua conocida actualmente, basada en esqueletos articulados, es †Gobius jarosi Přikryl et Reichenbacher, 2017 del Mioceno inferior.
Palabras clave: Gobiiformes fósiles; nueva especie; Gobiidae; Oxudercidae; branquiostegales
Traducción: Enrique Peñalver (Sociedad Española de Paleontología)
Résumé en Français
Une révision de soi-disant Pomatoschistus (Gobiiformes, Teleostei) de l’Éocène récent et de l’Oligocène ancien
Les Gobiiformes (Teleostei) sont parmi les groupes de vertébrés les plus diversifiés (en nombre d’espèces) et sont trouvés dans tous les habitats aquatiques. L’identification des espèces actuelles de Gobiiformes est principalement basée sur les caractères des tissus mous, et leur diversité et leur variabilité phénotypique compliquent fréquemment les déterminations spécifiques. Les fossiles ne préservent généralement pas les caractères des tissus mous, et une attribution certaine des Gobiiformes fossiles aux rangs familiaux et génériques peut être extrêmement problématique. “Pomatoschistus bleicheri (Sauvage)” de l’Oligocène inférieur de Rouffach (France) et “Pomatoschistus(?) cf. bleicheri (Sauvage)” de l’Éocène supérieur de l’île de Wight (Angleterre) illustrent ces difficultés. Ces découvertes sont d’un intérêt particulier, car elles pourraient représenter les plus anciens membres du clade Gobiidae + Oxudercidae (Gobiiformes avec cinq branchiostèges, 5brG) basés sur des restes squelettiques. Une révision du matériel-type de ces deux espèces révèle la présence d’un prémaxillaire avec un processus postmaxillaire, ce qui exclut l’attribution de ces fossiles au genre Pomatoschistus. Ils n’appartiennent même pas au clade 5brG car ils présentent six branchiostèges. Nous concluons que ces fossiles de France et d’Angleterre appartiennent à †Paralates Sauvage. Des différences dans la forme des os frontaux et dans le nombre de rayons de la nageoire caudale nous permettent d’attribuer le matériel français à †Pa. bleicheri Sauvage et les spécimens anglais à †Pa. chapelcorneri n. sp. La plus ancienne espèce connue actuellement dans le clade 5brG et basée sur des squelettes articulés est donc †Gobius jarosi Přikryl et Reichenbacher, 2017 du Miocène inférieur.
Mots-clés : Gobiiformes fossiles ; nouvelle espèce ; Gobiidae ; Oxudercidae ; branchiostèges
Translator: Antoine Souron
Deutsche Zusammenfassung
In progress
Translator: Eva Gebauer
Arabic
Translator: Ashraf M.T. Elewa
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Review: The Princeton Field Guide to Mesozoic Sea Reptiles
The Princeton Field Guide to Mesozoic Sea Reptiles
Article number: 26.1.1R
April 2023