DOI: 10.5061/dryad.7pvmcvf36
title: Data from: New osteological characteristics identify the first stem sleeper goby (Gobioidei, Odontobutidae) from the Upper Eocene

description: Amongst Gobioidei, one of the most diverse orders of teleost fishes, the
 families Odontobutidae (sleeper gobies) and Rhyacichthyidae are considered
 ‘basal gobioids’ and are therefore particularly interesting for
 understanding gobioid evolutionary history. However, their fossil record
 currently consists of only three extinct odontobutid species (two
 skeleton-based, one otolith-based). The aim of the study associated with
 this dataset (Bauer et al. 2025 in Papers in Palaeontology) was to
 investigate the possible relationship of one of the oldest known gobioid
 fossils, †Paralates chapelcornerensis from the Upper Eocene of southern
 England, to these families. To achieve this, osteological differences
 between Odontobutidae and Rhyacichthyidae were newly investigated using 19
 extant species from museum collections. This taxon set included both
 genera of the Rhyacichthyidae, five genera of the Odontobutidae (including
 Terateleotris; representing 78% of all odontobutid genera), one genus of
 the Eleotridae (Eleotris Bloch & Schneider 1801), two of the
 Butidae (Oxyeleotris Bleeker 1874; Bostrychus Lacepède 1801), one of the
 Gobiidae (Gobius Linnaeus 1758), as well as Trichonotus Bloch &
 Schneider, 1801 (Trichonotidae) and Apogon Lacepède 1801 (Apogonidae).
 Both Trichonotus and Apogon were selected as outgroups because the
 Trichonotidae is the sister group of the Gobioidei, and Apogonidae is
 closely related to Trichonotidae + Gobioidei (Thacker et al. 2015). To
 ensure that characters were consistent between species and capable of
 distinguishing between genera, three species each of Odontobutis and
 Trichonotus were included. The analysis identified 17 osteological
 characters (15 newly described) that are different between Odontobutidae
 and Rhyacichthyidae. These characters were used for comparative morphology
 and phylogenetic analyses (maximum parsimony, Bayesian inference) to
 determine whether †Paralates can be assigned to either the Odontobutidae
 or Rhyacichthyidae. This Dryad dataset provides an osteological atlas
 (Supplementary Data S1 in Bauer et al. 2025 in Papers in Palaeontology)
 illustrating the studied skeletal structures for †Paralates,
 Rhyacichthyidae and Odontobutidae, Eleotris (Eleotridae), Oxyeleotris and
 Bostrychus (both Butidae), Gobius (Gobiidae) and the used outgroups
 Trichonotus and Apogon.
description: Study of fossil
 material The fossil material included 51 articulated
 fossils, &gt; 100 isolated bones and four isolated otoliths of
 †<em>Paralates</em> from the Upper Eocene of the Chapelcorner
 Fish Bed (Chale, Isle of Wight, UK). The fossil material was studied and
 photographed under a Leica M165 C microscope equipped with a Jenoptik
 Gryphax Naos microscope camera. Photos were edited in Photoshop CS6 and
 Krita. The corresponding figures were prepared with Photoshop CS6 and
 Inkscape. Study of recent
 comparative material A total of 19 extant species were
 newly investigated. They encompass:
 Apogonidae<br> <em>Sphaeramia nematoptera</em>
 (Bleeker 1856)<br> <em>Apogon imberbis</em> (Linnaeus
 1758) Trichonotidae<br> <em>Trichonotus
 arabicus</em> Randall & Tarr 1994<br>
 <em>Trichonotus</em> cf. <em>setiger</em> Bloch
 & Schneider 1801<br> <em>Trichonotus nikii</em>
 Clark & von Schmidt 1966
 Rhyacichthyidae<br> <em>Protogobius attiti</em>
 Watson & Pöllabauer 1998<br> <em>Rhyacichthys
 guilberti</em> Dingerkus & Séret 1992<br>
 <em>Rhyacichthys aspro</em> (Valenciennes, 1837)
 Odontobutidae<br> <em>Microdous chalmersi</em>
 Nichols & Pope 1927<br> <em>Micropercops
 cinctus</em> (Dabry de Thiersant 1872)<br>
 <em>Odontobutis hikimius</em> Iwata & Sakai
 2002<br> <em>Odontobutis obscurus</em> Temminck
 & Schlegel 1845<br> <em>Odontobutis
 platycephala</em> Iwata & Jeon 1985<br>
 <em>Perccottus glenii</em> Dybowski 1877<br>
 <em>Terateleotris aspro</em> (Kottelat 1998)
 Eleotridae<br> <em>Eleotris pisonis</em>
 (Gmelin 1789) Butidae<br> <em>Bostrychus
 africanus</em> (Steindachner 1879)<br> <em>Oxyeleotris
 marmorata</em> (Bleeker 1852) Gobiidae<br>
 <em>Gobius incognitus</em> Kovačić & Šanda
 2016 For museum collection numbers of the species see
 Table 1 in Bauer et al. (2025) in <em>Papers in
 Palaeontology</em>. New micro-CT (µCT) scans were
 obtained for species that were not available on Morphosource from previous
 works. Only for <em>Sphaeramia nematoptera</em>,
 <em>Rhyacichthys aspro</em> and <em>Bostrychus
 africanus</em> no µCt scan could be acquired and these species were
 examined based on X-ray imaging. The software Amira 3D 2021.2 (Stalling
 <em>et al.</em> 2005) was used for analysis and segmentation
 of individual bones as well as generation of 3D models. Images of the
 models were made with the in-program snapshot function.
 A new MorphoSource project was created that contains µCT scans or
 X-ray images for all studied species as well as models of the studied
 bones. The reference is:<br> Bauer, E., Cerwenka, A. E., Schliewen,
 U. K. and Reichenbacher, B. 2025. Project: New osteological
 characteristics identify the first stem sleeper goby (Gobioidei,
 Odontobutidae) from the Upper Eocene [dataset]. MorphoSource. https://www.morphosource.org/projects/000642675 For MorphoSource DOI numbers of the µCT scans of the species see also Table 1 in Bauer et al. (2025) in <em>Papers in Palaeontology</em>. Osteological study The fossil material and µCt scans of the extant comparative specimens were used for osteological study. Six bony structures that were well-preserved in the fossil material of †<em>Paralates</em> were the focus: the first vertebra (atlas), the premaxilla, the palatine, the urohyal, the hyoid arch, and the pelvic girdle. These elements were segmented from the µCTs of the extant species and visualized in 3D. The outcome served to identify structures and character states that differed between Odontobutidae and Rhyacichthyidae. The results are compiled in an osteological atlas comprising eight plates (Supplementary Data S1 in Bauer et al. 2025 in <em>Papers in Palaeontology</em>). The original TIFF files of this atlas are provided in the Dryad dataset.
description: # Data from: New osteological characteristics identify the first stem
 sleeper goby (Gobioidei, Odontobutidae) from the Upper Eocene
 [https://doi.org/10.5061/dryad.7pvmcvf36](https://doi.org/10.5061/dryad.7pvmcvf36) ## Description of the data and file structure The fossil material and the µCt scans of the extant comparative material were used for osteological study. The focus was study of six bony structures that were well-preserved in the fossil material of †*Paralates*: the first vertebra (atlas), the upper jaw bone (premaxilla), the palatine, the urohyal, the hyoid arch, and the pelvic girdle. These bones were segmented from the µCTs of the extant species and visualized in 3D. The outcome served to identify structures and character states that differed between Odontobutidae and Rhyacichthyidae. To ensure that characters were consistent between species and capable of distinguishing between genera, three species of *Odontobutis* and three species of *Trichonotus* were included. The results are compiled in an osteological atlas comprising eight plates (=Supplementary Data S1 in Bauer et al. 2025 in *Papers in Palaeontology*). The original TIFF files of this osteological atlas are presented here in the Dryad dataset. **Atlas-Figures 1–6** illustrate a single morphological feature of the following specimens. Panel labels refer to species as follows: A, †*Paralates* sp. (A1-3 NHMUK PV P 77141) B, *Apogon imberbis* (ZSM 43837) C, *Trichonotus* cf. *setiger* (ZSM 43097) D, *Protogobius attiti* (MNHN-2019-0112) E, *Rhyacichthys guilberti* (MNHN-2019-0113-2) F, *Odontobutis platycephala* (NHMUK 1983.11.4.28-29) G, *Perccottus glenii* (ZSM 39696) H, *Microdous chalmersi* (NHMUK 1937.9.17.41-42) I, *Micropercops cinctus* (ZSM 39647) J, *Terateleotris aspro* (ZSM 43742) K, *Eleotris pisonis* (ZSM 41704) L, *Oxyeleotris marmorata* (ZSM 46847) M, *Gobius incognitus* (NMP 6V 146150). 
 
 The illustrated elements are: **Atlas-Figure_1.** First vertebra (atlas)
 **Atlas-Figure_2.** Premaxilla **Atlas-Figure_3.** Pelvic girdle and
 pelvic–pectoral articulation **Atlas-Figure_4.** Palatine
 **Atlas-Figure_5.** Urohyal **Atlas-Figure_6.** Hyoid arch. 
 
 **Atlas-Figures 7 and 8** illustrate that in three species each of
 *Odontobutis* and *Trichonotus*, the six studied bones were consistent
 between species and thus suitable for distinguishing genera.
 **Atlas-Figure_7.** Comparison of the first vertebra (atlas), premaxilla,
 pelvic girdle and pelvic–pectoral articulation, palatine, urohyal and
 hyoid arch among three species of *Odontobutis*. **Atlas-Figure_8.**
 Comparison of the first vertebra (atlas), premaxilla, pelvic girdle and
 pelvic–pectoral articulation, palatine, urohyal and hyoid arch among three
 species of *Trichonotus*. ## Captions of Atlas figures **ATLAS-FIG. 1.**
 First vertebra (atlas) of †*Paralates* sp. (NHMUK PV P 77141, arrows
 indicate anterior) and μCT scans of the first vertebra and character
 states of *Apogon imberbis*, *Trichonotus* cf. *setiger*, species of the
 Rhyacichthyidae and Odontobutidae, *Eleotris pisonis* (Eleotridae),
 *Oxyeleotris marmorata* (Butidae) and *Gobius incognitus* (Gobiidae).
 First vertebrae shown in: 1, anterior; 2, lateral (anterior to the left);
 3, dorsal view. All scale bars 1 mm. **ATLAS-FIG. 2.** Premaxilla of
 †*Paralates* *chapelcornerensis*/sp. (A1–3= NHMUK PV P 77105, A4,5= NHMUK
 PV P 77026, arrows indicate anterior) and μCT scans and character states
 of the premaxilla of *Apogon imberbis*, *Trichonotus* cf. *setiger*,
 species of the Rhyacichthyidae and Odontobutidae, *Eleotris pisonis*
 (Eleotridae), *Oxyeleotris marmorata* (Butidae) and *Gobius incognitus*
 (Gobiidae). Premaxillae shown in: 1, medial; 2, dorsal; 3, lateral view.
 All scale bars 1 mm. **ATLAS-FIG. 3.** Pelvic girdle and pelvic–pectoral
 articulation of †*Paralates* *chapelcornerensis* (NHMUK PV P 77028a) shown
 in: A1, dorso-ventral, A2, lateral, A3, dorsal view (arrows indicate
 anterior), and μCT scans and character states of the pelvic girdle and
 pelvic-pectoral articulation of *Apogon imberbis*, *Trichonotus* cf.
 *setiger*, species of the Rhyacichthyidae and Odontobutidae, *Eleotris
 pisonis* (Eleotridae), *Oxyeleotris marmorata* (Butidae) and *Gobius
 incognitus* (Gobiidae). Pelvic girdle for the extant species shown in:
 1, dorsal (anterior to the left), 2, lateral (anterior to the left),
 3, ventral view (anterior to the right); pelvic–pectoral articulation
 shown in: 4, lateral view (anterior to the left). Inset upper right corner
 shows generalized bauplan of pelvic bone for Rhyacichthyidae and
 Odontobutidae. All scale bars 1 mm. **ATLAS-FIG. 4.** Palatine of
 †*Paralates* sp. (NHMUK PV P 77036b, arrow indicates anterior) and μCT
 scans and character states of the palatine of *Apogon imberbis*,
 *Trichonotus* cf. *setiger*, species of the Rhyacichthyidae and
 Odontobutidae, *Eleotris pisonis* (Eleotridae), *Oxyeleotris marmorata*
 (Butidae) and *Gobius incognitus* (Gobiidae). Palatines shown in:
 1, lateral (anterior is upper right); 2, dorsal; 3, medial view (anterior
 is upper left). All scale bars 1 mm. **ATLAS-FIG. 5.** Urohyal of
 †*Paralates chapelcornerensis* (A1= NHMUK PV P 76993a, arrow indicates
 anterior), and μCT scans and character states of the urohyal of *Apogon
 imberbis*, *Trichonotus* cf. *setiger*, species of the Rhyacichthyidae and
 Odontobutidae, *Eleotris pisonis* (Eleotridae), *Oxyeleotris marmorata*
 (Butidae), and *Gobius incognitus* (Gobiidae). Urohyals shown in:
 1, lateral; 2, ventral view; anterior to the left. **ATLAS-FIG. 6.** Hyoid
 arch of †*P*. *chapelcornerensis* (NHMUK PV P 76990, arrow indicate
 anterior) and μCT scans and character states of the hyoid arch of *Apogon
 imberbis*, *Trichonotus* cf. *setiger*, species of the Rhyacichthyidae and
 Odontobutidae, *Eleotris pisonis* (Eleotridae), *Oxyeleotris marmorata*
 (Butidae), and *Gobius incognitus* (Gobiidae). Hyoid arches of extant
 species in: 1, lateral (anterior to the right); 2, dorsal (anterior to the
 left); 3, medial view (anterior to the left). All scale bars 1 mm.
 **ATLAS-FIG. 7.** Comparison of osteological structures and character
 states among three species of *Odontobutis*: *O. obscurus* (NHMUK
 1983.11.4.14-17), *O. platycephala* (NHMUK 1983.11.4.28-29) and
 *O. hikimius* (NHMUK 1999.4.20.1-2). A, urohyal in: A1, lateral;
 A2, ventral view. B, premaxilla in: B1, lateral; B2, dorsal; B3, medial
 view. C, palatine in: C1, lateral; C2, dorsal; C3, medial view. D, hyoid
 arch in: D1, lateral; D2, dorsal; D3, medial view. E, atlas in:
 E1, anterior; E2, lateral; E3, dorsal view. F, pelvic girdle in:
 F1, dorsal; F2, anterolateral; F3, ventral view. Arrows indicate anterior
 direction. All scale bars 1 mm. **ATLAS-FIG. 8.** Comparison of
 osteological structures among three species of *Trichonotus*: *T. nikii*
 (SMF 10255_1), *T*. cf. *setiger* (ZSM 43097) and *T. arabicus* (SMF
 26994). A, urohyal in: A1, lateral; A2, ventral view. B, premaxilla in:
 B1, lateral; B2, dorsal; B3, medial view. C, palatine in: C1, lateral;
 C2, dorsal; C3, medial view. D, hyoid arch in: D1, medial; D2, dorsal;
 D3, lateral view. E, atlas in: E1, anterior; E2, lateral; E3, dorsal view.
 F, pelvic girdle in: F1, dorsal; F2, anterolateral; F3, ventral view.
 Arrows indicate anterior direction. All scale bars 1 mm. **TABLE 1.** List
 of the studied species of the comparative material. 3D models of the
 investigated bones of the extant species are available from the
 MorphoSource project 000642675 of Bauer *et al*. (2025). See:
 [https://www.morphosource.org/projects/000642675?locale=en](https://www.morphosource.org/projects/000642675?locale=en) | **Species, collection number** | **Family** | **Collection number, MorphoSource DOI** | **Molecular data** | | :-------------------------------------------------- | :-------------- | :------------------------------------------------------------------------------------ | :-------------------- | | *Sphaeramia nematoptera* (Bleeker 1856) | Apogonidae | ZSM GO-0621, -0634; DOI: [10.17602/M2/M701284](https://doi.org/10.17602/M2/M701284) | Gierl *et al*. 2022 | | *Apogon imberbis* (Linnaeus, 1758) | Apogonidae | ZSM 43837; DOI: [10.17602/M2/M704361](https://doi.org/10.17602/M2/M704361) | Supplementary Data S2 | | *Trichonotus arabicus* Randall & Tarr, 1994 | Trichonotidae | SMF 26994; DOI: [10.17602/M2/M680497](https://doi.org/10.17602/M2/M680497) | - | | *Trichonotus* cf. *setiger* Bloch & Schneider, 1801 | Trichonotidae | ZSM 43097; DOI: [10.17602/M2/M701109](https://doi.org/10.17602/M2/M701109) | Supplementary Data S2 | | *Trichonotus nikii* Clark & von Schmidt, 1966 | Trichonotidae | SMF 10255(1); DOI: [10.17602/M2/M681004](https://doi.org/10.17602/M2/M681004) | - | | *Protogobius attiti* Watson & Pöllabauer, 1998 | Rhyacichthyidae | MNHN-2019-0112; DOI: [10.17602/M2/M123811](https://doi.org/10.17602/M2/M123811) | Gierl *et al*. 2022 | | *Rhyacichthys guilberti* Dingerkus & Séret, 1992 | Rhyacichthyidae | MNHN-2019-0113-2; DOI: [10.17602/M2/M123809](https://doi.org/10.17602/M2/M123809) | Gierl *et al*. 2022 | | *Rhyacichthys aspro* (Valenciennes, 1837) | Rhyacichthyidae | ZSM 44262; DOI: [10.17602/M2/M701274](https://doi.org/10.17602/M2/M701274) | Gierl *et al*. 2022 | | *Microdous chalmersi* Nichols & Pope, 1927 | Odontobutidae | BMNH 1937.9.17.41-42; DOI: [10.17602/M2/M689432](https://doi.org/10.17602/M2/M689432) | Supplementary Data S2 | | *Micropercops cinctus* (Dabry de Thiersant, 1872) | Odontobutidae | ZSM 39647; DOI: [10.17602/M2/M642729](https://doi.org/10.17602/M2/M642729) | Supplementary Data S2 | | *Odontobutis hikimius* Iwata & Sakai, 2002 | Odontobutidae | BMNH 1999.4.20.1-2; DOI: [10.17602/M2/M683184](https://doi.org/10.17602/M2/M683184) | Supplementary Data S2 | | *Odontobutis obscurus* Temminck & Schlegel, 1845 | Odontobutidae | BMNH 1983.11.4.14-17; DOI: [10.17602/M2/M689065](https://doi.org/10.17602/M2/M689065) | Gierl *et al*. 2022 | | *Odontobutis platycephala* Iwata & Jeon, 1985 | Odontobutidae | BMNH 1983.11.4.28-29; DOI: [10.17602/M2/M683240](https://doi.org/10.17602/M2/M683240) | Supplementary Data S2 | | *Perccottus glenii* Dybowski, 1877 | Odontobutidae | ZSM 39696; DOI: [10.17602/M2/M682159](https://doi.org/10.17602/M2/M682159) | Gierl *et al*. 2022 | | *Terateleotris aspro* (Kottelat, 1998) | Odontobutidae | ZSM 43742; DOI: [10.17602/M2/M681010](https://doi.org/10.17602/M2/M681010) | - | | *Eleotris pisonis* (Gmelin, 1789) | Eleotridae | ZSM 41704; DOI: [10.17602/M2/M123753](https://doi.org/10.17602/M2/M123753) | Gierl *et al*. 2022 | | *Bostrychus africanus* (Steindachner, 1879) | Butidae | ZSM 25113; DOI: [10.17602/M2/M701279](https://doi.org/10.17602/M2/M701279) | Supplementary Data S2 | | *Oxyeleotris marmorata* (Bleeker, 1852) | Butidae | ZSM 46847; DOI: [10.17602/M2/M680927](https://doi.org/10.17602/M2/M680927) | Gierl *et al*. 2022 | | *Gobius incognitus* Kovačić & Šanda, 2016 | Gobiidae | NMP 6V 146150; DOI: [10.17602/M2/M123810](https://doi.org/10.17602/M2/M123810) | Supplementary Data S2 | | †*Paralates bleicheri* Sauvage, 1883 | - | see Gierl & Reichenbacher (2017) | - | ### Files and variables #### File: Archive.zip **Description:** TIFF files of the Atlas-Figures 1–8 ## Code/software Adobe Reader or any Preview application can be used to open the files ## Access information Other publicly accessible locations of the data: * na Data was derived from the following sources: * na ##