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FIG. 15 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 15. — Viperinae ('Oriental vipers' group) from the early Miocene (MN 4) of MWQ. Middle trunk vertebra Pal. 1994 (2/2003 Reptile Joint) in right lateral (A), dorsal (B), ventral (C), cranial (D) and caudal (E) views. Posterior trunk vertebra Pal. 1506 (1/2001 Turtle Joint) with an indication of bifurcation in the distal tip of the hypapophysis in left lateral (F), dorsal (G), ventral (H), cranial (I) and caudal (J) views. Abbreviations: cd, condyle; ct, cotyle; d, diapophysis; hy, hypapophysis; na, neural arch; nc, neural canal; pctf, paracotylar foramen; pof, postzygapophyseal articular facet; pp, parapophyseal process; pr, prezygapophysis; prf, prezygapophyseal articular facet; prp, prezygapophyseal process; scf, subcentral foramen; scr, subcentral ridge; sctt, subcotylar tubercle; zy, zygosphene; zyg, zygantrum. Scale bars: 5 mm.

FIG. 13 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 13. — Vipera sp. ('European vipers' group) from the early Miocene (MN 4) of MWQ. Trunk vertebra Pal. 2022 (2/2003 Reptile Joint) in right lateral (A), dorsal (B), ventral (C) and cranial (D) views. Trunk vertebra Pal. 2023 (2/2003 Reptile Joint) in left lateral (E), dorsal (F), cranial (G) and caudal (H) views. Abbreviations: cd, condyle; ct, cotyle; hy, hypapophysis; nc, neural canal; ns, neural spine; pctf, paracotylar foramen; pof, postzygapophyseal articular facet; prf, prezygapophyseal articular facet; prp, prezygapophyseal process; scg, subcentral groove; scr, subcentral ridge; zy, zygosphene; zyg, zygantrum. Scale bars: 2 mm.

FIG. 10 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 10. — "Colubrinae" indet., type 1 from the early Miocene (MN 4) of MWQ. Middle trunk vertebra Pal. 1464 (1/2001 Turtle Joint) in right lateral (A), dorsal (B), ventral (C), cranial (D) and caudal (E) views. Middle trunk vertebra Pal. 1465 (1/2001 Turtle Joint) in right lateral (F), dorsal (G), ventral (H), cranial (I) and caudal (J) views. Posterior trunk vertebra Pal. 1466 (1/2001 Turtle Joint) in left lateral (K) and ventral (L) views. Caudal vertebra Pal. 1966 (2/2003 Reptile Joint) in right lateral (M), dorsal (N), ventral (O) and cranial (P) views. Abbreviations: cd, condyle; ct, cotyle; d, diapophysis; es, epizypapophyseal spine; hae, haemapophysis; hk, haemal keel; lf, lateral foramen; na, neural arch; nc, neural canal; ns, neural spine; p, parapophysis; pctf, paracotylar foramen; pl, pleurapophysis; pof, postzygapophyseal articular facet; prf, prezygapophyseal articular facet; prp, prezygapophyseal process; scf, subcentral foramen; scg, subcentral groove; scr, subcentral ridge; zy, zygosphene; zyg, zygantrum. Scale bar: 2 mm.

FIG. 12 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 12. — Natrix sp. and "Natricinae" indet. from the early Miocene (MN 4) of MWQ. Natrix sp.: trunk vertebra Pal. 1968 (2/2003 Reptile Joint) in left lateral (A), dorsal (B), ventral (C), cranial (D) and caudal (E) views. "Natricinae" indet.: trunk vertebra Pal. 1483 (1/2001 Turtle Joint) with preserved hypapophysis in left lateral (F), dorsal (G), ventral (H) and caudal (I) views. Abbreviations: ak, anterior keel; cd, condyle; ct, cotyle; es, epizypapophyseal spine; hy, hypapophysis; lf, lateral foramen;nc, neural canal; ns, neural spine; pctf, paracotylar foramen; po, postzygapophysis;pof, postzygapophyseal articular facet; prf, prezygapophyseal articular facet; prp, prezygapophyseal process; scf, subcentral foramen; scg, subcentral groove; scr, subcentral ridge; zy, zygosphene; zyg, zygantrum. Scale bars: 2 mm.

FIG. 9 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 9. — Coluber (s.l.) sp., from the early Miocene (MN 4) of MWQ. Anterior trunk vertebra Pal. 1454 (1/2001 Turtle Joint) in right lateral (A), dorsal (B), ventral (C), cranial (D) and caudal (E) views. Middle trunk vertebra Pal. 1456 (1/2001 Turtle Joint) in left lateral (F), dorsal (G), ventral (H), cranial (I) and caudal (J) views. Anterior caudal vertebra Pal. 1462 (1/2001 Turtle Joint) in right lateral (K), dorsal (L), ventral (M), cranial (N) and caudal (O) views. Abbreviations: cd, condyle; ct, cotyle; d, diapophysis; hae, haemapophysis; hk, haemal keel; hy, hypapophysis; lf, lateral foramen; na, neural arch; nc, neural canal; ns, neural spine; p, parapophysis; pctf, paracotylar foramen; pl, pleurapophysis; pof, postzygapophyseal articular facet; prf, prezygapophyseal articular facet; prp, prezygapophyseal process; scf, subcentral foramen; scg, subcentral groove; scr, subcentral ridge; zy, zygosphene; zyg, zygantrum. Scale bars: 2 mm.

FIG. 8 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 8. — Python sp. from the early Miocene (MN 4) of MWQ. Posterior trunk vertebra Pal. 1449 (1/2001 Turtle Joint) in right lateral (A), dorsal (B), ventral (C), cranial (D) and caudal (E) views. Abbreviations: cd, condyle; ct, cotyle; hk, haemal keel; na, neural arch; nc, neural canal; ns, neural spine; pd, paradiapophysis; po, postzygapophysis; pof, post zygapophyseal articular facet; pr, prezygapophysis; prf, prezygapophyseal articular facet; scf, subcentral foramen; scg, subcentral groove; scr, subcentral ridge; zy, zygosphene; zyf, zygosphenal facet; zyg, zygantrum. Scale bar: 5 mm.

FIG. 7 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 7. — Bavarioboa cf. hermi from the early Miocene (MN 4) of MWQ. Posterior trunk vertebra Pal. 1448 (1/2001 Turtle Joint) in left lateral (A), dorsal (B), ventral (C), cranial (D) and caudal (E) views. Abbreviations: ct, cotyle; hk, haemal keel; ir, interzygapophyseal ridge; na, neural arch; nc, neural canal; ns, neural spine; pd, paradiapophysis; pof, postzygapophyseal articular facet; prf, prezygapophyseal articular facet; prp, prezygapophyseal process; scf, subcentral foramen; scg, subcentral groove; scr, subcentral ridge; zy, zygosphene; zyg, zygantrum. Scale bar: 2 mm.

FIG. 6 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 6. — Ophisaurus and Anguinae indet. from the early Miocene (MN 4) of MWQ. Trunk vertebra Pal. 1407 (1/2001 Turtle Joint) of Ophisaurus sp. in (A) lateral, (B) dorsal, (C) ventral, (D) anterior and (E) posterior aspects. Anguinae indet.: right coronoid Pal. 1408 (1/2001 Turtle Joint) in (F) lateral, (G) medial and (H) dorsal aspects. Osteoderms Pal. 1577-1579 (2/2003 Reptile Joint) in (I-K) external aspects. Abbreviations: alp, anterolateral process; amp, anteromedial process; cd, condyle; ct, cotyle; dp, dorsal process; mr, medial ridge; muc, muscular crest; nc, neural canal; ns, neural spine; nt, notch; os, overlap surface; pof, postzygapophyseal articular facet; pp, posterior process; prf, prezygapophyseal articular facet; rd, ridge; scs, scupltured surface; syn, synapophysis. Scale bars: 2 mm.

FIG. 11 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 11. — Colubridae gen. et sp. indet. from the early Miocene (MN 4) of MWQ. Trunk vertebra Pal. 1484 (1/2001 Turtle Joint) in right lateral (A), dorsal (B) and cranial (C) views. Abbreviations: ct, cotyle; d, diapophysis; lf, lateral foramen; na, neural arch; nc, neural canal; ns, neural spine; pctf, paracotylar foramen; prf, prezygapophyseal articular facet; zy, zygosphene. Scale bar: 2 mm.

FIG. 3 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 3. — Amphisbaenia indet. from the early Miocene (MN 4) of MWQ. The trunk vertebra Pal. 1570 (2/2003 Reptile Joint) in (A) lateral, (B) dorsal, (C) ventral, (D) anterior and (E) posterior aspects. Abbreviations: cd, condyle; ct, cotyle; nc, neural canal; po, postzygapophysis; pof, postzygapophyseal articular fac- et; pr, prezygapophysis; prf, prezygapophyseal articular facet; scf, subcentral foramen; syn, synapophysis. Scale bar: 1 mm.

FIG. 16 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 16. — Elapidae gen. et sp. indet. from the early Miocene (MN 4) of MWQ. Trunk vertebra Pal. 1500 (1/2001 Turtle Joint) in right lateral (A), dorsal (B), ventral (C), cranial (D) and caudal (E) views. Abbreviations: cd, condyle; ct, cotyle; hy, hypapophysis; na, neural arch; nc, neural canal; ns, neural spine; pctf, paracotylar foramen; pof, postzygapophyseal articular facet; prf, prezygapophyseal articular facet; scg, subcentral groove; scr, subcentral ridge; sctt, subcotylar tubercle; zyg, zygantrum. Scale bar: 2 mm.

FIG. 14 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 14. — Viperinae ('Oriental vipers' group) from the early Miocene (MN 4) of MWQ. Left maxilla Pal. 1501 (1/2001 Turtle Joint) in rostral (A) and caudal (B) views. Fang Pal. 1502 (1/2001 Turtle Joint) in anterolateral (C) view. Anterior trunk vertebra Pal. 1503 (1/2001 Turtle Joint) in right lateral (D), dorsal (E), ventral (F) and cranial (G) views. Trunk vertebra Pal. 1504 (1/2001 Turtle Joint) in right lateral (H) and cranial (I) views. Vertebra from the middle part of the trunk section Pal. 1505 (1/2001 Turtle Joint) in right lateral (J), dorsal (K), ventral (L), cranial (M) and caudal (N) views. Abbreviations: can.den, dental canal; cd, condyle; ct, cotyle; fac. ecpt, facet for contact with ectopterygoid; hy, hypapophysis; ir, interzygapophyseal ridge; na, neural arch; nc, neural canal; ns, neural spine; pctf, paracotylar foramen; pd, paradiapophysis; po, postzygapophysis; pof, postzygapophyseal articular facet; pp, parapophyseal process; pr, prezygapophysis; pr.asc, ascending process; prf, prezygapophyseal articular facet; r, ridge; scf, subcentral foramen; scg, subcentral groove; scr, subcentral ridge; sctc, subcotylar canal; zy, zygosphene; zyg, zygantrum. Scale bars: A-C, 2 mm; D-N, 5 mm.

FIG. 2 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 2. — Lacertidae indet. from the early Miocene (MN 4) of MWQ. Left frontal Pal. 1565 (2/2003 Reptile Joint) in (A) dorsal and (B) ventral aspects. Lacertidae indet. tooth morphotype 1: right maxilla Pal. 1566 (2/2003 Reptile Joint) in (C) lateral and (D) medial aspects with detail of teeth. Left dentary Pal. 1567 (2/2003 Reptile Joint) in (E) lateral and (F) medial aspects with detail of teeth. Right dentary Pal. 1568 (2/2003 Reptile Joint) in (G) medial aspect with detail of teeth. Lacertidae indet. tooth morphotype 2: left dentary Pal. 1401 (1/2001 Turtle Joint) in (H) medial aspect with detail of teeth. Abbreviations: dc, dental crest; fs, frontal shield; lf, labial foramen (foramina); lp, lateral process; mc, mesial cusp; Mg, Meckel`s groove; mp, medial process; prs, prefrontal shield; sbs, subdental shelf; sc, sulcus; srs, supradental shelf; st, striations. Scale bars: A-D, H, 1 mm; D', G', 100 µm; E-G, H', 500 µm; F', 200 µm.

FIG. 1 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 1. — Topographic position of the Mokrá-Western Quarry with 1/2001 Turtle Joint and 2/2003 Reptile Joint (A). Position of the 1/2001 Turtle Joint at floor 380 m a.s.l. and field excavation of the fissure deposits in 2002 (B). The 2/2003 Reptile Joint has been discovered only few meters on the left from the 1/2001 Turtle Joint (modified after Ivanov et al. 2006).

FIG. 5 in Early Miocene squamate assemblage from the Mokrá-Western Quarry (Czech Republic) and its palaeobiogeographical and palaeoenvironmental implications

FIG. 5. — Pseudopus from the early Miocene (MN 4) of MWQ. Pseudopus laurillardi: parietal Pal. 1574 (2/2003 Reptile Joint) in (A) dorsal and (B) ventral aspects; parietal Pal. 1402 (1/2001 Turtle Joint) in (C) dorsal and (D) ventral aspects. Pseudopus sp.: left maxilla Pal. 1575 (2/2003 Reptile Joint) in (E) lateral and (F) medial aspects. Left quadrate Pal. 1576 (2/2003 Reptile Joint) in (G) lateral and (H) medial aspects. Right dentary Pal. 1403 (1/2001 Turtle Joint) in (I) lateral, (J) medial and (K) dorsal aspects. Trunk vertebra Pal. 1404 (1/2001 Turtle Joint) in (L) lateral, (M) dorsal, (N) ventral, (O) anterior and (P) posterior aspects. Trunk vertebra 1405 (1/2001 Turtle Joint) in (Q) anterior aspect. Abbreviations: are, arcuate edge; cc, cephalic condyle; cd, condyle; cf, facet for

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Attempting genetic inference from directional asymmetry during convergent hindlimb reduction in squamates

Loss and reduction of paired appendages is common in vertebrate evolution. How often does such convergent evolution depend on similar developmental and genetic pathways? For example, many populations of the Threespine Stickleback and Ninespine Stickleback (Gasterosteidae) have independently evolved pelvic reduction, usually based on independent mutations that caused reduced Pitx1 expression. Reduced Pitx1 expression has also been implicated in pelvic reduction in manatees. Thus, hind limb reduction stemming from reduced Pitx1 expression has arisen independently in groups that diverged tens to hundreds of millions of years ago, suggesting a potential for repeated use of Pitx1 across vertebrates. Notably, hindlimb reduction based on reduction of Pitx1 expression produces left-larger directional asymmetry in the vestiges. We used this phenotypic signature as a genetic proxy, testing for hindlimb directional asymmetry in six genera of squamate reptiles that independently evolved hindlimb reduction and for which genetic and developmental tools are not yet developed: Agamodon anguliceps, Bachia intermedia, Chalcides sepsoides, Indotyphlops braminus, Ophisaurus attenuatuas and O. ventralis, and Teius teyou. Significant asymmetry occurred in one taxon, Chalcides sepsoides, whose left-side pelvis and femur vestiges were 18% and 64% larger than right-side vestiges, respectively, suggesting modification of Pitx1 expression in that species. However, there was either right-larger asymmetry or no directional asymmetry in the other five taxa, suggesting multiple developmental genetic pathways to hindlimb reduction in squamates and vertebrates more generally.

Fig. 7 in An early Eocene pan-gekkotan from France could represent an extra squamate group that survived the K/Pg extinction

Fig. 7. SEM images of the jaw symphyseal region and details of tooth crowns in two extant gekkotans, Gonatodes albogularis (Duméril and Bibron, 1836) A) and Sphaerodactylus klauberi Grant, 1923 (B), showing a similar sulcus on the crown as present in Bauersaurus gen. nov.

Fig. 4 in An early Eocene pan-gekkotan from France could represent an extra squamate group that survived the K/Pg extinction

Fig. 4. Pan-gekkotan squamate Bauersaurus cosensis gen. et sp. nov. from Cos, France, early Eocene (MP 10–11). Tooth details of the holotype (UM- COS-1012) maxilla (A) and UM-COS-1013 dentary (B) in ventromedial (A1), ventral (A2), anteromedial (A3), dorsal (B1), anterior (B2) views (all micro-CT visualizations).

Fig. 6 in An early Eocene pan-gekkotan from France could represent an extra squamate group that survived the K/Pg extinction

Fig. 6. Maxillae of several gekkotans compared with Bauersaurus gen. nov. in lateral (left column) and medial (right column) views; 3D models of gekkotans were mirrored. Top to bottom, Sphaerodactylus grandisquamis spanius Stejneger, 1904, RT 14708 (Sphaerodactylidae), Holodactylus africanus Boettger, 1893, CAS 198932 (Eublepharidae), Paradelma orientalis (Günther, 1876) CAS 77652 (Pygopodide), Underwoodisaurus milii Bory de Saint-Vincent, 1823, CAS 74744 (Carphodactylidae), Pseudothecadactylus australis (Günther, 1877) MCZ Herp R- 35162 (Diplodactylide), Bauersaurus cosensis the holotype UM-COS-1012 (Pan-Gekkota). Bauersaurus cosensis is inferred as sister to Gekkota.

Fig. 1 in An early Eocene pan-gekkotan from France could represent an extra squamate group that survived the K/Pg extinction

Fig. 1. Pan-gekkotan squamate Bauersaurus cosensis gen. et sp. nov. from Cos, France, early Eocene (MP 10–11). Holotype (UM-COS-1012), right maxilla, in lateral (A1), medial (A2), dorsal (A3), ventral (A4), anteromedial (A5), and dorsomedioposterior (A6) views (all micro-CT visualizations).