Table 1.
Specimen data and measurements.
Figure 1.
Anterior is to the left; scale bar = 0.5 mm. A–C from U. woodmasoni (TMM M-10001); D–F from U. melanogaster (TMM M-10045); and G–I from B. rhodogaster (TMM M-10024). A,D,G in dorsal view; B,E,H in ventral view; and C,F,I in lateral view. The posterior tip of the nasal process is broken in the B. rhodogaster specimen. l.f = lateral foramen; Na.Pmx = nasal process (keel) of premaxilla; pl.c = posterolateral canal; ros = rostral tip; sep.c = septal canal; sub.f = subnarial foramen; sub.fen = subnarial fenestra; trv.p = transverse process; vp.f = ventral premaxillary foramen; Vo.Pmx = vomerine process of premaxilla.
Figure 2.
Left lateral view of articulated Uropeltis and Brachyophidium skulls.
Anterior up; scale bars = 1.0 mm. (A) U. woodmasoni, TMM M-10006; (B) U. rubromaculata, TMM M-10028; (C) B. rhodogaster, TMM M-10011. a.m.f. = anterior maxillary foramen; Ang = angular; com = compound; den = dentary; ecpt = ectopterygoid; fr = frontal; js.r = juxtastapedial recess; lc.f = lacrimal foramen; ls = laterosphenoid region; m.m.f = middle maxillary foramen; max = maxilla; Me.c.f = foramen associated with canal for Meckel's cartilage; na = nasal; o.f = optic foramen; oo.c = otooccipital complex; p.m.f = posterior maxillary foramen; pa = parietal; pfr = prefrontal; pl = palatine; pmx = premaxilla; pro.c = prootic canal; pt = pterygoid; smx = septomaxilla; spl = splenial; q = quadrate; v2.f = foramen for maxillary branch of trigeminal nerve; v2m.f = foramen for branch of maxillary branch of trigeminal nerve; v3.f = foramen for mandibular branch of trigeminal nerve.
Figure 3.
Anterior is to the left unless noted; scale bar = 0.5 mm. All elements from the left side of the skull. A–C from U. woodmasoni (TMM M-10001); D–F from U. melanogaster (TMM M-10045); and G–I from B. rhodogaster (TMM M-10022). A,D,G in lateral view; B,E,H in medial view (anterior is to the right); and C,F,I in dorsal view. a.m.f = anterior maxillary foramen; alv.c = alveolar canal; ant.med.p = anteromedial process; asc.Mx = ascending process; Ect.Mx = ectopterygoid process of maxilla; f.jug+X = foramen for jugular vein and vagus nerve; m.m.f = middle maxillary foramen; p.m.f = posterior maxillary foramen; Pl.Mx = palatine process (posteromedial process) of maxilla; Pmx.Mx = premaxillary process of maxilla; shelf = shelf medial to articulation facet for prefrontal.
Figure 4.
Ventral view of articulated Uropeltis and Brachyophidium skulls.
Anterior up; scale bars = 1.0 mm. (A) U. woodmasoni, TMM M-10006; (B) U. rubromaculata, TMM M-10028; (C) B. rhodogaster, TMM M-10011. Ang = angular; com = compound; den = dentary; ecpt = ectopterygoid; js.r = juxtastapedial recess; max = maxilla; oo.c = otooccipital complex; pa = parietal; pl = palatine; pmx = premaxilla; pro.c = prootic canal; pt = pterygoid; smx = septomaxilla; spl = splenial; sub.f = subnarial foramen; q = quadrate; vn.o = vomeronasal opening; vo = vomer; vp.f = ventral premaxillary foramen; v2.f = foramen for maxillary branch of trigeminal nerve; v3.f = foramen for mandibular branch of trigeminal nerve.
Figure 5.
Dorsal view of articulated Uropeltis and Brachyophidium skulls.
Anterior up; scale bars = 1.0 mm. (A) U. woodmasoni, TMM M-10006; (B) U. rubromaculata, TMM M-10028 (C) B. rhodogaster, TMM M-10011. Ecpt = ectopterygoid; fr = frontal; low.j = lower jaw; max = maxilla; na = nasal; oo.c = otooccipital complex; pa = parietal; pfr = prefrontal; pmx = premaxilla; pl = palatine; pt = pterygoid; smx = septomaxilla; q = quadrate; R.c = Rieppel's canal; v2m.f = foramen for branch of maxillary branch of trigeminal nerve.
Figure 6.
Dorsal view of articulated Rhinophis skulls.
Anterior up; scale bars = 1.0 mm. (A) R. blythii, TMM M-10030; (B) R. drummondhayi, TMM M-10046; (C) R. philippinus, TMM M-10037; (D) R. homolepis, TMM M-10041. Ecpt = ectopterygoid; fr = frontal; low.j = lower jaw; max = maxilla; na = nasal; oo.c = otooccipital complex; pa = parietal; pfr = prefrontal; pl = palatine; pmx = premaxilla; pt = pterygoid; smx = septomaxilla; q = quadrate; R.c = Rieppel's canal; v2m.f = foramen for branch of maxillary branch of trigeminal nerve.
Figure 7.
Ventral view of articulated Rhinophis skulls.
Anterior up; scale bars = 1.0 mm. (A) R. blythii, TMM M-10030; (B) R. drummondhayi, TMM M-10046; (C) R. philippinus, TMM M-10037; (D) R. homolepis, TMM M-10041. Ang = angular; com = compound; den = dentary; ecpt = ectopterygoid; fr = frontal; js.r = juxtastapedial recess; max = maxilla; oo.c = otooccipital complex; pa = parietal; pfr = prefrontal; pl = palatine; pmx = premaxilla; pro.c = prootic canal; pt = pterygoid; smx = septomaxilla; spl = splenial; sub.fen = subnarial fenestra; q = quadrate; vn.o = vomeronasal opening; vo = vomer; vp.f = ventral premaxillary foramen; v2.f = foramen for maxillary branch of trigeminal nerve; v3.f = foramen for mandibular branch of trigeminal nerve.
Figure 8.
Left lateral view of articulated Rhinophis skulls.
Anterior up; scale bars = 1.0 mm. (A) R. blythii, TMM M-10030; (B) R. drummondhayi, TMM M-10046; (C) R. philippinus, TMM M-10037; (D) R. homolepis, TMM M-10041. a.m.f. = anterior maxillary foramen; com = compound; den = dentary; ecpt = ectopterygoid; fr = frontal; js.r = juxtastapedial recess; lc.f = lacrimal foramen; ls = laterosphenoid region; m.m.f = middle maxillary foramen; max = maxilla; Me.c.f = foramen associated with canal for Meckel's cartilage; na = nasal; o.f = optic foramen; oo.c = otooccipital complex; p.m.f = posterior maxillary foramen; pa = parietal; pfr = prefrontal; pl = palatine; pmx = premaxilla; pro.c = prootic canal; pt = pterygoid; smx = septomaxilla; q = quadrate; v2.f = foramen for maxillary branch of trigeminal nerve; v2m.f = foramen for branch of maxillary branch of trigeminal nerve; v3.f = foramen for mandibular branch of trigeminal nerve.
Figure 9.
Lack of contact between prefrontal and supraorbital process of parietal of Uropeltis woodmasoni (TMM M-10006).
Reconstruction from CT scan, left lateral view, anterior to the left. pfr = prefrontal; s.o.p = supraorbital process of the parietal.
Figure 10.
Anterior is to the right in D,H–K; anterior is to the left in all others; scale bar = 0.5 mm. A–D from the left side of U. woodmasoni (TMM M-10001); E–H from the left side of U. melanogaster (TMM M-10045); and I–L from the right side of B. rhodogaster (TMM M-10022). A,E,I in lateral view; B,F,J in lateral view; C,G,K in ventral view; and D,H,L in medial view. Arrow denotes broken posteromedial portion of bone in U. melanogaster specimen. ad.p = anterodorsal process of lateral process; av.p = anteroventral process of lateral process; lat.p = lateral process (lateral wall); na.bt = nasal buttress; pal.t = palatal tubercle; pm.p = posteromedial process; Pmx.p = premaxillary process of septomaxilla; vm.t = ventromedial tubercle; vn.f = vomeronasal foramen.
Figure 11.
Magnified view of the palate of U. woodmasoni (TMM M-10010) with disarticulated left maxilla and palatine in ventral view.
Scale bars = 0.5 mm. (A) Partial disarticulation reveals contact of the septomaxilla (arrow) with the medial surface of the prefrontal. Note broken palatine process of vomer on left side of animal. Anterior is toward the upper left corner. (B) The groove for the cartilaginous portion of the crista trabecularis is formed between the sphenoid region of the otooccipital complex and the frontal. Arrow points to the ventrolateral part of the frontal-parietal suture. Anterior is to the left. c.trab = ossified base of crista trabecularis; fr = frontal; Fr.Pfr = frontal process of prefrontal; groove = groove for the cartilaginous portion of the crista trabecularis; mx = maxilla; pa = parietal; pfr = prefrontal; pl = palatine; pt = pterygoid; smx = septomaxilla; s.o.p = supraorbital process of the parietal; vo = vomer.
Figure 12.
Anterior is to the left unless noted; scale bar = 0.5 mm. All elements from the left side of the skull. A–C from U. woodmasoni (TMM M-10001); D–F from U. melanogaster (TMM M-10045); and G–I from B. rhodogaster (TMM M-10022). A,D,G in dorsal view; B,E,H in lateral view; and C,F,I in medial view (anterior is to the right). can = anterior opening of the canal within the medial process of the nasal; md.p = medial process; Pmx.Na = premaxillary process of nasal; shelf = shelf that is continuous with pre-orbital ridge of frontal; vl.p = ventrolateral process at triple junction.
Figure 13.
Anterior is to the left in A,D,E; anterior is to the right in B,C,F; scale bar = 0.5 mm. A–C from the left side of the skull in U. woodmasoni (TMM M-10001); D–F from the right side of the skull in U. melanogaster (TMM M-10045); and G–I from the left side of the skull in B. rhodogaster (TMM M-10027). A,C,E in lateral view and B,D,F in medial view. Fr.Pfr = frontal process of prefrontal; knob = medially projecting knob at base of frontal process of prefrontal; l.f.p = lateral foot plate; m.f.p = medial foot process.
Figure 14.
Anterior is to the left unless noted; scale bar = 0.5 mm. All elements from the left side of the skull. A–C from U. woodmasoni (TMM M-10001); D–F from U. melanogaster (TMM M-10045); and G–I from B. rhodogaster (TMM M-10022). A,D,G in dorsal view; B,E,H in lateral view; and C,F,I in medial view (anterior is to the right). Fr.c = frontal canal; l.f.f = lateral frontal flange; m.f.f = mesial frontal flange; o.f = optic foramen; ol.Fr. = olfactory process of frontal; p.o.r = pre-orbital ridge of frontal; So.g = groove for supraorbital process of parietal; trab.g = groove for cartilaginous portion of crista trabecularis.
Figure 15.
Anterior is to the left; scale bar = 0.5 mm. A–C from U. woodmasoni (TMM M-10001); D–F from U. melanogaster (TMM M-10045); and G–I from B. rhodogaster (TMM M-10022). A,D,G in dorsal view; B,E,H in ventral view; and C,F,I in lateral view. Arrow points to slit-like opening along posterior midline of parietal. Pa.sh = parietal shelf; s.o.p = supraorbital process of the parietal; tab = tab-like process that articulates with otic region; V2.f = notch that contributes to V2 foramen.
Figure 16.
Anterior is to the left in A–F; anterior is to the right in I; scale bar = 0.5 mm. A–C from the right side of U. woodmasoni (TMM M-10001); D–F from the right side of U. melanogaster (TMM M-10045); and G–I from the left side of B. rhodogaster (TMM M-10022). A,D,G in dorsal view; B,E,H in ventral view; and C,F,I in medial view. al.p = anterior lateral process; d.Pmx.Vo = dorsal premaxillary process of vomer; dm.f = dorsomedial foramen of posterolateral crest; md.w = medial wall; pl.cr = crest on posterolateral process; pl.p = posterior lateral process; Pl.Vo = palatine process of vomer; Pmx.Vo = premaxillary process of vomer; tab = bone tab projecting into vomero-nasal opening; v.Pmx.Vo = ventral premaxillary process of vomer.
Figure 17.
Magnified view of the palate of Uropeltis.
Anterior is to the left; scale bars = 1.0 mm. (A) U. woodmasoni, TMM M-10010. Note missing max, pl, ecpt, and pt on left side. (B) U. rubromaculata, TMM M-10028. Note that right lower jaw is present. ang = angular; den = dentary; ecpt = ectopterygoid; fr = frontal; max = maxilla; pal.tub = palatine tubercle of septomaxilla; pfr = prefrontal; pl = palatine; pmx = premaxilla; pt = pterygoid; smx = septomaxilla; spl = splenial; sph = sphenoid region of the otooccipital complex; vo = vomer.
Figure 18.
Magnified view of the palate of Rhinophis.
Anterior is to the left; scale bars = 1.0 mm. (A) R. blythi. Note that the left lower jaw is present. (B) R. drummondhayi. ang = angular; den = dentary; ecpt = ectopterygoid; max = maxilla; pl = palatine; pmx = premaxilla; pt = pterygoid; smx = septomaxilla; spl = splenial; sph = sphenoid region of the otooccipital complex; vo = vomer.
Figure 19.
Magnified view of the palate of Rhinophis.
Anterior is to the left; scale bars = 1.0 mm. (A) R. philippinus. Note missing max, ecpt, and pt on right side; (B) R. homolepis. Note that right com and ang are present. ang = angular; ecpt = ectopterygoid; max = maxilla; pfr = prefrontal; pl = palatine; pmx = premaxilla; pt = pterygoid; smx = septomaxilla; sph = sphenoid region of the otooccipital complex; vo = vomer.
Figure 20.
Magnified view of the palate of Brachyophidium rhodogaster.
Anterior is to the left; scale bar = 1.0 mm. Note that posterior half of skull, including otooccipital region, left ecpt, and left pt, is missing. ecpt = ectopterygoid; fr = frontal; max = maxilla; pl = palatine; pmx = premaxilla; pt = pterygoid; vp.f = ventral premaxillary foramen; vn.o = vomeronasal opening; vo = vomer.
Figure 21.
Anterior is to the left in A–F; anterior is to the right in I; scale bar = 0.5 mm. A–C from the right side of U. woodmasoni (TMM M-10001); D–F from the right side of the skull of U. melanogaster (TMM M-10045); and G–I from the left side of the skull of B. rhodogaster (TMM M-10022). A,D,G in dorsal view; B,E,H in ventral view; and C,F,I in medial view. ch.Pl = choanal process of palatine; lat.f = tiny lateral foramen; lat.p = lateral process of palatine; Pt.Pl = pterygoid process of palatine; Vo.Pl = vomerine process of palatine; v2m.f = foramen for branch of the trigeminal nerve (CN V2).
Figure 22.
Disarticulated ectopterygoids and pterygoids.
Anterior is to the right unless noted; scale bars = 0.5 mm. A,E from U. woodmasoni (TMM M-10001); B,F from U. melanogaster (TMM M-10045); C,D from B. rhodogaster (TMM M-10016); and G from B. rhodogaster (TMM M-10022). Right ectopterygoids in ventral (A–C) and dorsal (D) views; left pterygoids (E,G; anterior to the left) and right pterygoid (F) in dorsal views. Ect.Pt = ectopterygoid process of pterygoid; Mx.Ect = maxillary process of ectopterygoid; Pl.Pt = palatine process of pterygoid; post.p = posterior process of pterygoid; Pt.Ect = pterygoid process of ectopterygoid.
Figure 23.
Disarticulated otooccipital complex of Uropeltis woodmasoni (TMM M-10021).
Anterior up unless noted; scale bar = 1.0 mm. (A) Dorsal, (B) ventral, and (C) lateral (anterior to the left) views, note broken i.ch.p. a.sc = anterior semi-circular canal; ap.l = apertura lateralis recessus scalae tympani; c.cer = cerebral carotid foramen; c.trab = ossified crista trabecularis; clt.p = cultriform process; f.end = endolymphatic foramen; f.jug = jugular foramen; i.ch.p = interchoanal process of sphenoid; js.r = juxtastapedial recess; l.sc = lateral semi-circular canal; ls.f = laterosphenoid foramen; o.c = occipital condyle; p.ao.vc = primary anterior opening of vidian canal; p.sc = posterior semi-circular canal; po.vc = posterior opening of vidian canal; pro.c = prootic canal; R.c = Rieppel's canal; s.ao.vc = secondary anterior opening of vidian canal; so.t = spheno-occipital tubercle?; V2 = foramen for maxillary branch of trigeminal nerve; V3 foramen for mandibular branch of trigeminal nerve; VI = foramen for abducens nerve; VII = foramen for facial nerve seven; X = vagus nerve.
Figure 24.
CT images of the otooccipital complex of Uropeltis woodmasoni (TMM M-10006).
Scale bar = 1.0 mm. (A) Anterior view; (B) Anterior cut-away at level of otic capsule; (C) Lateral view, anterior to left. a.sc = anterior semi-circular canal; ap.m = apertura medialis recessus scalae tympani; c.cer = cerebral carotid foramen; c.trab = ossified crista trabecularis; clt.p = cultriform process; f.end = endolymphatic foramen; f.jug = jugular foramen; i.a.m = internal auditory meatus; i.ch.p = interchoanal process of sphenoid; l.sc = lateral semi-circular canal; o.c = occipital condyle; p.ao.vc = primary anterior opening of vidian canal; po.vc = posterior opening of vidian canal; pro.c = prootic canal; s.ao.vc = secondary anterior opening of vidian canal; s.cir = semi-circular canal; stat = statolithic mass; t.f.c = trigeminal facialis chamber (for trigeminal branches two and three); VI = abducens nerve; VII = facial nerve seven (to prootic opening); VIIIv = vestibular branch of auditory nerve; X = vagus nerve; XII = hypoglossal nerve.
Figure 25.
Disarticulated quadrates and stapes.
Anterior is to the left unless noted; scale bars = 0.5 mm. A from U. woodmasoni (TMM M-10001); B from U. melanogaster (TMM M-10045); C from B. rhodogaster (TMM M-10027); and D–F from CT scans of U. woodmasoni (TMM M-10006). Left quadrates in lateral view (A–C), and right stapes in ventrolateral (K, anterior to the right), dorsal (L, anterior to the right), and anterolateral (M, lateral to the left) views. fp.St = stapedial footplate; m.con = mandibular condyle of quadrate; sh.St = stapedial shaft; sst.p = suprastapedial process (caudal process) of quadrate; ty.cr = tympanic crest of quadrate.
Figure 26.
Magnified view of the posterior end of the crista circumfenestralis of Uropeltis woodmasoni (TMM M-10008) in dorsolateral view.
Anterior is to the right; scale bar = 0.5 mm. This specimen has a double opening for the passage of cranial nerve X, the jugular vein, and associated tissue. a.sc = anterior semi-circular canal; ap.l = apertura lateralis recessus scalae tympani; com = compound bone; f.jug = jugular foramen; l.sc = lateral semi-circular canal; o.c.n = neck of occipital condyle; p.sc = posterior semi-circular canal; sst.q = suprastapedial process of the quadrate; X = foramen for vagus nerve.
Figure 27.
Otooccipital complex of Brachyophidium rhodogaster (TMM M-10023).
Anterior up unless noted; scale bar = 1.0 mm. (A) Dorsal, (B) ventral, and (C) lateral (anterior to left) views. a.sc = anterior semi-circular canal; c.cer = cerebral carotid foramen; c.trab = ossified crista trabecularis; clt.p = cultriform process; f.end = endolymphatic foramen; f.jug = jugular foramen; js.r = juxtastapedial recess; l.sc = lateral semi-circular canal; ls.f = laterosphenoid foramen; o.c = occipital condyle; p.ao.vc = primary anterior opening of vidian canal; p.sc = posterior semi-circular canal; po.vc = posterior opening of vidian canal; pro.c = prootic canal; R.c = Rieppel's canal; s.ao.vc = secondary anterior opening of vidian canal; s.cir = semi-circular canal; so.t = spheno-occipital tubercle?; V2 = foramen for maxillary branch of trigeminal nerve; V3 foramen for mandibular branch of trigeminal nerve; VI = foramen for abducens nerve; VII = foramen for facial nerve seven; X = foramen for vagus nerve.
Figure 28.
Disarticulated compounds and dentaries.
Anterior is to the left in A,C,E,G,I,K,L; anterior is to the right in B,D,F,H,J,K; scale bars = 0.5 mm. Left compound bones in lateral (A,C,E) and medial (B,D,F) views. Left dentaries in lateral (G,I) and medial (H,J) views, and right dentary in lateral (K) and medial (L) views. A,B and G,H are from U. woodmasoni (TMM M-10001); C,D and I,J are from U. melanogaster (TMM M-10045); and E,F and K,L are from B. rhodogaster (TMM M-10022). ch.t.f = chorda tympani foramen; Com.Den = compound process of the dentary; cor.p = coronoid process of the compound; ct.f = foramen for the chorda tympani; Den.Com = dentary process of the compound; Me.c = Meckel's canal; Me.g = Meckel's groove; mn.f = mandibular foramen; r.a.p = retroarticular process of the compound.
Figure 29.
Majority Rule Consensus trees (50%) for the phylogenetic analysis of morphological characters for 16 uropeltid species.
(A) All 33 characters included (see Methods S2); (B) characters 4,6,7,11,13 excluded.
Figure 30.
Strict Consensus trees with Bremer support values for the phylogenetic analysis of morphological characters for 16 uropeltid species.
(A) All 33 characters included (see Methods S2); (B) characters 4,6,7,11,13 excluded.