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Data from: A new tusked cistecephalid dicynodont (Therapsida, Anomodontia) from the upper Permian Upper Madumabisa Mudstone Formation, Luangwa Basin, Zambia
Cistecephalids are among the most distinctive Permian dicynodonts because of their highly derived skulls and postcrania, which indicate a fossorial ecology. Four cistecephalid species have been described from India, South Africa, and Tanzania; a fifth putative species has been reported from the Luangwa Basin of Zambia but never formally described. Here we present a detailed description of the Luangwa Basin cistecephalid, which we name Kembawacela kitchingi gen. et. sp. nov. The most obvious diagnostic character of K. kitchingi is the presence of caniniform tusks in most specimens. Other important characters include a pineal foramen located at the posterior end of the skull roof; an interparietal that has a pair of anterior processes that extend onto the dorsal surface of the skull, flanking the pineal foramen (but otherwise is restricted to the occipital surface); an undivided nuchal crest; and a trough on the ventral surface of the mid-ventral vomerine plate. Phylogenetic analysis reconstructs Kembawacela as a basal cistecephalid and confirms that Cistecephalidae is a well-supported clade. However, relationships within the clade received low branch support. Increased knowledge of cistecephalid diversity shows that they vary in functionally relevant characters, such as degree of inflation of the bony vestibule and the morphology of the scapula and humerus, indicating the need for a more nuanced approach to the relationship between form, function, and ecology in the clade. The highly allopatric distribution of cistecephalid species suggests that they experienced strong interspecific competition for limited resources and had limited dispersal ability, similar to extant subterranean mammals.
TABLE 1 in The sternum and interclavicle of Aelurognathus tigriceps (Broom & Haughton, 1913) (Therapsida: Gorgonopsia), with comments on sternal evolution in therapsids
<p>TABLE 1. — Representative measurements from the associated skeleton of <i>Aelurognathus tigriceps</i> (Broom & Haughton, 1913) (NHCC LB350). All measurements are in millimeters. Symbols: <b>‡</b>, estimated; <b>*</b>, measurement taken between proximal and distal articulation surfaces.</p><table><thead><tr><th>Skeleton</th><th><b>Mesurements</b></th></tr></thead><tbody><tr><th>Skull preorbital length</th><td>167‡</td></tr><tr><th colspan="2">Humerus (left)</th></tr><tr><th>length*</th><td>211</td></tr><tr><th>width across distal epicondyles</th><td>97</td></tr><tr><th>minimum midshaft diameter</th><td>22</td></tr><tr><th colspan="2">Humerus (right)</th></tr><tr><th>length*</th><td>214</td></tr><tr><th>width across distal epicondyles</th><td>112</td></tr><tr><th>minimum midshaft diameter</th><td>23</td></tr><tr><th colspan="2">Femur</th></tr><tr><th>midshaft diameter, transverse</th><td>37</td></tr><tr><th>midshaft diameter, anteroposterior</th><td>22</td></tr><tr><th>head transverse width</th><td>57</td></tr></tbody></table>
Figure 10 in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 10. Specimens tentatively considered juvenile representatives of Digalodon rubidgei. RC 303 in dorsal (a) and right lateral (b) views. RC 306 in dorsal (c) and right lateral (d) views. Scale bars equal 1 cm.
Figure 9. USNM 22941, a in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 9. USNM 22941, a "large emydopoid" from Richmond considered a possible sexually dimorphic female of Digalodon rubidgei in dorsal (a), palatal (b), left lateral (c), and occipital (d) views. Scale bar equals 1 cm.
Figure 6 in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 6. Photographs of B 42, a referred specimen of Digalodon rubidgei, in dorsal (a), palatal (b), right lateral (c), left lateral (d), occipital (e), and anterior (f) views. Scale bar equals 1 cm. Abbreviations: ae, anterior emargination of palatal rim; apr, anterior palatal ridge; co, crista oesophagea; ip, interparietal; lar, lateral anterior palatal ridge; lpf, lateral palatal foramen; pa, parietal; pla, pila antotica; ve, vertebra; vf, vascular foramen.
Figure 11 in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 11. Photographs of "large emydopoid" material from the Graaff-Reinet area referable to Compsodon helmoedi. RC 641, a specimen referable to Compsodon helmoedi in dorsal (a) view. RC 736, a specimen referable to Compsodon helmoedi in dorsal (b) and right lateral (c) views. Scale bars equal 1 cm.
Figure 7 in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 7. Photographs of BP/1/157, a referred specimen of Digalodon rubidgei, in dorsal (a) and left lateral (b) views. Abbreviations: cp, caniniform process; ds, tip of dentary symphysis; nb, nasal boss. Scale bar equals 1 cm.
Figure 3 in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 3. Photograph (a) and interpretive drawing (b) of RC 76, the holotype of Digalodon rubidgei, in ventral view. Gray indicates matrix, hatching indicates damaged bone surface, and cross-hatching indicates plaster. Scale bar equals 1 cm. Abbreviations: apr, anterior palatal ridge; apt, anterior pterygoid ramus; bo, basioccipital; bt, basal tuber; co, crista oesophagea; cp, caniniform process; ec, ectopterygoid; ipv, interpterygoid vacuity; j, jugal; mpr, posterior median palatal ridge; mx, maxilla; op, opisthotic; pl, palatine; pmx, premaxilla; ps, parasphenoid; q, quadrate; qpt, quadrate pterygoid ramus; sq, squamosal; st, stapes; t, tusk; v, vomer; vf, vascular foramen.
Figure 2 in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 2. RC 76, the holotype of Digalodon rubidgei, in left lateral (a, photograph; b, interpretive drawing) and right lateral (c, photograph; d, interpretive drawing) views. Gray indicates matrix, hatching indicates damaged bone surface, and cross-hatching indicates plaster. Scale bar equals 1 cm. Abbreviations: cp, caniniform process; ec, ectopterygoid; f, frontal; j, jugal; la, lacrimal; lcf, lacrimal foramen; mx, maxilla; na, nasal; pa, parietal; pmx, premaxilla; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sq, squamosal; t, tusk; vf, vascular foramen.
Figure 8 in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 8. Photograph of RC 469, a specimen referable to Digalodon rubidgei, in dorsal view. Scale bar equals 1 cm.
Figure 1 in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 1. Photograph (a) and interpretive drawing (b) of RC 76, the holotype of Digalodon rubidgei, in dorsal view. Gray indicates matrix, hatching indicates damaged bone surface, and cross-hatching indicates plaster. Scale bar equals 1 cm. Abbreviations: f, frontal; ip, interparietal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pf, pineal foramen; pmx, premaxilla; po, postorbital; pp, preparietal; pr, prootic; prf, prefrontal; sq, squamosal; ta, tabular.
Figure 5 in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 5. Phylogenetic position of Digalodon rubidgei within Dicynodontia based on the results of the phylogenetic analysis. Eo. = Eodicynodon.
Figure 4 in Redescription of Digalodon rubidgei, an emydopoid dicynodont (Therapsida, Anomodontia) from the Late Permian of South Africa
Figure 4. Photograph (a) and interpretive drawing (b) of RC 76, the holotype of Digalodon rubidgei, in occipital view. Gray indicates matrix, hatching indicates damaged bone surface, and cross-hatching indicates plaster. Scale bar equals 1 cm. Abbreviations: bo, basioccipital; dn, dorsolateral notch in squamosal; eo, exoccipital; fm, foramen magnum; ip, interparietal; op, opisthotic; pa, parietal; pe, paroccipital eminence; ptf, post-temporal fenestra; q, quadrate; so, supraoccipital; sq, squamosal; st, stapes; ta, tabular.
FIG. 5 in Taxonomic revision of therocephalians (Therapsida: Theriodontia) from the Lower Triassic of Antarctica
FIG. 5. (A) AMNH FARB 9548, Baurioidea indet., crushed skull and lower jaw in ventral view, with associated postcrania. Abbreviations: C1, first upper canine; C2, second upper canine; PC1, root of first upper postcanine tooth; PC6?, root of possible sixth upper postcanine tooth. (B) AMNH FARB 9525, Baurioidea indet., partial skull and lower jaw in right lateral view. Abbreviations: c, impression of lower canine; C, root of upper canine; I?, root of possible upper incisor. Numbers refer to apomorphies listed in table 1. Scale bars = 10 mm.
FIG. 3. AMNH FARB 9550 in Taxonomic revision of therocephalians (Therapsida: Theriodontia) from the Lower Triassic of Antarctica
FIG. 3. AMNH FARB 9550, Eutherocephalia indet., partially articulated posterior skeleton in ventral view. Numbers refer to apomorphies listed in table 1. Gray areas indicate impression. Abbreviations: tr. int.: internal trochanter; tr. maj.: trochanter major; vert. l.: lumbar vertebrae; vert. s.: sacral vertebrae. Scale bar = 10 mm.
FIG. 2 in Taxonomic revision of therocephalians (Therapsida: Theriodontia) from the Lower Triassic of Antarctica
FIG. 2. Consensus cladogram summarizing the results of several recent analyses of therocephalian relationships (Hopson and Barghusen, 1986; van den Heever, 1994; Botha et al., 2007; Abdala, 2007; Huttenlocker, 2009; Huttenlocker et al., 2011). Numbers in parentheses indicate apomorphies observable in the studied specimens and listed in table 1.
FIG. 1 in Taxonomic revision of therocephalians (Therapsida: Theriodontia) from the Lower Triassic of Antarctica
FIG. 1. Geology and geography of the Permo-Triassic vertebrate-bearing localities in the vicinity of the Beardmore and Shackleton glaciers, central Transantarctic Mountains. Star indicates position of therocephalian localities (Kitching Ridge, Halfmoon Bluff, Thrinaxodon Col) in the lower Fremouw Formation of Cumulus Hills, Shackleton Glacier region (inset). Gray areas represent exposures of the Buckley Formation; striped areas represent exposures of the Fremouw Formation; hatched areas represent igneous intrusions or metamorphic pre-Devonian basement; dotted areas represent surficial cover. Scale bars are in kilometers. (Map modified from Axsmith et al., 2000 and Collinson et al., 2006.)
FIG. 6. AMNH FARB 9542 in Taxonomic revision of therocephalians (Therapsida: Theriodontia) from the Lower Triassic of Antarctica
FIG. 6. AMNH FARB 9542, cf. Ericiolacerta parva, isolated pterygoid in ventral view. Numbers refer to apomorphies listed in table 1. Abbreviation: cr. v.-int. Pt, intermediate ventral crest of pterygoid. Scale bar = 5 mm.
FIG. 7. AMNH FARB 9542 in Taxonomic revision of therocephalians (Therapsida: Theriodontia) from the Lower Triassic of Antarctica
FIG. 7. AMNH FARB 9542, cf. Ericiolacerta parva, isolated left dentary in lateral view. Numbers refer to apomorphies listed in table 1. Abbreviations: i1, root of first lower incisor; i2, root of second lower incisor; pc1?, possible first lower postcanine tooth; pc6?, possible sixth lower postcanine tooth. Scale bar = 5 mm.
Fig. 3 in A New Nonmammalian Eucynodont (Synapsida: Therapsida) from the Triassic of Northern Gansu Province, China, and its Biostratigraphic and Biogeographic Implications
Fig. 3. Beishanodon youngi, gen. et sp. nov. (holotype PKUP V3007): skull in dorsal (A) ventral (B), and lateral (C) views.
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