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Figure 7 in Phylogenetic implications of the morphology of the braincase of caecilian amphibians (Gymnophiona)

Figure 7. The morphology of the braincase (sphenethmoid and os basale) and stapes, as revealed through microcomputed tomography, of Epicrionops bicolor (BM 1946.9.5.66). A, the braincase in lateral view; B, dorsal view; and C, ventral view (arrowhead indicates location of the constriction of floor of the os basale, if present). D, the sphenethmoid in anterior view; and E, posterior view. F, the os basale in anterior view; and G, posterior view (asterisk indicates the location of the incision of the margin of the otic capsule by the fenestra vestibuli). H, the right medial surface of the os basale revealing the foramina of the antotic wall and of the medial wall of the otic capsule. I–L, the left stapes in natural position. I, lateral view; J, medial view; K, anterior view; and L, posterior view. Scale bars = 1 cm. See Figure 2 for abbreviations.

Figure 3 in Phylogenetic implications of the morphology of the braincase of caecilian amphibians (Gymnophiona)

Figure 3. The morphology of the braincase (sphenethmoid and os basale) and stapes, as revealed through microcomputed tomography, of Gymnopis multiplicata (BM 1907.10.9.10). A, the braincase in lateral view; B, dorsal view; and C, ventral view (arrowhead indicates location of the constriction of floor of the os basale, if present). D, the sphenethmoid in anterior view; and E, posterior view. F, the os basale in anterior view; and G, posterior view (asterisk indicates the location of the incision of the margin of the otic capsule by the fenestra vestibuli). H, the right medial surface of the os basale revealing the foramina of the antotic wall and of the medial wall of the otic capsule. I–L, the left stapes in natural position. I, lateral view; J, medial view; K, anterior view; and L, posterior view. Scale bars = 1 cm. See Figure 2 for abbreviations.

Figure 9 in Phylogenetic implications of the morphology of the braincase of caecilian amphibians (Gymnophiona)

Figure 9. The morphology of the braincase (sphenethmoid and os basale) and stapes, as revealed through microcomputed tomography, of Siphonops annulatus (UMMZ 150624). A, the braincase in lateral view; B, dorsal view; and C, ventral view (arrowhead indicates location of the constriction of floor of the os basale, if present). D, the sphenethmoid in anterior view; and E, posterior view. F, the os basale in anterior view; and G, posterior view (asterisk indicates the location of the incision of the margin of the otic capsule by the fenestra vestibuli). H, the right medial surface of the os basale revealing the foramina of the antotic wall and of the medial wall of the otic capsule. I–L, the left stapes in natural position. I, lateral view; J, medial view; K, anterior view; and L, posterior view. Scale bars = 1 cm. See Figure 2 for abbreviations.

Figure 8 in Phylogenetic implications of the morphology of the braincase of caecilian amphibians (Gymnophiona)

Figure 8. The morphology of the braincase (sphenethmoid and os basale), as revealed through microcomputed tomography, of Crotaphatrema lamottei (UMMZ 174496). A, the braincase in lateral view; B, dorsal view; and C, ventral view (arrowhead indicates location of the constriction of floor of the os basale, if present). D, the sphenethmoid in anterior view; and E, posterior view. F, the os basale in anterior view; and G, posterior view (asterisk indicates the location of the incision of the margin of the otic capsule by the fenestra vestibuli). H, the right medial surface of the os basale revealing the foramina of the antotic wall and of the medial wall of the otic capsule. Scale bars = 1 cm. See Figure 2 for abbreviations.

Figure 6 in Phylogenetic implications of the morphology of the braincase of caecilian amphibians (Gymnophiona)

Figure 6. The morphology of the braincase (sphenethmoid and os basale) and stapes, as revealed through microcomputed tomography, of Gegeneophis ramaswami (KU 203038). A, the braincase in lateral view; B, dorsal view; and C, ventral view (arrowhead indicates location of the constriction of floor of the os basale, if present). D, the sphenethmoid in anterior view; and E, posterior view. F, the os basale in anterior view; and G, posterior view (asterisk indicates the location of the incision of the margin of the otic capsule by the fenestra vestibuli). H, the right medial surface of the os basale revealing the foramina of the antotic wall and of the medial wall of the otic capsule. I–L, the left stapes in natural position. I, lateral view; J, medial view; K, anterior view; and L, posterior view. Scale bars = 1 cm. See Figure 2 for abbreviations.

Figure 12. The 50 in Phylogenetic implications of the morphology of the braincase of caecilian amphibians (Gymnophiona)

Figure 12. The 50% majority-rule trees resulting from the parsimony (A–C) and 95% maximum clade credibility tree resulting from the Bayesian phylogenetic analyses of morphology (D) including the new braincase and stapes characters. A, all characters, all taxa; B, exclusion of certain eye characters (see text for details), yielding strong congruence with family level systematics; C, exclusion of certain eye characters and Sylvacaecilia grandisonae, which yields improved resolution to the family level with little discrepancy with molecular analyses (bootstrap values greater than 50% shown on the nodes); D, Bayesian analysis excluding certain eye characters and Sy. grandisonae, and the resolution of the nine families under this topology (Dermophiidae, Herpelidae, and Siphonopidae are paraphyletic). Node letters correspond to the text. 1, Rhinatrematidae; 2, Ichthyophiidae; 3, Scolecomorphidae; 4, Herpelidae; 5, Typhlonectidae; 6, Caeciliidae; 7, Indotyphlidae; 8, Siphonopidae; and 9, Dermophiidae.

Figure 11. Residual plots from the regression analyses for the four measurements that had low r in Phylogenetic implications of the morphology of the braincase of caecilian amphibians (Gymnophiona)

Figure 11. Residual plots from the regression analyses for the four measurements that had low r-squared values in their respective regression plots. Species circled represent those that deviate from the expected value of each measurement predicted by the regression plot. Data point labels correspond to the first letter of the genus followed by the first three letters of the species. A, residuals from the regression plot of dorsomedial process length against skull length, for which Typhlonectes natans (T.natans), Chthonerpeton indistinctum (C.indistinctum), and Crotaphatrema lamottei (C.lamottei) possess a process that is shorter than predicted; B, residuals from the regression plot of nasal septum length against skull length, for which T. natans and Ch. indistinctum possess a septum that is shorter than predicted; C, residuals of the regression plot of lateral wall length against skull length, for which Scolecomorphus kirkii (S.kirkii), Scolecomorphus vittatus, and Cr. lamottei possess walls that are longer than predicted; and D, residuals of the regression plot of occipital condyle protrusion against skull length, for which Epicrionops bicolor (E.bicolor), Rhinatrema bivittatum (R.bivittatum), Ichthyophis beddomei (I.beddomei), Ichthyophis kohtaoensis, Caudacaecilia asplenia (C.asplenia), and Scolecomorphus vittatus (S.vittatus) possess less protruded condyles than predicted.

Figure 14 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 14. Fifty-nine major collecting localities of Praomys reported herein, sequentially numbered from north to south. Numbers refer to the localities gazetted in Appendix 1 and to those used to define the 21 OTUs (see Material and Methods).

Figure 13 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 13. Interpretations of area relationship among Tanzanian mountains, as derived from documented occurrences (Appendix 4) of terrestrial small mammals in all montane settings (A, C) or those restricted to montane forest (B, D) and based on two measures of faunal similarity, Sørensen's (A, B) and Raup-Crick's (C, D) indices. Mountain abbreviations: EUS, East Usambara Mts; MAL, Malundwe Mts; NGU, Nguu Mts; NGR, Nguru Mts; NHI, Northern Highlands (Mt Kilimanjaro, Mt Meru, Ngorongoro Crater Highlands); NPA, North Pare Mts; RUB, Rubeho Mts; SHI, Southern Highlands (Livingstone Mts, Poroto Mts, Mt Rungwe); SPA, South Pare Mts; TAI, Taita Hills; UDZ, Udzungwa Mts; UKA, Ukaguru Mts; ULU, Uluguru Mts; WUS, West Usambara Mts.

Figure 12 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 12. Morphometric comparisons of samples of Hylomyscus and Praomys that co-occur in montane forest across the Eastern Arc Mountains (South Pare Mts to Udzungwa Mts) into the Southern Highlands (Livingstone Mts and Mt Rungwe) of Tanzania. Top graphs: north–south transects of morphometric differentiation as indexed by the first canonical variate extracted; vertical lines correspond to the sample range and filled circles to the sample mean (Abbreviations: EUS, East Usambara Mts; LIV, Livingstone Mts; NGR, Nguru Mts; RUN, Mt Rungwe; SPA, South Pare Mts; UDZ, Udzungwa Mts; UKA, Ukaguru Mts; ULU, Uluguru Mts; WUS, West Usambara Mts). Bottom phenograms: Cluster diagrams (UPGMA) based on Mahalanobis squared distances between the nine OTU centroids as derived from discriminant function analysis; only nodes that were consistently recovered by bootstrapping are indicated (± 70%, 1000 iterations), leaving poorly defined levels of phenetic affinity as unresolved. Both multivariate perspectives document abrupt shifts in craniodental differentiation between populations of Praomys coincident with the Makambako Gap (here interpreted as different species, P. melanotus and P. taitae) but not those of Hylomyscus (interpreted as the single species H. arcimontensis by Carleton & Stanley, 2005). See text for discussion.

Figure 11 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 11. Plot of elevational ranges (in metres, above sea level) of Hylomyscus arcimontensis (including localities in the EAM and Southern Highlands) and the three species of the Praomys delectorum species group. Vertical line embraces the elevational range of each species and filled circles indicate elevations of individual collecting localities (see Appendix X).

Figure 9 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 9. Examples of the Praomys delectorum group as photographed in the field: Top, P. melanotus from Ngozi Crater in the Poroto Mountains, Southern Highlands; Bottom, P. taitae from Mount Malundwe, central Eastern Arc Mountains. Darker pigmentation of the epidermal surfaces is typical of individual P. melanotus compared with P. taitae; also note the extension of dark hairs onto the metatarsum as characteristic of P. melanotus versus a wholly white metatarsum and phalanges in most P. taitae. Hindfoot length averages about 23–25 mm; photographs by W. T. Stanley.

Figure 7 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 7. Lateral view (about 2¥) of adult crania of the three species of the Praomys delectorum group (same specimens as illustrated in Fig. 6): A, P. delectorum; B, P. melanotus; C, P. taitae.

Figure 10 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 10. Distributions of the three species of the Praomys delectorum group based on specimens documented herein (see species accounts). The patchy occurrence of Praomys across this region is closely linked to equally patchy associations of wet montane forest (areas shaded pale grey, as extracted from World Wildlife Fund Ecoregions, USGS Digital Atlas of Africa).

Figure 8 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 8. Dorsal (left set) and ventral (right set) views of adult round skins (about 0.5¥), illustrating 'typical' pelage patterns of the three species of the Praomys delectorum group recognized here: AA′, P. delectorum (FMNH 181237; Total Length = 223 mm), a male from Chisongeli Forest, Mt Mulanje, 1575 m, Malawi; BB′, P. melanotus (FMNH 205299; Total Length = 234 mm), a male from Ngozi Crater, Poroto Mts, 2250 m, Tanzania; CC′, P. taitae (FMNH 150346; Total Length = 240 mm), a male from 4.5 km ESE of Amani, East Usambara Mts, 900 m, Tanzania.

Figure 5 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 5. Cluster diagram (UPGMA) based on average Mahalanobis distances between the 21 OTU centroids as derived from discriminant function analysis (coefficient of cophenetic correlation = 0.938; also see Fig. 4). Bootstrap values are indicated for those nodes that were consistently defined (± 70%, 1000 iterations); mountain system and OTU identifier are indicated for each terminal OTU; species recognized herein are identified for those stems that subtend major geographical associations among the analytical samples.

Figure 6 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 6. Dorsal (top row) and ventral (bottom row) views (about 2¥) of adult crania of the three species of the Praomys delectorum group recognized here: A, P. delectorum (FMNH 181214; ONL = 29.0 mm), a male from Lichenya Hut, Mt Mulanje, 1840 m, Malawi; B, P. melanotus (FMNH 163682; ONL = 29.9 mm), a female from 5 km east of Ilolo, Rungwe Forest Reserve, Mt Rungwe, 1870 m, Tanzania; C, P. taitae (FMNH 153979; ONL = 28.7 mm), a male from 3 km east and 0.7 km north of Mhero, Chome Forest Reserve, South Pare Mts, 2000 m, Tanzania.

Figure 4 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 4. Three scatter plots depicting results of discriminant function analysis performed on 18 log-transformed craniodental variables, as measured on 541 intact adult specimens representing 21 OTUs of the Praomys delectorum complex. A, projection of individual specimen scores onto the first two canonical variates (CV) extracted. B, projection of centroids for the 21 OTUs onto the first two canonical variates (see Material and Methods for general locality and sample sizes; bars indicate ± 1 standard error of the mean for a sample centroid); OTU numbers 1–15 are placed approximate to the entire cluster in view of the extensive overlap and visual congestion among those analytical samples. C, multivariate disposition of the four type specimens entered as unknowns and based on a posteriori classification using the discriminant function coefficients; confidence envelopes represent 1 standard deviation around a grand centroid and enclose approximately 68% of specimen scores for each. See Table 4 for variable correlations and variance explained.

Figure 3 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 3. Two scatter plots depicting results of 18 log-transformed craniodental variables as measured on 155 intact specimens representing samples from the Udzungwa Mts (OTU 15) and regions of the type localities of Praomys melanotus (OTUs 16–18) and P. taitae (OTU 1). The factor scores of the type specimens of melanotus and taitae are indicated by small crosses; see Table 3 for variable correlations and variance explained. A, projection of unrotated specimen scores onto the first and second principal components (PC) extracted; major axes of the species constellations and regression statistics are indicated (see text for discussion). Specimens from the Udzungwa Mts, a locality which Allen & Loveridge (1933) had allocated to their new species melanotus, are aligned with examples of P. taitae. B, projection of specimen scores, after varimax rotation, onto the first and second principal components (PC) extracted. Reorientation of major axes of the species constellations maximizes age variation along PC I, whereas taxon differences are consolidated mostly along PC II (see text for discussion).

Figure 1 in Species limits within the Praomys delectorum group (Rodentia: Muridae: Murinae) of East Africa: a morphometric reassessment and biogeographical implications

Figure 1. Mountainous regions of East Africa inhabited by populations of the Praomys delectorum complex, covering an area approximately 2–17°S by 30–41°E. Type localities are illustrated for the four species-group taxa currently assigned to Praomys delectorum: delectorum Thomas, 1910 (Malawi, Mulanje Plateau, 1675 m); melanotus Allen & Loveridge, 1933 (Tanzania, Poroto Mountains, Nyamwanga, 1950 m); octomastis Hatt, 1940 (Tanzania, Old Mbulu Reserve, 1829 m); taitae Heller, 1912 (Kenya, Taita Hills, Mount Mbololo, 1524 m). The Eastern Arc Mountains (sensu Wasser & Lovett 1993) are those disconnected ranges in Tanzania and southern Kenya that trend in a north-eastern direction from the Makambako Gap, including the Udzungwa Mountains through the Taita Hills.