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Crocodylian princess in Taiwan: Revising the taxonomic status of Tomistoma taiwanicus from the Pleistocene of Taiwan and its paleobiogeographic implications
<p><em>Toyotamaphimeia</em>, deriving from a mythological princess in Japan into a crocodile, refers to an extinct crocodylian lineage. Here, we reexamine the holotype of a long-forgotten species: <em>Tomistoma taiwanicus </em>from the Pleistocene of Tainan (Taiwan) and revise its taxonomic status to <em>Toyotamaphimeia taiwanicus </em>comb. nov., leading to the first recognized species of <em>Toyotamaphimeia </em>outside Japan. Our phylogenetic analyses also support this taxonomic assignment and, more interestingly, further suggest an East Asian lineage. In addition, <em>Toyotamaphimeia taiwanicus</em> represents a tropical species, resolving a long-lasting puzzle of why <em>Toyotamaphimeia </em>only inhabited the much higher latitudinal area (Japan). Given the large body size of <em>Toyotamaphimeia taiwanicus</em> (about 7 m) and geologically older than <em>Toyotamaphimeia machikanensis</em> from Japan, we propose a novel evolutionary scenario: the genus <em>Toyotamaphimeia </em>originated in Taiwan and evolved to a large body size with gigantothermic physiology, allowed an Out-of-Taiwan scheme, and dispersed further north to Japan. Our taxonomic identification shows the presence of an extinct endemic crocodylian species from the Pleistocene of Taiwan with large-scale paleogeographic implications, and this study, with our recent progress in vertebrate paleontology in Taiwan, should provoke more in-depth paleontological research on elucidating a larger picture of the Pleistocene extinction.</p>
FIGURE 1 in New taxonomic status of the Pseudotachycines procerus guizhouensis Zhu & Shi 2022 (Rhaphidophoridae: Aemodogryllinae: Aemodogryllini)
FIGURE 1. Pseudotachycines guizhouensis (male). A. habitus, dorsal view; B. habitus, lateral view; C. habitus, ventral view; D. head, frontal view; E. vertex of head, showing conical tubercles; F. hind tarsus, lateral view; G. epiproct, dorsal view; H. paraproct, lateral view.
FIGURE 3 in New taxonomic status of the Pseudotachycines procerus guizhouensis Zhu & Shi 2022 (Rhaphidophoridae: Aemodogryllinae: Aemodogryllini)
FIGURE 3. Pseudotachycines guizhouensis (female). A. habitus, dorsal view; B. habitus, lateral view; C. habitus, ventral view; D. hind tarsus, lateral view; E. ovipositor, lateral view; F. subgenital plate, ventral view.
Figure 1. Phylogenetic relationships between Anthidiellum troodicum, A in Taxonomic status of the disjunct populations of the resin bee Anthidiellum breviusculum (Pérez, 1890) s.l. in the Mediterranean (Apoidea: Anthidiini)
Figure 1. Phylogenetic relationships between Anthidiellum troodicum, A. africanum sp. nov. and A. breviusculum as inferred from COI (mitochondrial cytochrome c oxidase I) DNA sequences. The phylogram shows the best-scoring maximum likelihood tree. Numbers shown at nodes are maximum likelihood bootstrap values based on 1000 bootstrap replicates. A set of 36 COI sequences of Anthidiellum strigatum from different parts of its distribution range was used as outgroup.
Figure 6 in Taxonomic status of the disjunct populations of the resin bee Anthidiellum breviusculum (Pérez, 1890) s.l. in the Mediterranean (Apoidea: Anthidiini)
Figure 6. Distribution of Anthidiellum africanum sp. nov. (green dots), A. breviusculum (blue dots) and A. troodicum (red dots).
Figure 3 in Taxonomic status of the disjunct populations of the resin bee Anthidiellum breviusculum (Pérez, 1890) s.l. in the Mediterranean (Apoidea: Anthidiini)
Figure 3. Face of the female of (a) Anthidiellum breviusculum from Spain, (b) A. africanum sp. nov. from Algeria, and (c) A. troodicum from Turkey and (d) from Israel. Note the finer punctation of the clypeus in A. breviusculum (a) and A. africanum sp. nov. (b), and the coarser punctation in A. troodicum (c and d).
Figure 2 in Taxonomic status of the disjunct populations of the resin bee Anthidiellum breviusculum (Pérez, 1890) s.l. in the Mediterranean (Apoidea: Anthidiini)
Figure 2. Anthidiellum africanum sp. nov. (a) Female (holotyope) from Algeria. (b) Male (paratype) from Morocco.
Figure 5 in Taxonomic status of the disjunct populations of the resin bee Anthidiellum breviusculum (Pérez, 1890) s.l. in the Mediterranean (Apoidea: Anthidiini)
Figure 5. Variability of the punctation and colouration of the scutum in Anthidiellum troodicum. While the scutum is mostly densely punctured (a, male from western Turkey), scattered punctation is sometimes observed, particularly in some southern populations of the Levant b, male from Jordan). It is not yet understood whether this difference has taxonomic relevance.
Figure 4 in Taxonomic status of the disjunct populations of the resin bee Anthidiellum breviusculum (Pérez, 1890) s.l. in the Mediterranean (Apoidea: Anthidiini)
Figure 4. Proboscis of (a) Anthidiellum africanum sp. nov. from Algeria, (b) A. breviusculum from France and (c) A. troodicum from Turkey. The arrows show the absence/presence of hairs with hooked apices on the labial palpi.
FIGURE 9 in Taxonomic status of the enigmatic Natrix sexcarinata Wagler, 1824 (Serpentes: Colubridae: Colubrinae)
FIGURE 9. Aulcate (A) and asulcate (B) sides of the hemipenis of the Phrynonax sexcarinatus (MNRJ 18023) specimen from the Carajás National Forest, Parauapebas municipality, in the state of Pará, Brazil. Scale = 1 cm.
FIGURE 7 in Taxonomic status of the enigmatic Natrix sexcarinata Wagler, 1824 (Serpentes: Colubridae: Colubrinae)
FIGURE 7. Cumulative frequency for the number of anterior keeled dorsal scale rows (top; only for females), number of right supralabials contacting the orbit (center), and number of posterior keeled dorsal scale rows (bottom; only for females) for Phrynonax polylepis subpopulations.
FIGURE 5 in Taxonomic status of the enigmatic Natrix sexcarinata Wagler, 1824 (Serpentes: Colubridae: Colubrinae)
FIGURE 5. Association between snout-vent length and color pattern changes in females (left) and males (right) (A) and differences in the number of right infralabials (left), left infralabials (center) and snout-vent lengths (right) (B) in Phrynonax polylepis from the Amazon.
FIGURE 4 in Taxonomic status of the enigmatic Natrix sexcarinata Wagler, 1824 (Serpentes: Colubridae: Colubrinae)
FIGURE 4. Phrynonax polylepis distribution based on examined samples. Different colors represent subpopulations labeled as distinct Operational Taxonomic Unities, following possible natural dispersion barriers across the main interfluves along the Amazon Basin (see Material and Methods for additional explanations).
FIGURE 1 in Taxonomic status of the enigmatic Natrix sexcarinata Wagler, 1824 (Serpentes: Colubridae: Colubrinae)
FIGURE 1. Illustration employed in the description of Natrix sexcarinata Wagler, 1824 based on a specimen from the banks of the Amazon River in Brazil, modified from the original.
FIGURE 2 in Taxonomic status of the enigmatic Natrix sexcarinata Wagler, 1824 (Serpentes: Colubridae: Colubrinae)
FIGURE 2. Illustration employed in the description of Natrix cinnamomea Wagler, 1824 based on a specimen from the Amazon Forest without detailed location, modified from the original.
FIGURE 8 in Taxonomic status of the enigmatic Natrix sexcarinata Wagler, 1824 (Serpentes: Colubridae: Colubrinae)
FIGURE 8. Dorsal (A), ventral (B) and lateral (C) views of the head and dorsal (D) and ventral (E) views of the body of the Phrynonax sexcarinatus (MNRJ 20302) neotype from Porto Trombetas, Oriximiná municipality, in the state of Pará, Brazil. Specimens measured 1128 mm snout-vent length and 446 mm tail length.
FIGURE 3 in Taxonomic status of the enigmatic Natrix sexcarinata Wagler, 1824 (Serpentes: Colubridae: Colubrinae)
FIGURE 3. General view of Phrynonax polylepis in life from Parauapebas, in the state of Pará (A); Sinop, in the state of Mato Grosso (B); from Bujari, in the state of Acre (C); and Belterra, in the state of Pará (D), all in Brazil. Color Pattern A is depicted in Figures A–B, while Color Pattern B is depicted in figures C–D. Modified images from original photographs by Fábio Giordano (A), Antonino Gonçalves Medina (B), Martin Acosta (C), and Miguel A. Casado (D). All photos are available at .
FIGURE 3 in Taxonomic status of Pseudophilautus conniffae (Anura: Rhacophoridae) an endemic shrub frog restricted to South-Western wet zone of Sri Lanka
FIGURE 3. PC1 vs. PC2 factor scores of the principal components analysis of P. conniffae, P. limbus, P. stictomerus and P. folicola, show P. conniffae and P. limbus clustered together and two species, P. stictomerus and P. folicola, separated well from each other in PC space.
FIGURE 2 in Taxonomic status of Pseudophilautus conniffae (Anura: Rhacophoridae) an endemic shrub frog restricted to South-Western wet zone of Sri Lanka
FIGURE 2. Maximum likelihood phylogram based on mitochondrial 16 rRNA showing the phylogenetic position of P. conniffae. Numbers above branches indicate bootstrap support values on right (ML BS ≥ 75) and Bayesian posterior probabilities on left (BI PP ≥ 0.95), respectively; low support values (ML BS <75, BI PP <0.95) are denoted by "-". Recovered species delimitation based on overall significance of results for molecular species delimitation methods ABGD and mPTP using 16S rRNA are shown as grey color rectangles on the right.
FIGURE 1. A in Taxonomic status of Pseudophilautus conniffae (Anura: Rhacophoridae) an endemic shrub frog restricted to South-Western wet zone of Sri Lanka
FIGURE 1. A. Map showing the sampling localities of P. conniffae; B-D. dorsolateral view of the three specimens in life, JPF 41, JPF 64, JPF 65; E. Habitat of P. conniffae.
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Allen Brain Atlas
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DANDI Archive for NWB datasets
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International Brain Laboratory public data
The International Brain Laboratory public data releases expose standardized mouse decision-making experiments, including Neuropixels recordings, widefield calcium imaging, behavior, and session metadata accessed through the ONE API.
OpenNeuro
OpenNeuro is a free, open platform for sharing neuroimaging datasets, with public search, dataset pages, and download paths for web, S3, DataLad, and the OpenNeuro CLI.