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-Snout morphology of Pelturagonia anolophium (female holotype, MZB 14992) and Phoxophrys tuberculata (UTA 65254). Rostral scales shaded gray.

-Snout morphology of Pelturagonia anolophium (female holotype, MZB 14992) and Phoxophrys tuberculata (UTA 65254). Rostral scales shaded gray.

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Article
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We review morphology and systematics of Phoxophrys using new specimens of previously rare species. In addition to external characters, we relied heavily on skull morphology visualized using computed tomography data of all currently recognized species in this genus. Phylogenetic analysis of ND4, 12S, and 16S mDNA sequences reveal that Ph. tuberculat...

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... and infraorbital series.-Number and condition of scales below the eye have considerable diagnostic value among draconines (e.g., Wood et al. 2009;Mahony 2010;Harvey et al. 2017a,b). Between the orbital margin and supralabials, species of Phoxophrys invariably have an enlarged row of infraorbitals (Fig. 3). The infraorbitals may contact the supralabials or be separated from them by a row of small lorilabials. Like Inger (1960), we use the term infraorbitals for both types of scales and use the term lorilabial when we need to draw a distinction between these two types of scales. Inger's (1960) key uses variation in these scales at two ...
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... are unmodified (A) or tubercular (B) anterior to a large tubercular scale below the rictal fold. Harvey et al. (2017a) recently used counts of tubercular sublabial scales to diagnose species of Dendragama. These same scales have diagnostic value in Phoxophrys. All Phoxophrys have a prominent tubercular scale positioned below the rictal fold (Fig. 3). Most species also have 2-7 similar tubercular scales on sublabials anterior to this subrictal scale. However, Ph. cephalum lacks the additional scales and only has the single subrictal tubercular ...
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... we now know that condition of the dorsal crest is sexually dimorphic in this species. As Inger (1960) observed, females lack a crest or, at least, have a poorly defined one. Males have a continuous, projecting dorsal crest of small triangular scales extending from the pectoral gap to the base of the tail ( Fig. 1; see also Manthey 2010: Fig. RA03453-4, who illustrates an additional male specimen from ''near Solok'' with a clearly visible dorsal crest extending onto the tail). Thus, when identifying male Phoxophrys, this character causes confusion at Couplet 2 of Inger's key, potentially leading some to misidentify Ph. tuberculata as Ph. ...
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... Referring to illustrations provided in their supplementary material, skulls with either state have a high ventral flange that abruptly stops near the quadrate. The pterygoids of Phoxophrys anolophium, Ph. borneensis, Ph. cephalum, and Ph. spiniceps lack the flange or have a low ridge that ends in an inconspicuous, shallow dip near the quadrate (Fig. 13). In a broader analysis of agamids, the new character defined here could be treated as a third character state for Character 14 of Stilson et al. (2017). Among other Draconinae, the pterygoids of only Lophocalotes and Ptycholaemus lack the ventral flange. Pterygoids of Phoxophrys nigrilabris and Ph. tuberculata have well-developed ...
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... Bornean Phoxophrys have large tablike angular processes, with high vertical development in a transverse plane. In these species, the anterior face of the process is concave. In sharp contrast, Ph. tuberculata lacks an angular process (Fig. 13). Neither of these character states is common among other draconines. Among the species we examined, only the four species of Dendragama entirely lack angular processes, whereas only Aphaniotis acutirostris, Pseudocalotes kingdonwardi, and Lyriocephalus scutatus have transverse, vertical processes. The remaining species have horizontal ...
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... maximum-likelihood and Bayesian trees differed only in placement of Calotes versicolor. With a posterior probability of 100%, the Bayesian analysis recovered C. versicolor as sister to all other draconines except for Draco and Bornean Phoxophrys, whereas the maximum-likelihood analysis placed C. versicolor above mainland Pseudocalotes, albeit with a bootstrap value of 65. Figure 13 does not show intergeneric relationships among the various species of Dendragama, Lophocalotes, and Pseudocalotes. With the exception of the three species of Dendragama, Harvey et al. (2017bHarvey et al. ( , 2018) published these intergeneric relationships. ...
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... Lophocalotes, and Pseudocalotes. With the exception of the three species of Dendragama, Harvey et al. (2017bHarvey et al. ( , 2018) published these intergeneric relationships. Interested readers may also consult our supplementary material to view the unedited maximum-likelihood and Bayesian trees used to construct the simplified phylogeny in Fig. 13. Molecular data did not support monophyly of Phoxophrys as defined by Inger (1960). Although we lack DNA sequences for half of the species of Phoxophrys, we found Ph. tuberculata to be more closely related to Dendragama, Lophocalotes, and insular Pseudocalotes than to the Bornean species Ph. anolophium and Ph. nigrilabris (Fig. 14). ...
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... arched supraciliaries producing a serrate edge to the supercilium (1.1) and a bony ridge that curves ventrad from the postciliary ornament to extend vertically down the postorbital bone (18.2). Pelturagonia nigrilabris is the most distinctive member of Pelturagonia and has evolved two to four times as many apomorphic characters as its congeners (Fig. ...
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... of the holotype followed by UTA 21846 then UTA 21844 and MZB 14993 in parentheses: SVL 79.5 (39.2, 44.5, 42.8), body length 38.8 (19.0, 21.0, 19.8), pectoral width 15.4 (7.2, 8.3, 7.7), length of tail 113 (63,66,65), head length 17.9 (10.7, 11.6, 10.9), head width 17.0 (9.3, 10.0, 9.5), diameter of orbit 8.2 (5.4, 5.7, 5.3), width of snout 5.4 (3.4, 3.9, 3.7), width of medial rostral 1.1 (not measured for paratypes), length of shank 17.9 (8.8, 10.8, 9.2), height of longest nuchal crest scale 1.4 (not measured for paratypes), height of longest dorsal crest scale 1.2 (not measured for paratypes), length of Finger III 9 (5.5, 6, 6), length of Finger IV 10 (6.2, 7, 7), length of Toe IV 14 (8.5, ...
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... and infraorbital series.-Number and condition of scales below the eye have considerable diagnostic value among draconines (e.g., Wood et al. 2009;Mahony 2010;Harvey et al. 2017a,b). Between the orbital margin and supralabials, species of Phoxophrys invariably have an enlarged row of infraorbitals (Fig. 3). The infraorbitals may contact the supralabials or be separated from them by a row of small lorilabials. Like Inger (1960), we use the term infraorbitals for both types of scales and use the term lorilabial when we need to draw a distinction between these two types of scales. Inger's (1960) key uses variation in these scales at two ...
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... are unmodified (A) or tubercular (B) anterior to a large tubercular scale below the rictal fold. Harvey et al. (2017a) recently used counts of tubercular sublabial scales to diagnose species of Dendragama. These same scales have diagnostic value in Phoxophrys. All Phoxophrys have a prominent tubercular scale positioned below the rictal fold (Fig. 3). Most species also have 2-7 similar tubercular scales on sublabials anterior to this subrictal scale. However, Ph. cephalum lacks the additional scales and only has the single subrictal tubercular ...
Context 12
... we now know that condition of the dorsal crest is sexually dimorphic in this species. As Inger (1960) observed, females lack a crest or, at least, have a poorly defined one. Males have a continuous, projecting dorsal crest of small triangular scales extending from the pectoral gap to the base of the tail ( Fig. 1; see also Manthey 2010: Fig. RA03453-4, who illustrates an additional male specimen from ''near Solok'' with a clearly visible dorsal crest extending onto the tail). Thus, when identifying male Phoxophrys, this character causes confusion at Couplet 2 of Inger's key, potentially leading some to misidentify Ph. tuberculata as Ph. ...
Context 13
... Referring to illustrations provided in their supplementary material, skulls with either state have a high ventral flange that abruptly stops near the quadrate. The pterygoids of Phoxophrys anolophium, Ph. borneensis, Ph. cephalum, and Ph. spiniceps lack the flange or have a low ridge that ends in an inconspicuous, shallow dip near the quadrate (Fig. 13). In a broader analysis of agamids, the new character defined here could be treated as a third character state for Character 14 of Stilson et al. (2017). Among other Draconinae, the pterygoids of only Lophocalotes and Ptycholaemus lack the ventral flange. Pterygoids of Phoxophrys nigrilabris and Ph. tuberculata have well-developed ...
Context 14
... Bornean Phoxophrys have large tablike angular processes, with high vertical development in a transverse plane. In these species, the anterior face of the process is concave. In sharp contrast, Ph. tuberculata lacks an angular process (Fig. 13). Neither of these character states is common among other draconines. Among the species we examined, only the four species of Dendragama entirely lack angular processes, whereas only Aphaniotis acutirostris, Pseudocalotes kingdonwardi, and Lyriocephalus scutatus have transverse, vertical processes. The remaining species have horizontal ...
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... maximum-likelihood and Bayesian trees differed only in placement of Calotes versicolor. With a posterior probability of 100%, the Bayesian analysis recovered C. versicolor as sister to all other draconines except for Draco and Bornean Phoxophrys, whereas the maximum-likelihood analysis placed C. versicolor above mainland Pseudocalotes, albeit with a bootstrap value of 65. Figure 13 does not show intergeneric relationships among the various species of Dendragama, Lophocalotes, and Pseudocalotes. With the exception of the three species of Dendragama, Harvey et al. (2017bHarvey et al. ( , 2018) published these intergeneric relationships. ...
Context 16
... Lophocalotes, and Pseudocalotes. With the exception of the three species of Dendragama, Harvey et al. (2017bHarvey et al. ( , 2018) published these intergeneric relationships. Interested readers may also consult our supplementary material to view the unedited maximum-likelihood and Bayesian trees used to construct the simplified phylogeny in Fig. 13. Molecular data did not support monophyly of Phoxophrys as defined by Inger (1960). Although we lack DNA sequences for half of the species of Phoxophrys, we found Ph. tuberculata to be more closely related to Dendragama, Lophocalotes, and insular Pseudocalotes than to the Bornean species Ph. anolophium and Ph. nigrilabris (Fig. 14). ...
Context 17
... arched supraciliaries producing a serrate edge to the supercilium (1.1) and a bony ridge that curves ventrad from the postciliary ornament to extend vertically down the postorbital bone (18.2). Pelturagonia nigrilabris is the most distinctive member of Pelturagonia and has evolved two to four times as many apomorphic characters as its congeners (Fig. ...
Context 18
... of the holotype followed by UTA 21846 then UTA 21844 and MZB 14993 in parentheses: SVL 79.5 (39.2, 44.5, 42.8), body length 38.8 (19.0, 21.0, 19.8), pectoral width 15.4 (7.2, 8.3, 7.7), length of tail 113 (63,66,65), head length 17.9 (10.7, 11.6, 10.9), head width 17.0 (9.3, 10.0, 9.5), diameter of orbit 8.2 (5.4, 5.7, 5.3), width of snout 5.4 (3.4, 3.9, 3.7), width of medial rostral 1.1 (not measured for paratypes), length of shank 17.9 (8.8, 10.8, 9.2), height of longest nuchal crest scale 1.4 (not measured for paratypes), height of longest dorsal crest scale 1.2 (not measured for paratypes), length of Finger III 9 (5.5, 6, 6), length of Finger IV 10 (6.2, 7, 7), length of Toe IV 14 (8.5, ...

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