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Phymaturus robustus LOBO, BARRASSO, HIBBARD, QUIPILDOR, SLODKI, VALDECANTOS & BASSO, 2021

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Higher TaxaLiolaemidae, Iguania, Sauria, Squamata (lizards)
Subspecies 
Common Names 
SynonymPhymaturus robustus LOBO, BARRASSO, HIBBARD, QUIPILDOR, SLODKI, VALDECANTOS & BASSO 2021
Phymaturus zapalensis — LOBO & QUINTEROS 2005 (part)
Phymaturus zapalensis — LOBO et al. 2012 (part)
Phymaturus sp. 17 — MORANDO et al. 2013 
DistributionArgentina (Neuquén)

Type locality: National Road No 40 km 2306, “Las Coloradas” (39°48′01.2′′S, 70°36′38.1′′W; 946 m), Catán Lil Department, Neuquén Province, Argentina.  
Reproduction 
TypesHolotype. Adult male IBIGEO‐R 5556, collected on 12 November 2016 by T. Hibbard, F. Lobo, M. Quipildor, D. Slodki, and S. Valdecantos.
Paratypes. 3 females and 10 males. IBIGEO‐R 5547, 5553, and 5580 females; IBIGEO‐R 5548–52, 5557–58, and 5579 males. MACN 50903–04 (previously IBIGEO‐R 5554–55) males, all collected with the holotype. 
DiagnosisDiagnosis (Table 2). Phymaturus robustus sp. nov. belongs to the patagonicus group, as it exhibits flat and imbricated superciliary scales, smooth tail scales, and a set of en‐ larged scales projected over the tympanic region. Within the patagonicus group, Phymaturus robustus sp. nov. be‐ longs to the payuniae clade, indicated by a dark lateral (flank) band and the absence of ventral tail pattern with scattered small dark spots. As mentioned above, the payuniae clade is formed by eight described species: P. cacivioi, P. delheyi, P. nevadoi, P. payuniae, P. rahuensis, P. sitesi, and P. zapalensis (with the addition of P. niger sp. nov. de‐ scribed above). Phymaturus robustus sp. nov. can be dis‐ tinguished from these species as follows:
Phymaturus cacivioi: in this species, melanic individuals are common, but they are absent in P. robustus sp. nov. The throat pattern is densely disrupted in P. cacivioi (Fig. 6B), while it is sparse and formed by thick lines in P. robustus sp. nov. (Fig. 6D; Table 2). Most individuals of P. robustus sp. nov. exhibit a conspicuous black line behind the mental scale on the midline (Fig. 6D), a character not shared with P. cacivioi. Dorsal white spots are large (formed by 9–16 scales) in P. robustus sp. nov., but small in P. cacivioi (formed by no more than 6 scales; Table 2). In P. robustus sp. nov., white dorsal spots never form transversal lines like they do in P. cacivioi (Table 2). The dorsal tail pattern of P. robustus sp. nov. is either variegated or absent, while it is ringed in P. cacivioi (Table 2). The ventral pattern of the tail is absent in P. robustus sp. nov., unlike in P. cacivioi. The hemipenes of P. robustus are more curved than those of P. cacivioi (Fig. 8B).
Phymaturus delheyi: Some female P. robustus sp. nov. exhibit precloacal pores, a phenomenon not reported for P. delheyi. There are no dorsal ocelli in P. robustus sp. nov., which are common in females of P. delheyi. The dorsal white spots are large (formed by 9–16 scales) in P. robustus but small in P. delheyi (formed by no more than 6 scales; Table 2). The dorsal tail pattern of P. robustus sp. nov. is variegated or absent, unlike in P. delheyi, in which it is spotted. The belly of males of P. robustus sp. nov. are orange, but yellow in P. delheyi (Table 2). The sulcus spermaticus is open in P. robustus sp. nov. but closed in P. delheyi (Table 2). The sulcus bifurcation in P. robustus sp. nov. hemipenes is distal, unlike in P. delheyi (Table 2).
Phymaturus nevadoi: Dorsal ocelli are absent in P. robustus sp. nov. but common in female P. nevadoi. The throat pattern in P. robustus sp. nov. (Fig. 6D) is sparce, formed by thick lines, whereas P. nevadoi’s throat pattern is scarce, but formed by thin lines (Fig. 6C; Table 2). Also, individuals of P. robustus sp. nov. display a conspicuous black line behind the mental scale on the midline (Fig. 6D), absent in P. nevadoi (Table 2). The dorsal tail pattern of P. robustus sp. nov. is either variegated or absent, different from the spotted pattern of P. nevadoi (Table 2). The ventral pattern of the tail is absent in P. robustus sp. nov., different from that of P. nevadoi. Phymaturus robustus sp. nov. exhibits a higher number of scales around midbody than P. nevadoi (x̅ = 222.2 ± 13.1 and x̅ = 203.7 ± 9.9, respectively). Phymaturus robustus sp. nov. is larger than P. nevadoi (x̅=96.9±4.8mmSVLandx̅=86.5±4.2mmSVL,respectively). The sulcus spermaticus is open in the hemipenes of P. robustus sp. nov. but closed in P. nevadoi. The sulcus bifurcation is distal in P. robustus sp. nov., unlike in P. nevadoi (Table 2).
Phymaturus payuniae: Dorsal ocelli are absent in P. robustus sp. nov. but common in females of P. payuniae. The throat pattern in P. robustus (Fig. 6D) is sparce, formed by thick lines, whereas in P. payuniae it is also sparce, but it is formed by thin lines (Fig. 6C; Table 2). Another difference is that individuals of P. robustus sp. nov. display a conspicuous black line behind the mental scale on the midline (Fig. 6D), absent in P. payuniae. The dorsal tail pattern of P. robustus is either variegated or absent, different from the spotted pattern of P. payuniae (Table 2). The ventral pattern of the tail is absent in P. robustus sp. nov., unlike in P. payuniae (Table 2). The belly of male P. robustus sp. nov. is orange, but it is light gray/light yellow in P. payuniae (Table 2). Phymaturus robustus sp. nov. is larger than P. payuniae(x̅=96.9±4.8mmSVLandx̅=80.3±9.5mmSVL, respectively). The sulcus spermaticus is open in the hemipenis of P. robustus sp. nov. but closed in P. payuniae. The sulcus bifurcation is distal in P. robustus sp. nov., unlike in P. payuniae (Table 2).
Phymaturus rahuensis: Some female P. robustus sp. nov. possess precloacal pores, a phenomenon not reported for P. rahuensis. Dorsal ocelli are absent in P. robustus sp. nov. but common in female P. rahuensis (Table 2). Most individuals of P. robustus sp. nov. show a conspicuous black line behind the mental scale on the midline (Fig. 6D), absent in P. rahuensis. Dorsal white spots are large (formed by 9–16 scales) in P. robustus sp. nov. but small in P. rahuensis (Table 2). In P. robustus sp. nov., white dorsal spots never form transversal lines, like they do in certain P. rahuensis individuals. The dorsal tail pattern of P. robustus sp. nov. is either variegated or absent, different from that of P. rahuensis, which is ringed (Table 2). The belly of male P. robustus sp. nov. is orange, but it is yellow in P. rahuensis. The belly of female P. robustus sp. nov. is pink, while it is gray/yellow in female P. rahuensis (Table 2). Phymaturus robustus sp. nov. has fewer scales around midbody than P. rahuensis (x̅ = 222.2 ± 13.1 and x̅ = 240.4 ± 18.8, respectively) and is larger than P. rahuensis (x̅ = 96.9 ± 4.8 mm SVL and x̅ = 89.9 ± 6.1 mm SVL, respectively). Hemipenes are more curved in P. robustus sp. nov. (Fig. 8B) than in P. rahuensis (in lateral view; Table 2).
Phymaturus sitesi: Some female P. robustus sp. nov. possess precloacal pores, a phenomenon not reported for P. sitesi. The throat pattern in P. robustus sp. nov. (Fig. 6D) is sparce, formed by thick lines, but dense or absent in P. sitesi (Table 2). Both sexes of P. robustus show throat patterns, while individuals of P. sitesi do not. Most individuals of P. robustus sp. nov. exhibit a conspicuous black line behind the mental scale on the midline (Fig. 6D), absent in P. sitesi. Dorsal white spots are large (formed by 9–16 scales) in P. robustus sp. nov. but small in P. sitesi (formed by no more than 6 scales; Table 2). The dorsal
tail pattern of P. robustus sp. nov. is either variegated or absent, while it is spotted in P. sitesi (Table 2). The belly of male P. robustus sp. nov. is orange, but it is yellow in P. sitesi. Phymaturus robustus sp. nov. is larger than P. sitesi (x̅ = 96.9 ± 4.8 mm SVL and x̅ = 82.6 ± 4.2 mm SVL, respectively). The sulcus spermaticus is open in P. robustus sp. nov. but closed in P. sitesi. The sulcus bifurcation in P. robustus sp. nov. hemipenes is distal, unlike P. sitesi (Table 2).
Phymaturus zapalensis: The throat pattern in P. robustus sp. nov. (Fig. 6D) is sparce and formed by thick lines, while in P. zapalensis it is dense/dense disrupted (like in P. niger sp. nov.; Fig. 6A; Table 2). Phymaturus robustus sp. nov. displays a conspicuous black line behind the mental scale on the midline (Fig. 6D), which is absent in P. zapalensis. The dorsal white spots are large (formed by 9–16 scales) in P. robustus sp. nov., but small in P. zapalensis (formed by no more than 6 scales; Table 2). In P. robustus sp. nov., white dorsal spots never form transversal lines like they do in certain individuals of P. zapalensis. Phymaturus robustus sp. nov. is larger than P. zapalensis (x̅ = 96.9 ± 4.8 mm SVL and x̅ = 85.9 ± 8.4 mm SVL, respectively). The hemipenes in P. robustus sp. nov. are more curved than in P. zapalensis (in lateral view; Table 2).
Phymaturus niger sp. nov.: Phymaturus robustus sp. nov. is easily distinguished from its sister taxon Phymaturus niger sp. nov. by several characters, including a wider body shape (abdominal width). Also, the dorsal white spots of P. robustus sp. nov. are large, while they are small in P. niger sp. nov. (Table 2). Phymaturus robustus sp. nov. never presents melanism. The mid‐dorsal region of the trunk exhibits enlarged white spots arranged irregularly in P. robustus sp. nov., while they are small and dispersed in P. niger sp. nov. The dorsal tail pattern is variegated or absent in P. robustus sp. nov., while it is ringed in P. niger sp. nov. (Table 2). Coloration of the abdomen and precloacal area is orange in P. robustus sp. nov. males but yellow in P. niger sp. nov. There are fewer enlarged scales on the anterior margin of the tympanic region in P. robustus sp. nov. (x̅=5.2±1.4)than in P.niger sp.nov.(x̅=6.8±1.3). Hemipenes in P. robustus sp. nov. have a wider sulcus spermaticus than those of P. niger sp. nov., as well as an inconspicuous margin of apical lobes and larger calices, irregularly distributed over its asulcate face (Fig. 8; Table 2).
Description of holotype (Fig. 5). Male. SVL 101.3 mm. Head length 17.5 mm. Head width 17.4 mm. Head height (at parietal) 10.6 mm. Axilla–groin 53.9 mm (53.2% of SVL). Tail length (complete, not regenerated) 131.3 mm (1.30 times SVL). Body moderately wide, trunk width: 42.5 mm (41.9% of SVL). Twenty smooth dorsal head scales. Seven, six, and nine scale organs in postrostrals. Nasal bordered by seven scales, not in contact with rostral. Canthal separated from nasal by two scales. Loreal region flat. Eight enlarged supralabial scales, none contacting subocular. Seven enlarged infralabials. Tympanic region oval (height 4.5 mm; width 2.8 mm) with seven enlarged, flat, keeled, perpendicularly projecting scales on anterior margin. Auricular scale absent. Nine convex, juxtaposed temporals. Distance from tympanic region to posterior ciliary scales commissure 6.4 mm. Rostral undivided. Mental subpentagonal, in contact with four scales. Interparietal bordered by eight scales, larger than postparietals. Supraorbital semicircles inconspicuous. Distinctly enlarged supraoculars absent. Nine imbricate, flat superciliaries, the sixth juxtaposed on both extremes by fifth and seventh. Subocular not fragmented, separated from supralabials by two rows of lorilabials at level of its anterior half and one row at level of posterior half. Nine lorilabials, seventh and ninth contacting subocular. Preocular larger than canthal, separated by one scale. Preocular separated from lorilabial row by one scale. Scales of throat round, flat, juxtaposed. Eighty‐seven gulars between auditory meata. Lateral nuchal folds well developed with granular scales over longitudinal fold. Antehumeral pocket well developed. Eighty‐eight scales between tympanic region and shoulder. Sixty‐two scales between antehumeral fold and shoulder. In ventral view, anterior and posterior gular folds present, anterior margins with three enlarged scales on borders. Enlarged scales in center of chest absent. Dorsal scales round, smooth, juxtaposed. Fifty‐one dorsal scales along midline counted in length equivalent to head length. Scales around midbody 236. Ventral scales larger than dorsal scales. Ventral scales between mental and precloacal pores 205. Eight precloacal pores in undivided row without supernumerary pores. Two conspicuous, enlarged postcloacal scales. Brachial and antebrachial scales smooth, posterior margins round. Supracarpals laminar, round, smooth. Subdigital lamellae of fingers with three keels. Number of subdigital lamellae of fingers (left manus) I: 11; II: 18; III: 23; IV: 24; V: 17. Supradigital lamellae convex, imbricate. Infracarpals and infratarsals with round margins, 2–3 keels. Supracarpals and supratarsals smooth, posterior margins rounded. Subdigital lamellae of toes (left pes) I: 14; II: 18; III: 25; IV: 30; V: 22. Claws moderately long (fourth toe claw 2.5 mm).


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Comment 
EtymologyThe specific name, “robustus” (latin: hard and strong), refers to its body size and shape. It is the largest species of the payuniae clade. 
References
  • Lobo, F. and S. Quinteros 2005. A MORPHOLOGY-BASED PHYLOGENY OF PHYMATURUS (IGUANIA: LIOLAEMIDAE) WITH THE DESCRIPTION OF FOUR NEW SPECIES FROM ARGENTINA. Papéis Avulsos de Zoologia (São Paulo) 45(13):143-177 - get paper here
  • Lobo, F., Barrasso, D. A., Hibbard, T., Quipildor, M., Slodki, D., Valdecantos, S., & Basso, N. G. 2021. Morphological and genetic divergence within the Phymaturus payuniae clade (Iguania: Liolaemidae), with the description of two new species. South American Journal of Herpetology 20 (1): 42-66 - get paper here
  • LOBO, FERNANDO; CRISTIAN ABDALA & SOLEDAD VALDECANTOS 2012. Morphological diversity and phylogenetic relationships within a South-American clade of iguanian lizards (Liolaemidae: Phymaturus). Zootaxa 3315: 1–41 - get paper here
  • Lobo, Fernando; Robert E. Espinoza, Eduardo A. Sanabria, and Lorena B. Quiroga 2012. A New Phymaturus (Iguania: Liolaemidae) from the Southern Extreme of the Argentine Puna. Copeia 2012 (1): 12-22. - get paper here
  • Morando, Mariana; Luciano J. Avila, Cristian H.F. Perez, Monty A. Hawkins, Jack W. Sites 2013. A molecular phylogeny of the lizard genus Phymaturus (Squamata, Liolaemini): Implications for species diversity and historical biogeography of southern South America. Molecular Phylogenetics and Evolution 66 (3): 694-714 - get paper here
 
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