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Pseudocalotes rhammanotus HARVEY, HAMIDY, KURNIAWAN, SHANEY & SMITH, 2014

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Higher TaxaAgamidae (Draconinae), Sauria, Iguania, Squamata (lizards)
Common NamesE: Stitched-back False Garden Lizard 
SynonymPseudocalotes rhammanotus HARVEY, HAMIDY, KURNIAWAN, SHANEY & SMITH 2014 
DistributionIndonesia (S Sumatra)

Type locality: montane forest along the ridge of a mountain south of Danau Ranau (= Lake Ranau), Lampung, Sumatra, Indonesia, 4.9394° S, 103.85292° E, 1237 m elevation.  
Reproductionoviparous. Each oviduct of the holotype contains a single large, oval egg with a thick fibrous shell that has not yet calcified (dimensions of egg from left oviduct 17.3 X 6.8 mm; clutch size two). The left ovary contains one large and subcircular yellow egg (largest diameter 4.8 mm) and four small cream-colored eggs (0.7–1.5 mm). 
TypesHolotype: MZB 10804, an adult female, 1954 hrs, 17 June 2013, found by Elijah Wostl, accompanied by other field party members from BC, UB, and UTA. 
DiagnosisDiagnosis. A species of Pseudocalotes reaching at least 196 mm (67.9 mm SVL) and distinguished from congeners by the following combination of characters: (1) discrete interparietal present; (2) canthals six; (3) enlarged, heavily keeled to subpyramidal posttemporal and posttympanic scales present; postrictal modified scale absent; (4) gulars relatively small, homogenous, no transition to small scales on poorly developed gular pouch; (5) short antehumeral fold with noticeably smaller scales below it (Fig. 7); (6) dorsolateral row of six heavily keeled scales; dorsolateral series reduced to single scale on neck; (7) scales on lower flanks feebly keeled and homogenous (8) 51 scales around midbody (9) dorsal crest of projecting scales extending to base of tail; all projecting scales of crest separated by medial contact between scales of paravertebral series; (10) ventrals smaller than dorsals; (11) subdigital lamellae at base of Toe III bicarinate with preaxial and postaxial keels equally developed; (12) dorsum brown and green, banded in unique holotype; (13) ventral body white with small brown spots densest laterally; (14) gular pouch white medially grading to dirty brown laterally; (15) tongue cream, throat black.

Comparisons. Pseudocalotes rhammanotus can be distinguished from other agamids on Sumatra by its clearly visible tympanum, enlarged row of scales below the orbit bordering the supralabials, relatively wide gap between the dorsal and nuchal crests, heterogenous dorsal squamation, tail about twice as long as SVL, keeled subdigital lamellae, dorsals larger than ventrals, absence of large spines in the nuchal and postorbital regions, relatively narrow head, and other characters of external morphology described in the differential diagnosis of Hallermann & Böhme (2000; see also Mahony 2010).

Three other congeners occur on Sumatra. Of these, Pseudocalotes rhammanotus is most similar to P. tympanistriga. However, unlike P. tympanistriga (characters in parentheses), P. rhammanotus has an antehumeral fold with noticeably smaller scales below it (fold and transition to smaller scales in this region absent), 6/6 heavily keeled scales between the limbs (4 or 5), a wide nuchal gap of 7 scales (narrow, 2 or 3 scales), fewer vertebrals (20 vs. 24–31, 27 ± 3, n = 5), vertebrals that project from the dorsum (vertebrals enlarged with upturned tips but without entire scale projecting from dorsum), fewer subdigital lamellae (24 vs. 26–32, 28 ± 3, n = 5), and fewer infralabials (8 vs. 9–11). Unlike P. cybelidermus and P. guttalineatus (characters in parentheses), P. rhammanotus lacks an enlarged, postrictal modified scale (present, slightly ventral and posterior to border of auditory meatus); has much smaller gulars covering most of the throat without any transition to smaller scales medially on the gular pouch (large scales laterally, transitioning to small granular scales medially on the gular pouch; Fig. 4); 36 gulars from the preaxial margin of the arm to the mental (31 or fewer gulars); only 20 projecting scales making up the dorsal crest (24–34), because medial contact between scales of the paravertebral series separates each vertebral from the one behind it (similar separation limited to posterior body in P. guttalineatus, absent in P. cybelidermus except as apparent anomalies); only a single heavily keeled scale in the dorsolateral series in front of the arm (four or five); a poorly developed longitudinal skin fold of smaller scales spanning the area between the posterior margin of the jaw and anterior margin of the scapula (fold and transition to smaller scales in this region absent); subequal keels on the bicarinate lamellae at the base of Toe III (preaxial keel much more developed, postaxial keel absent on some lamellae, very low on others); and cream tongue (yellow-orange).

Like Pseudocalotes rhammanotus and P. tympanistriga, P. dringi, P. khaonanensis, and P. saravacensis lack modified subdigital lamellae at the base of Toe III. Although reported to have bicarinate subdigital lamellae at the base of Toe III, P. flavigula actually lacks or has greatly reduced postaxial lamellae similar to P. cybelidermus and P. guttalineatus. With 51 scales around midbody, P. rhammanotus has more midbody scales than P. flavigula (38–40) and fewer than either P. khaonanensis (72–75) or P. saravacensis (68). Unlike P. dringi, P. rhammanotus has six canthals (five), heavily keeled gulars (smooth), projecting scales forming the dorsal crest (described as “very low dorsal crest formed by the keeled but not enlarged middorsal scale row” in the original description), and an antehumeral fold (antehumeral fold absent). Like P. rhammanotus, P. austeniana, P. kakhienensis, and P. kingdonwardi have antehumeral folds, however the dorsals of these species are strongly heterogenous in size and shape. All remaining species of Pseudocalotes have modified lamellae at the bases of their third toes. 
CommentBehavior: When threatened, P. rhammanotus darkens in color and gapes its mouth exposing the sharply contrasting cream-colored tongue and black throat.

Abundance: only known from the type specimen (Meiri et al. 2017). 
EtymologyThe new name rhammanotus is an adjective derived from the Greek nouns rhamma meaning suture and notos meaning back. In Pseudocalotes rhammanotus, scales of the paravertebral series separate all scales of the dorsal crest and remind one of sutures along the back of this species. 
  • HARVEY, MICHAEL B.; AMIR HAMIDY, NIA KURNIAWAN, KYLE SHANEY & ERIC N. SMITH 2014. Three new species of Pseudocalotes (Squamata: Agamidae) from southern Sumatra, Indonesia. Zootaxa 3841 (2): 211–238 - get paper here
  • HARVEY, MICHAEL B.; KYLE SHANEY, AMIR HAMIDY, NIA KURNIAWAN, ERIC N. SMITH 2017. A new species of Pseudocalotes (Squamata: Agamidae) from the Bukit Barisan Range of Sumatra with an Estimation of its phylogeny. Zootaxa 4276 (2): 215–232 - get paper here
  • Meiri, Shai; Aaron M. Bauer, Allen Allison, Fernando Castro-Herrera, Laurent Chirio, Guarino Colli, Indraneil Das, Tiffany M. Doan, Frank Glaw, Lee L. Grismer, Marinus Hoogmoed, Fred Kraus, Matthew LeBreton, Danny Meirte, Zoltán T. Nagy, Cristiano d 2017. Extinct, obscure or imaginary: the lizard species with the smallest ranges. Diversity and Distributions - get paper here
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