Phymaturus (Diaz Gomez, 2008)

Lobo, Fernando, Barrasso, Diego A., Hibbard, Thomas & Basso, Néstor G., 2016, On the evolution and diversification of an Andean clade of reptiles: combining morphology and DNA sequences of the palluma group (Liolaemidae: Phymaturus), Zoological Journal of the Linnean Society 176 (3), pp. 648-673 : 666-670

GENUS PHYMATURUS

Below we present the list of new morphological characters that are added to the morphological data matrix of Lobo et al. (2012a). Several of these characters were discovered at the time of the 2012 analysis, but they were not included because they lacked a wide revision across taxa. *Asterisks refer to characters that are informative for the palluma group only.

*Character 208: White scales spread out over dorsum and flanks (binary polimorphic): 0, absent; 1, present. Character shared by P. punae , P. sp. lar, P. sp. gua. Not fixed in any species.

Character 209: Occurrence of melanistic individuals (binary polimorphic): 0, absence; 1, presence. Character shared by P. ceii , P. cf. ceii , and P. tenebrosus . Not fixed in any species.

**Character 210: White spots on dorsum of trunk arrangement (binary polimorphic): 0, irregularly distributed; 1, forming transversal rows ( P. cf. ceii , P. felixi , and a few individuals of P. calcogaster and P. manuelae ). Not fixed in any species.

Character 211: Tails spotted white in a similar way to dorsum of trunk: 0, absent; 1, present. Character shared by P. delheyi , P. nevadoi , P. payuniae , and P. sitesi .

Character 212: Light brown over dorsum of trunk: 0, absent; 1, present. Character shared by P. sitesi , P. tenebrosus , and P. zapalensis .

*Character 213: Dorsal mesenterium of large intestine pigmentation ( Fig. 6A and B ): 0, absent (translucid); 1, present (melanic pigmented).

*Character 214: External wall of rectum (multistate polymorphic; Fig. 6C ): 0, smooth; 1, shallowly striated; 2, deeply striated.

*Character 215: Internal mucosa of rectum ( Fig. 6D ): 0, simple, without folds; 1, small longitudinal folds; 2, large longitudinal folds.

*Character 216: Pigmentation of connective dorsal fascia of axial musculature ( Fig. 7 ): 0, no pigmentation; 1, dorsal fascia of transversospinalis pigmented; 2, dorsal fascia of both epiaxial muscles pigmented (light brown); 3, dorsal fascia of longissimus dorsi brown pigmented and transversospinalis without pigmentation; 4, dorsal fascia dark brown, but lateral fascia still evident; 5, dorsal fascia dark brown, lateral fascia difficult to recognize.

*Character 217: Anterior projection of dorsal fascia melanism: 0, only reaching level of shoulders; 1, reaching level over nuchal musculature.

*Character 218: Neural spines ( Fig. 7B and D ): 0, neural spines inconspicuous; 1, neural spines of vertebrae notable from the transversospinalis pigmentation.

*Character 219: Number of pigmented longitudinal/ lateral fascia separating epaxial muscles ( Fig. 7 ): 0, absent; 1, five; 2, six; 3, seven; 4, eight; 5, nine.

*Character 220: Dorsal tibial scale shape (binary polymorphic; Fig. 8 ): 0, round (juxtaposed); 1, subpentagonal elongated in the proximal–distal axis (subimbricated).

*Character 221: Relative size of dorsal and anteri- or tibial scales (binary polymorphic): 0, similar size; 1, dorsal tibial scales larger than anterior scales.

*Character 222: Tibial scale structures ( Fig. 8B ): 0, dorsal tibial scales with spines and scale organs on their distal tips; 1, dorsal tibial scales with scale organs (not spines).

*Character 223: Granular scales among dorsal tibials ( Fig. 8F ): 0, absent; 1, present (binary polymorphic).

*Character 224: Posterior tibial scales (binary polymorphic): 0, enlarged in the distal third of tibia length; 1, enlarged along its entire length.

*Character 225: Pigal scales of the posterior half of the precloacal region: 0, smaller in females than in males; 1, similar size in both sexes.

*Character 226: Fecundity (species average number of embryos per female): 0, 1.00; 1, 1.5–1.66; 2, 1.75– 1.77; 3, 1.80; 4, 2.00–2.25; 5, 2.66; 6, 3.0. This character would be analysed as a continuous character, but sample sizes are limited, and because Phymaturus species are considered ‘vulnerable’ in Abdala et al. (2012), collecting pregnant females to achieve more accurate information is not recommended. Most species exhibit states 2–4.

Character 227: Ocelli formed by the incomplete and irregular confluence of white spots: 0, absence; 1, presence.

Character 228: Females with bright yellow over cloaca and thighs and posterior region of abdomen orange: 0, absent; 1, present ( P. tenebrosus and P. zapalensis ).

Character 229: Ontogenetic sequence of transversal row of white spots (step matrix/Sankoff).

Character 230: Ontogenetic sequence of black transversal stripes (step matrix/Sankoff).

Character 231: Ontogenetic sequence of lightbrown ocelli (step matrix/Sankoff).

Character 232: First chevron location: 0, beyond second caudal vertebra; 1, second caudal vertebra.

*Character 233: Number of temporal scales with two or more scale organs/number of temporal scales with only one scale organ ( Fig. 9A ): 0, <0.20; 1, ≥0.20.

*Character 234: Nasal salt excretion: 0, absent; 1, present (Lobo et al., 2012c: fig. 5). At this time found only in the Puna clade: P. aguedae , P. bibronii , P. denotatus , P. extrilidus , P. laurenti , P. paihuanense , and P. sp. lar.

*Character 235: Scale organs on rostral scale (binary polymorphic): 0, present; 1, absent.

*Character 236: Number or scales between the row of precloacal pores and the anterior border of cloaca ( Fig. 9B ): 0, 0–4 scales; 1, ≥5 scales. The plesiomorphic condition is exhibited by Ctenoblepharys and Liolaemus (state 0), see Etheridge (1995).

*Character 237: Part of adult female population with the whole body brown ‘chocolate’ coloration (binary polymorphic; Fig. 9C ): 0, absence; 1, presence. Character shared by P. dorsimaculatus , P. tromen ; P. verdugo .

*Character 238: Supratemporal shape: 0, wide, with its lower margin convex, resembling the shape of a knife; 1, slender, upper and lower margins straight and parallel.

*Character 239: Supratemporal length: 0, long, its anterior tip reaches at least a position at the level of the posterior medial margin of parietal, close to the epipterygoid connection; 1, short, not reaching this level, just half of the squamosal process of the parietal.

*Character 240: Orbitosphenoid ossification ( Figs 10A and B ): 0, reaching dorsally the ventral margin of fenestra epioptica; 1, not reaching this fenestra.

*Character 241: Posteromedial process of ectopterygoid (multistate polymorphic; Figs 10C and D ): 0, absent; 1, round and short; 2, forming a curved spine with a cartilage at its free ending.

*Character 242: Lateral and lateral–dorsal antebrachial scales: 0, round and smooth; 1, round to leaf-shaped, with a posterior small spine; 2, leafshaped, with a conspicuous posterior spine.

*Character 243: Enlarged external postcloacal scales in males: 0, absent; 1, present.

*Character 244: Scale organs under spines of dorsal and lateral tail scales: 0, absent; 1, one or two scale organs. Present in P. sp. lar, absent in P. dorsimaculatus .

Character 245: Mid-dorsal longitudinal band formed by enlarged white spots irregularly distributed (binary polymorphic): 0, absence; 1, presence.

*Character 246: Fusion of first postocular with subocular scale (binary polymorphic): 0, absent; 1, present. Not fixed in any species. Significant variation within the patagonicus group; only detected in P. sp. gua within the palluma group.

*Character 247: Wide opening of nares with its anterior lower border projected inside the opening, forming a flat and wide surface (binary polymorphic): 0, absent; 1, present. Not fixed in any species.

*Character 248: Irregularly divided row of precloacal pores (not at the middle; binary polymorphic): 0, absent; 1, present. Not fixed in any species.

*Character 249: Males with yellow dorsum becoming white on the neck (binary polymorphic): 0, absent; 1, present ( Fig. 9D ). Not fixed in any species.

*Character 250: Dorsal and lateral tail scales with longitudinal rugosities: 0, absent; 1, present ( palluma group apomorphy). Not listed in Lobo & Quinteros (2005), but described in the discussion under the section ‘Observations on the ontogenetic shift of morphological characters’. In advanced unborn fetuses and juveniles of the palluma groups these scales are smooth (as in juvenile and adult individuals of the patagonicus group), becoming rugose later.

*Character 251: Juvenile pattern provided of paravertebral irregular shaped markings: 0, absence; 1, presence.

*Character 252: Melanic spots on sides of neck and shoulders (binary polymorphic): 0, absence; 1, presence ( Fig. 9E ). Not fixed in any species.

*Character 253: Hemipenis pigmentation: 0, sulcus melanic, with pigmentation invading surface of lobes; 1, sulcus completely melanic, with pigmentation projected in both branches of its bifurcation, lobes immaculate ( Fig. 9F ); 2, completely unpigmented.

Figures

Figure 6. A, transparent mesenterium of Phymaturus palluma (MCN 2651; character 213). Scale bar: 10 mm. B, same mesenterium, but completely melanic in Phymaturus patagonicus (MCN 3561). Scale bar: 10 mm. C, external surface of rectum of Phymaturus verdugo (MCN 1961; character 214). Scale bar: 5 mm. D, Longitudinal folds of the internal mucosa of rectum in Phymaturus payuniae (MCN 2879; character 215). Scale bar: 5 mm.

Figure 7. Epiaxial musculature fascia (characters 216–219): A, no pigmentation of fascia dorsal among axial muscles of Phymaturus roigorum (MCN 2102); B, lateral pigmentation between axial muscles forming longitudinal lines, neural spines delineated in black in Phymaturus aguanegra (MCN 3288). C, lateral and dorsal fascia of axial muscles pigmented in Phymaturus spurcus (FML 1244). D, same condition, but less melanic, observed in Phymaturus tenebrosus (MCN 1263). Neural spines highlighted in white. Scale bars: 10 mm. Abbreviations: ld, longissimus dorsi muscle; ts, transversospinalis muscle.

Figure 8. A, tibial dorsal scalation of Phymaturus patagonicus (MCN 3561). B, central area of the same view of P. patagonicus (same specimen) tibia, showing scales in detail (character 220). Almost every dorsal tibial scale carries a spine and a scale organ on the distal tip of the scale (arrow; character 222). C, tibial dorsal scalation of Phymaturus palluma (MCN 3131). D, central area of the same view of P. palluma (same specimen). Scale organs hidden under the scale spines. E, tibial dorsal scalation of Phymaturus denotatus (MCN 3160). F, central area of the same view of P. denotatus (same specimen). The arrow indicates granular scales spread out among tibial scales (character 223). Scale bars: A, C, E, 5 mm; B, D, F, 2 mm.

Figure 9. A, temporal scales of Phymaturus dorsimaculatus (MCN 1569) showing two organ scales (character 233); scale bar, 2 mm. B, cloacal region of Phymaturus verdugo (MCN 1960; Character 236); scale bar, 5 mm. C, female of Phymaturus verdugo (MCN 1958; snout–vent length, SVL = 99.8 mm; character 237). D, Phymaturus querque male (MCN 3863; SVL = 104.9 mm; character 249). E, Phymaturus paihuanense female (SSUC–Re 0422; photo by A. Laspiur; character 252). F, hemipenis of Phymaturus roigorum (MCN 1963; character 253); scale bar, 5 mm.

Figure 10. A, orbitosphenoid bone in Phymaturus aguanegra (MCN 968; character 240); the arrow indicates the cartilaginous margin of the fenestra epioptica. B, Phymaturus zapalensis (MCN 1486). C, ectopterygoid of Phymaturus spurcus (MCN 1249; character 241). D, same bone in Phymaturus laurenti (MCN 326). Abbreviations: ecp, ectopteygoid; fep, fenestra epioptica; fm, fenestra metoptica; fop, fenestra optica; fpr, fenestra prootica; j, jugal; pt, pterygoid. Scale bars: 1 mm.