Prasinohaema, GREER, 1974

Slavenko, Alex, Tamar, Karin, Tallowin, Oliver J S, Kraus, Fred, Allison, Allen, Carranza, Salvador & Meiri, Shai, 2022, Revision of the montane New Guinean skink genus Lobulia (Squamata: Scincidae), with the description of four new genera and nine new species, Zoological Journal of the Linnean Society 195 (1), pp. 220-278 : 234-237

PRASINOHAEMA GREER, 1974

(CLADE II)

( FIG. 5 ; SUPPORTING INFORMATION, FIG. S6 ; TABLE 1 ) Prasinohaema Greer, 1974 . Australian Journal of Zoology Supplementary Series (31): 1–67.

Type species: Lygosoma flavipes Parker, 1936 , by original designation.

Diagnosis: Large (adult SVL up to 103 mm; Meiri, 2018) arboreal skinks with short limbs (forelimbs 27.7–31.4% of SVL, hindlimbs 29.9–34.1% of SVL); lobules present on anterior edge of ear opening; two pairs of chin shields in medial contact; three supralabials posterior to subocular supralabial; chin shields separated from infralabials by a row of genials; lower eyelid with window variable in size, opaqueness and scaliness; temporal region fragmented (> 3 scales); nasal scale undivided; frontoparietals unfused; viviparous; litter size 2–9; green blood serum and tissues; tail prehensile with a glandular tip; subdigital lamellae greatly expanded basally.

Prasinohaema differs from Lobulia and Papuascincus by having green blood serum and tissues ( Greer, 1974), a prehensile tail with a glandular tip and basally expanded subdigital lamellae, by having the chin shields separated from the infralabials by a row of genials (vs. chin shields abutting the infralabials) and by having a fragmented temporal region (vs. the standard three-scale arrangement). It further differs from Papuascincus by having two pairs of chin shields in medial contact (vs. one), unfused (vs. fused) frontoparietals, an undivided (vs. divided) nasal scale and by its viviparous (vs. oviparous) reproductive mode.

Species included: Prasinohaema flavipes ( Parker, 1936) ; Prasinohaema prehensicauda (Loveridge, 1897) .

Species incertae sedis: Prasinohaema parkeri ( Smith, 1937) was originally placed in Prasinohaema by Greer (1974), seemingly based on having basally enlarged subdigital lamellae and transverse cross-bands on the dorsum, a coloration pattern it shares with Pr. prehensicauda and Pr. flavipes , but also with Pr. semoni which is phylogenetically distant from the former two species ( Fig. 1 ). However, no information was given in Smith (1937) regarding the condition of its tail or the colour of its blood serum or tissues, data for the latter of which would not have been available for Greer in his revision ( Greer, 1974) since the species was never collected after its original description. Furthermore, Pr. parkeri lacks lobules on the anterior edge of the ear opening and has a unique arrangement of the frontal (contacting the three vs. two anteriormost supraoculars) and prefrontals (fused with the anterior loreals). Pr. parkeri is only known from its type specimen ( Meiri et al., 2018) collected in the Utakwa River ( Smith, 1937), presumably along the southern slopes of the Sudirman Range ( Wollaston, 1914). Although the presence of basally expanded subdigital lamellae and cross-bands may suggest an affinity with Pr. prehensicauda and Pr. flavipes , these traits are also common in at least some other New Guinean skinks (e.g. basally expanded subdigital lamellae in Li. longiceps , cross-bands and basally expanded subdigital lamellae in Pr. semoni ), and therefore its placement in Prasinohaema is uncertain. Similarly, the presence of green blood serum and tissues alone would not be enough to place it in Prasinohaema , as both Pr. semoni and Pr. virens possess this trait but are otherwise morphologically and phylogenetically distant from Pr. prehensicada and Pr. flavipes ( Figs 1–2 ).

Distribution: The two species in the genus ( Pr. flavipes and Pr. prehensicauda ) are widespread in the montane regions of Papua New Guinea. Prasinohaema prehensicauda is present in the New Guinea Highlands and on the Papuan Peninsula, whereas Pr. flavipes also occurs on the Huon Peninsula.

Remarks: Two other species are currently assigned to the genus Prasinohaema : Pr. semoni and Pr. virens . These species emerge in our analyses as phylogenetically distant from the type species of the genus, Pr. flavipes ( Fig. 1 ; Rodriguez et al., 2018), rendering the former concept of the genus polyphyletic. They also differ widely morphologically ( Fig. 2 ), reproductively ( Pr. virens is oviparous, whereas Pr. semoni , Pr. flavipes and Pr. prehensicauda are viviparous; Fig. 2 ) and in elevational range ( Pr. semoni and Pr. virens are lowland species, whereas Pr. flavipes and Pr. prehensicauda are montane species; Fig. 4 ). Many of these differences, particularly in Pr. virens , were mentioned by Greer even in his original description of the genus ( Greer, 1974). Therefore, we stress that Prasinohaema is in need of taxonomic revision. Prasinohaema semoni and Pr. virens likely need to be assigned to new genera, although this is beyond the scope of the current work.

Figures

Figure 1. A. Bayesian Inference phylogenetic tree based on the concatenated data set of three mitochondrial and two nuclear markers, with BI posterior probabilities and ML bootstrap support values shown at each node, respectively. B, Bayesian Inference multilocus species tree, with BI posterior probabilities shown at each node. Colours in both panels correspond to each of seven clades representing seven different genera: Lobulia (Clade I, red), Prasinohaema (Clade II, green), Nubeoscincus (Clade III, light blue), Papuascincus (Clade IV, blue), Palaia (Clade V, orange), Alpinoscincus (Clade VI, yellow) and Ornithuroscincus (Clade VII, purple).

Figure 2. Time-calibrated phylogenetic tree, based on one specimen per species, with posterior probability values indicated above the nodes. Blue bars represent 95% HPD of divergence date estimations. Colours correspond to the same clade assignments as in Figure 1. The matrix on the right represents character states of nine morphological characters for each species. Blank squares represent missing data.

Figure 4. Ancestral-state reconstruction of elevations occupied by this clade of skinks. A, tips are coloured according to the known minimum elevation (m a.s.l.) for each species. Pie charts at each node denote the posterior probabilities of each state, based on 100 simulated character histories. B, phenogram showing evolution of minimum elevation along the timecalibrated phylogeny under a BM model, with branches coloured according to minimum elevation (m a.s.l.).

Figure 5. Representatives of each of the seven revised genera, not to scale: (A) Lobulia huonensis (BPBM 40322); (B) Prasinohaema flavipes (BPBM 40369); (C) Nubeoscincus glacialis (BPBM 14712); (D) Papuascincus lineage VII (BPBM 44749); (E) Palaia pulchra (BPBM 38789); (F) Alpinoscincus alpinus (BPBM 44218); (G) Ornithuroscincus pterophilus (BPBM 45705). Images by Allen Allison.

Figure 6. Pictures in life of (A) Lobulia brongersmai (BPBM 34733); (B) Lobulia elegans (BPBM 18692); (C) Lobulia fortis (paratype; BPBM 41130); (D) Lobulia huonensis (holotype; BPBM 40322); (E) Lobulia lobulus (BPBM 47837); (F) Lobulia marmorata (holotype; BPBM 34150). Images A, B, F by Fred Kraus, images C–E by Allen Allison.