Toxicocalamus lamingtoni (Kinghorn, 1928) Kraus & Kaiser & O’Shea, 2022

Kraus, Fred, Kaiser, Hinrich & O'Shea, Mark, 2022, Hidden diversity in semi-fossorial Melanesian forest snakes: A revision of the Toxicocalamus loriae complex (Squamata, Elapidae) from New Guinea, Vertebrate Zoology 72, pp. 997-1034 : 997

Toxicocalamus lamingtoni (Kinghorn, 1928) comb. nov.

Figs 1D, D ', 2G, G’, H, H’, 3D, D’, 4G, H

Apisthocalamus lamingtoni Kinghorn, 1928: 290.

Apistocalamus lamingtoni - Roux, 1934: 79.

Toxicocalamus (Apistocalamus) loriae (part) - McDowell, 1969: 456.

Toxicocalamus loriae X T. stanleyanus (part) - McDowell, 1969: 485.

Toxicocalamus loriae Clade 3 - Strickland et al., 2016: 671.

Types and collection.

The specimens on which Kinghorn (1928) based his description of T. lamingtoni (an adult male, AMS R9351; two juveniles, AMS R9352 and R61072) were obtained by C. Terence McNamara (born ca. 1900), the Resident Magistrate (later referred to as District Commissioner) of Mount Lamington District, Northern Division, Papua ( Troughton 1946), during August and September 1927. Of these, Kinghorn designated the male as the holotype and commented on the two juveniles, which we therefore consider to be paratypes. Our examination shows that both juveniles are immature females. The collector appears to have sent one additional specimen from the same locality in the same time frame (AMS R9851), but this has no type status.

Etymology.

Kinghorn (1928: 291) stated that the specimens on which his description was based were all collected in "Mount Lamington district, Northern Division, Papua." It is possible that the author chose the name of the district, which itself takes its name from Mt. Lamington (8.94°S, 148.16°E, elevation 1680 m), a stratovolcano in Oro Province, Papua New Guinea, as the name for the new species. However, this would ordinarily be indicated by the adjectival suffix - ensis, which Kinghorn did not use. He may have been unaware of proper Latinized name formation, as he incorrectly named other species for localities using the genitive case (- i or - ae). Regardless, the person after whom these localities were named is Lord Lamington, Charles Wallace Alexander Napier Cochrane-Baillie (1860-1940), was the 2nd Baron Lamington and a British colonial administrator, who served as the 8th Governor of Queensland (1896-1901) and the 14th Governor of Bombay (1903-1907). The description was published in English.

Diagnosis.

A modestly sized member of the T. loriae Group (male SVL up to 428 mm, female SVL up to 500 mm), with the following unique combination of characters: cloacal plate single; a single intergenial separating posterior genials, widest posteriorly. Preocular elongate, approximately twice as long as high, contacting nasal but not internasal; one postocular; two (92%) or three (8%) posterior temporals; 160-178 ventrals in nine males, 186-195 in nine females, sexually dimorphic without overlap; 41-53 subcaudals in males, 26-34 in females, sexually dimorphic without overlap; SCR 19.3-23.0% in males, 12.2-14.9% in females, sexually dimorphic without overlap; females with very short tails relative to males (TLR sexually dimorphic without overlap, 16.7-20.8% in adult males, 9.0-11.6% in adult females); pale markings on prefrontals absent, even in juveniles; tail spine brown, same colour as remainder of tail; venter uniformly yellow; juveniles with brown anterior supralabials; and head pattern in juveniles typically consisting of a complete, broad, pale band across the nape, parietals, temporals, and last two supralabials, with remainder of head anterior to that lacking pale markings.

Comparisons with other species.

Toxicocalamus lamingtoni is unique within the T. loriae Group and distinguished from all other members of the genus except T. buergersi , T. cratermontanus , and T. stanleyanus in having a single cloacal plate; from these last three species T. lamingtoni is easily distinguished by having the preocular and prefrontal distinct (vs. fused). It is further distinguished from T. loriae in having only a single intergenial (vs. two in T. loriae ), a dark-brown (vs. white in T. loriae ) tail spine, brown anterior supralabials in juveniles (vs. yellow in T. loriae ), and the broad yellow nuchal collar in juveniles (vs. narrow and incomplete in T. loriae ); from T. nymani by its uniformly yellow venter in adults (vs. black or very dark brown in adult T. nymani ), single postocular (usually two in T. nymani ), dark-brown (vs. white in T. nymani ) tail spine, brown anterior supralabials in juveniles (vs. yellow in T. nymani ), and the broad yellow nuchal collar in juveniles (vs. narrow and incomplete in T. nymani ); from T. loennbergii by having two (vs. three in T. loennbergii ) posterior temporals, lacking (vs. possessing) a dark vertebral stripe, and having a dark-brown (vs. white in T. loennbergii ) tail spine; from T. nigrescens by its smaller size (SVL up to 500 mm in T. lamingtoni and 635 mm in T. nigrescens ) and in having a uniformly yellow (vs. grey) venter; and from T. mattisoni in having the preocular contact the nasal (vs. separated by prefrontal contact with the second supralabial in T. mattisoni ) and its uniformly yellow venter (vs. pale grey or yellow with grey band in T. mattisoni ).

Redescription of the holotype.

Adult male, 342 mm SVL + 78 mm TL = 420 mm TTL. Rostral broader than high, notched ventromedially; internasals angulate, semi-triangular, wider than long; prefrontals distinct from preoculars, approximately square but angled posteriorly, slightly longer than wide (Fig. 1D, D '), bordered below by preocular and nasal; preoculars elongate, narrower anteriorly, approximately 2.0-2.5 times as long as deep (Fig. 2G, G ', H, H’), bordered anteriorly by nasal, below by second and third supralabials; frontal shield-shaped, lateral margins angled obliquely, not fused with supraoculars, anterior margin extending slightly anterior to remainder of scale medially; parietals approximately twice as long as wide. Nasals divided by large nares, without grooves above or below naris, though this area dimpled or creased. Postoculars one, irregularly hexagonal in shape, approximately same size as eye; one elongate anterior temporal above fifth and sixth supralabials, separating latter from parietal; two posterior temporals on right (one above the other, with upper larger), three on left (anteriormost smallest followed posteriorly by a larger upper and smaller lower temporal), in either configuration lowest abutting posterodorsal margin of sixth supralabial. Supralabials six, third and fourth entering eye; infralabials six, first four in contact with anterior genial. Mental small, shallow, triangular, wider than deep, bordered behind by first supralabials; anterior genials larger and longer than posterior genials, in medial contact along entire length; posterior genials in narrow anterior contact, otherwise separated by single elongate intergenial, which is widest posteriorly; three gulars separate intergenial from first ventral in the midline; first sublabial separates posterior genial from fifth infralabial (Fig. 3D, D '). Eye relatively small; pupil round.

Dorsal scale rows 15-15-15, smooth, not notched posteriorly, without apical pits. Ventrals 173, each approximately four times wider than long; vent covered by single scale; subcaudals 46, paired. Tail tipped by a pointed conical spine.

In preservative (88 years after collection), dorsum uniformly brown-grey, paler laterally. Venter uniformly pale yellow; medial brown markings scattered on several anterior subcaudals, posterior subcaudals largely brown. Anterior five supralabials and rostral uniformly dark brown, last supralabial brown with large yellow blotch. Head otherwise uniformly dark brown. Chin and throat pale yellow suffused with brown on mental, anterior gulars, and first four supralabials. Tail spine brown, not distinct in colour from remainder of tail but slightly paler at tip. Iris black.

Variation.

Nasals divided by large nares, without grooves above or below naris, though these areas often dimpled or creased. Postoculars one, except two in AMNH R-101103, irregularly hexagonal in shape, smaller than or occupying approximately same area as eye; two (63%) or three (37%) posterior temporals, either one above the other, with upper larger, or with anteriormost smallest followed posteriorly by a larger upper and smaller lower temporal, in either configuration lowest abutting posterodorsal margin of sixth supralabial. Supralabials six, except two specimens with five on one side; third and fourth supralabials contacting eye, except third or second and third in specimens with five supralabials. Anterior genials usually larger and longer than posterior genials but may be subequal; posterior genials entirely separated by single elongate intergenial (n = 5) or in medial contact for first quarter to first three-quarters of length (n = 12); intergenial one (except AMNH R-101103, which has an additional tiny intercalary scale anteriorly), widest posteriorly.

Dorsal scale rows invariably 15-15-15. Ventrals 160-178 (170 ± 5) in nine males, 186-195 (190 ± 3) in nine females; subcaudals 41-53 (46 ± 4) in nine males, 26-34 (29 ± 2) in nine females; SCR 19.3-23.0% (21.4 ± 1.2%) in males and 12.2-14.9% (13.2 ± 0.8%) in females. Tail tipped by a blunt to pointed conical spine. Maximum male SVL 428 mm, TLR = 16.7-20.8% (18.8 ± 1.4%); maximum female SVL 500 mm, TLR = 9.0-11.6% (10.3 ± 0.9%).

In preservative, dorsum uniformly grey or brown-grey in recent specimens, fading to uniform medium brown in specimens retained longer in alcohol. Venter uniformly pale yellow; most larger specimens and one neonate have some brown markings on the posterior subcaudals or midventrally on more anterior subcaudals, but these are never densely arrayed. In the Garaina sample, all supralabials and rostral pale yellow ventrally; in samples from south of there supralabials and rostral often densely suffused with brown or grey; in populations from Mt. Lamington and Cape Nelson, anterior 4-5 supralabials and rostral uniform black or dark brown, posterior supralabials mostly yellow. Yellow markings typically absent on nasals and prefrontals, though vaguely developed on prefrontals in two specimens. Nuchal collar evident in specimens <260 mm SVL but absent or very obscure in specimens> 330 mm SVL, better developed in southern samples; collar narrow in AMNH R-101100 (SVL = 160 mm) but very wide in AMS R9352 (SVL = 163 mm), AMS R61027 (SVL = 167 mm), and BPBM 36171 (SVL = 190 mm), extending from behind head anteriorly across most of parietals, anterior temporals, and supralabials 5 and 6 (Fig. 4G, H ). AMS R61027 also has a yellow blotch centrally located on the anterior frontal and posterior prefrontals. Chin and throat uniformly pale yellow in Garaina samples, with brown suffusion on anterior of chin in all other specimens. Conical tail spine invariably brown, not distinct in colour from remainder of tail.

In life, field notes described BPBM 39813 (a juvenile) as "Slate gray above with yellow nuchal collar. Venter pale gray, with each scale darker anteriorly and lighter posteriorly".

Range.

Restricted to the northern versant of the Owen Stanley Mts. in Oro Province and southern Morobe Province, Papua New Guinea, at elevations from 100-940 m (Fig. 6B ).

Ecological notes.

AMNH R-101100 was ploughed out of an old clump of sugar cane in a field being cleared for a new tea plantation. BPBM 43032 (SVL = 480 mm) contains four shelled eggs.

Figures

Figure 1. Dorsal views of heads of Toxicocalamus loriae Group snakes, presented as both photographic and line-drawing illustrations. A, A ' holotype of T. loriae (MSNG 29141), Haveri, Bartholomew Range, Central Province, PNG. B, B ' lectotype of T. nymani comb. nov. (UUMZ 290 / 2387), Sattelberg, Huon Peninsula, Morobe Province, PNG. C, C ' lectotype of T. loennbergii comb. nov. (BMNH 1946.1.18.24), Fakfak, Onin Peninsula, West Papua Province, Indonesia. D, D ' holotype of T. lamingtoni comb. nov. (AMS R 9351), Mount Lamington, Oro Province, PNG. Images not to scale. Scale abbreviations: anterior temporals (AT), frontal (F), internasals (IN), nasals (N), parietals (P), prefrontals (PF), postoculars (PO), preoculars (PR), posterior temporals (PT), rostral (R), supraoculars (SO).

Figure 2. Right and left lateral views of heads of Toxicocalamus loriae Group snakes, presented as both photographic and line-drawing illustrations. A, A ', B, B ' holotype of T. loriae (MSNG 29141). C, C ', D, D ' lectotype of T. nymani comb. nov. (UUMZ 290 / 2387). E, E ', F, F ' lectotype of T. loennbergii comb. nov. (BMNH 1946.1.18.24). and G, G ', H, H ' holotype of T. lamingtoni comb. nov. (AMS R 9351). Images not to scale. Scale abbreviations: anterior genial (AG), anterior temporal (AT), frontal (F), numbered infralabials (IN 1 - 6), internasal (IN), mental (M), nasal (N), parietal (P), posterior genial (PG), postocular (PO), preocular (PR), posterior temporals (PT), numbered supralabials (SL 1 - 6), supraocular (SO).

Figure 3. Ventral views of heads of Toxicocalamus loriae Group snakes, presented as both photographic and line-drawing illustrations. A, A ' holotype of T. loriae (MSNG 29141). B, B ' lectotype of T. nymani comb. nov. (UUMZ 290 / 2387). C, C ' lectotype of T. loennbergii comb. nov. (BMNH 1946.1.18.24). D, D ' holotype of T. lamingtoni comb. nov. (AMS R 9351). Images not to scale. Scale abbreviations: anterior genials (AG), intergenials (IG), numbered infralabials (IL 1 - 6), mental (M), posterior genials (PG), sublabials (Sb).

Figure 4. Dorsal and right lateral views of heads of juveniles of Toxicocalamus loriae Group snakes. A, B T. loriae (BPBM 10966), Agaun, Milne Bay Province, PNG. C, D T. nymani comb. nov., spotted form (BPBM 5442), Kalolo, Morobe Province, PNG. E, F T. nymani, dark form (BPBM 23699), Wau, Morobe Province, PNG. G, H T. lamingtoni comb. nov. (AMS R 9352), Mt. Lamington, Oro Province, PNG. I, J T. vertebralis sp. nov. (KU 129086), Wau, Morobe Province, PNG. K, L T. spilorhynchus sp. nov. (AMNH R- 107204), Garaina, Morobe Province, PNG. M, N T. atratus sp. nov. (CAS 118958), Mintima, Chimbu Province, PNG.

Figure 6. Type localities (stars) and other collection sites (circles) of Toxicocalamus loriae Group species. A Map of New Guinea and its satellite islands, showing the positions of the inset maps and the type locality of T. mintoni (magenta), an endemic of Sudest Island, Milne Bay Province, PNG. Scale = 500 km. B Papua New Guinea and adjacent areas of West New Guinea, Indonesia, with white lines demarcating provincial boundaries. Scale = 250 km. C West Papua Province, Indonesia (WP), showing the type locality of T. loennbergii comb. nov. (orange). Other species treated in this report are T. loriae (dark blue), T. lamingtoni comb. nov. (yellow), T. spilorhynchus sp. nov. (red), T. nymani comb. nov. (neon green), T. vertebralis sp. nov. (pink), and T. atratus sp. nov. (light blue). Additional related species in the T. loriae clade are T. goodenoughensis (brown), T. mattisoni (violet), T. nigrescens (black), T. pachysomus (dark green), and T. " Toxicocalamus loriae " Clade 4 (white). The type locality of the recently described dubious taxon T. longhagen (tan) is also indicated. Scale = 250 km. Provinces of PNG are Central (Ce), Chimbu (Ch; also spelled Simbu), Gulf (Gu), Hela (He; created in 2012), Eastern Highlands (EH), East Sepik (ES), Enga (En), Jiwaka (Ji; created in 2012), Madang (Ma), Milne Bay (MB), Morobe (Mo), National Capital District (NCD), Oro (Or; also known as Northern Province); Sandaun (Sa; formerly West Sepik), Southern Highlands (SH), Western (We), and Western Highlands (WH). Provinces of West New Guinea are Highland Papua (HP; created in 2022), Papua (Pa), and South Papua (SP; created in 2022). Sites of sympatry are Wau (W; neon green + pink) with two species and Garaina (G; neon green + red + yellow) with three species.