Toxicocalamus loennbergii (Boulenger, 1908) 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 loennbergii (Boulenger, 1908) comb. nov.

Figs 1C, C ', 2E, E’, F, F’, 3C, C’

Apisthocalamus loennbergii Boulenger, 1908: 248.

Apistocalamus loennbergi - Sternfeld, 1913: 387.

Apistocalamus Apistocalamus lönnbergi - de Rooij, 1917: 260.

Apistocalamus loriae (part) - McDowell, 1967: 537.

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

Types and collection.

As reported by Boulenger (1908) in the original description, four specimens were collected by the British naturalist and explorer Antwerp Edgar Pratt (1852-1924) during his sojourns in New Guinea with his collector sons Charles Henry Pratt (1886-1936) and Felix Thomas Biet Pratt (1891-1955). Pratt spent the months of May-June 1905 and December 1907-January 1908 near Fakfak on the Onin Peninsula (formerly Fakfak Peninsula), an extension of the Bomberai Peninsula in Dutch New Guinea ( Papua Insects Foundation 2015). The specimens on which Boulenger based his description (four females, BMNH 1946.1.18.24-26, MCZ R-76634) were originally accessioned as BMNH 1908.6.30.7-10 with one of them (BMNH 1908.6.30.10) exchanged with the MCZ in 1963. These would have been collected on Pratt’s earlier voyage, given that specimens collected in January 1908 could not have reached Boulenger in time for a March 1908 publication date, because delivery of passengers (or specimens) from New Guinea to London on the prevalent early 20th Century steamship route via Australia and the Suez Canal would have taken at least 75 days ( Phillips 2002). Thus, we presume the type material of T. loennbergii was collected in May or June 1905.

Because Boulenger (1908) did not designate a holotype, McDowell (1969: 456) designated BMNH 1946.1.18.24 as the “holotype” of A. loennbergii . This designation is invalid because a holotype cannot be designated from among syntypes after the original description. We nevertheless follow McDowell and designate BMNH 1946.1.18.24 as the lectotype of A. loennbergii . This is the largest specimen (SVL = 565 mm). The remaining syntypes (BMNH 1946.1.18.25-26 and MCZ R-76634) become paralectotypes.

Etymology.

Named by Boulenger (1908) for Professor Einar Lönnberg (1865-1942), the Swedish zoologist who described Pseudapistocalamus nymani . The description was published in English.

Diagnosis.

A modestly sized member of the T. loriae Group (maximum SVL 565 mm, only females known) with the following unique combination of characters: two scales covering vent; four infralabials contacting anterior genial; a single intergenial separating posterior genials, widest posteriorly. Preocular elongate, approximately twice as long as wide, contacting nasal (62%) or not (38%), not contacting internasal; relatively short snout (SNL/SNW x̄ = 0.95, range = 0.93-0.99); relatively small eye (EY/SNL x̄ = 0.16, range = 0.15-0.18); one postocular (fused to supraocular on one side of one specimen); three posterior temporals; 214-220 ventrals in four females; 23-32 subcaudals; SCR 9.7-12.7%; dark vertebral stripe; large pale blotch on parietal; pale markings on prefrontals absent (50%), small or vaguely developed (25%), or well developed (25%), best developed in the smallest specimen; tail spine white, paler than remainder of tail; and venter uniformly yellow.

Comparisons with other species.

Toxicocalamus loenn-bergii can be distinguished from all other members of the T. loriae Group except some T. loriae and juvenile T. nymani by its uniformly yellow venter. It can be distinguished from T. loriae in having only a single intergenial (two in T. loriae ) and from juvenile T. nymani by having four infralabials in contact with the anterior genial (three in 87.5% of T. nymani ), a single postocular (usually two in T. nymani ), and a dark vertebral stripe (absent in T. nymani ). It can further be distinguished from T. nigrescens by its greater number of ventrals (214-220 vs. 184-193 in female T. nigrescens ), and dark vertebral stripe (absent in T. nigrescens ); and from T. mattisoni in having the preocular contact the nasal (separated by prefrontal contact with the second supralabial in T. mattisoni ), its greater number of ventrals (214-220 vs. 170-181 in female T. mattisoni ), and dark vertebral stripe (absent in T. mattisoni ).

Description of the lectotype.

An adult female, 565 mm SVL + 55 mm TL = 620 mm TTL. Rostral wider than tall, notched ventromedially; internasals angulate, semi-triangular, wider than long. Prefrontals distinct from preoculars, approximately square, rounded posterolaterally, bordered below by preocular and nasal, in point contact with second supralabial on right side; preoculars elongate, narrower anteriorly, approximately twice as long as tall (Fig. 1C, C '), bordered anteriorly by nasal, below by second and third supralabials (Fig. 2E, E ', F, F’). Nasal divided by large naris, without grooves above or below naris. Postoculars one on left, fused to supraocular on right; irregularly hexagonal in shape, longer than tall, occupying approximately same area as eye. 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; one elongate anterior temporal above fifth and sixth supralabials, separating sixth from parietal; three posterior temporals, lowest abutting posterodorsal margin of sixth supralabial. Supralabials six, third and fourth contacting eye; infralabials six, first four in contact with anterior genial. Mental small, shallow, triangular, wider than long, bordered posteriorly by first supralabials; anterior genials slightly larger than posterior genials, in medial contact along their entire length; posterior genials in medial contact for first 20% of their length; intergenial single, diamond-shaped, widest posteriorly; five gulars separate intergenials from first ventral in the midline; first sublabial separates posterior genial from fifth infralabial on right but not on left (Fig. 3C, C '). Eye relatively small; pupil round.

Dorsal scales smooth, not notched posteriorly, without apical pits, in 15-15-15 rows; Ventrals 214, each approximately four times wider than long; two scales covering vent; subcaudals 23, paired. Tail tipped by a blunt conical spine.

In preservative (108 years after collection), dorsum reddish brown, darker mid-dorsally, with darker-brown vertebral stripe one scale wide; each dorsal scale with darker brown edges, imparting a reticulated appearance to dorsum, especially laterally. Venter uniformly pale yellow; subcaudals lightly edged in brown at their medial junctures. Supralabials pale yellow ventrally, suffused with brown dorsally. Top of head with vague traces of yellow blotches on parietals and anterior temporals; no pale nuchal collar. Chin and throat pale yellow, with small amount of brown suffusion on mental, sutures between infralabials, and anterior suture of anterior genial. Tail spine brown above and white below. Iris black.

Variation.

All specimens are female. Prefrontals are bordered below by the preocular and the nasal, but they are in point contact with the second supralabial on one side in two specimens; preoculars are bordered below by the second and third supralabials, except by only the third supralabial on the right side of BMNH 1946.1.18.25. One postocular, except when it is fused to the supraocular on the right side of BMNH 1946.1.18.24, smaller than or occupying approximately same area as the eye. Six supralabials, except seven on the right side of MCZ R-76634; third and fourth supralabials contact the eye, except when Supralabial 2 also contacts it, as on the right side of MCZ R-76634. The posterior genials are entirely separated by a single elongate intergenial in MCZ R-76634, in point contact anteriorly in BMNH 1946.1.18.26, and in medial contact for the first 20-35% of length in the remaining two specimens.

Dorsal scales invariably in 15-15-15 rows. Ventrals 214-220 (216 ± 2); subcaudals 23-32 (28 ± 3); SCR 9.7-12.7% (11.3 ± 1.1%). Tail tipped by a blunt to pointed conical spine. Maximum SVL 565 mm, TLR 8.9-11.2% (10.2 ± 0.9%).

In preservative, all specimens are coloured as the holotype dorsally and ventrally. Subcaudals are lightly edged in brown at their medial junctures but dusted with brown posteriorly in MCZ R-76634. Supralabials are pale yellow ventrally, variably suffused with brown on dorsal portions of some or all supralabials (absent on second supralabial in two specimens). In the smallest specimen (BMNH 1946.1.18.25, SVL = 373 mm), yellow markings occupy a band across the prefrontals and the anterior portion of the frontal, there is a large yellow blotch on each parietal and a complete yellow collar. In BMNH 1946.1.18.26, there is a yellow blotch on each anterior temporal in addition to these other markings. In larger specimens, these markings become suffused with brown and may disappear, with the markings on the prefrontals disappearing first. Conical tail spine white or brown above and white below (BMNH 1946.1.18.24).

Range.

Known only from the type locality, north of Fakfak town on the Onin Peninsula, West Papua Province, Indonesia, at an elevation of 520 m (Fig. 6C ).

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 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.