Toxicocalamus nymani ( Loennberg , 1900) 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 nymani ( Loennberg, 1900) comb. nov.

Figs 1B, B ', 2C, C’, D, D’, 3B, B’, 4C, D

Pseudapistocalamus Nymani Lönnberg, 1900: 578.

Pseudapisthocalamus nymani - Boulenger, 1908: 249.

Pseudapistocalamus nymani - de Rooij, 1917: 263.

Pseudapistocalamus nymanni - Klemmer, 1963: 328.

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

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

Types and collection.

The Swedish zoologist Einar Lönnberg (1865-1942) described this species on the basis of three specimens from Sattelberg, German New Guinea, at the eastern end of the Huon Peninsula (now in Morobe Province, PNG). These were collected by the Swedish botanist Erik Nyman (1866-1900), who visited the area from June-November 1899 ( Nordstedt 1899, 1900; Svedmark 1900). Whereas Lönnberg (1900) did not designate a holotype, he provided a little more information for the largest specimen, a female with approximate measurements of SVL = 445 mm and TL = 47 mm, and which contained eggs of about 13 mm length. De Rooij (1917) examined what she called the "type-specimen [...] from the Upsala Museum" and confirmed the size of the eggs. The measurements of SVL = 398 mm and TL = 47 mm she provided for the species just before her discussion of the “type-specimen” do not correlate with the Uppsala specimen and may refer to one of the other specimens from "New Guinea" she examined. We have examined the sole specimen of this series remaining at the Uppsala Museum, an untagged specimen that was loaned to FK as UUZM 290 in 2003 and to MOS as UUZM 2387 in 2013. Our measurements of this female specimen are SVL = 435 mm and TL = 42 mm, which is very close to the size reported by Lönnberg for his largest specimen, given more than a century of possible shrinkage in ethanol. Of the other two specimens referred to in the original description, one is now deposited as BMNH 1946.1.17.57 (originally catalogued as BMNH 1900.9.21.6) and is noted in their catalogue and on their jar label to be "one of the types." It is also a female, and its measurements are SVL = 325 mm and TL = 43 mm, so it is clearly too small to be the large specimen noted by Lönnberg (1900). We are unable to locate the third specimen or any other museum specimen from Sattelberg, so we presume the third syntype is lost. As the larger specimen of the two extant syntypes and as the one with nearly the same measurements as those provided in the original description, we designate UUZM 290/2387 as the lectotype of Pseudapistocalamus Nymani Lönnberg, 1900. This renders the second remaining specimen from the type series (BMNH 1946.1.17.57) a paralectotype.

Etymology.

The species was named for Erik Nyman, the collector of the type series. It also serves as a tribute to a remarkable scientist, who died in 1900 on the journey home to Europe. The description was published in -English.

Diagnosis.

A modestly sized member of the T. loriae Group (maximum SVL in males 422 mm, in females 540 mm) with the following unique combination of characters: two scales covering vent; three infralabials contacting anterior genial; posterior genials entirely separated (45%) by a single intergenial or in anterior (53%) or entire (2%) contact with each other; intergenial usually widest posteriorly or centrally. Preocular approximately as long as wide and never twice as long as wide, usually contacting nasal (94%), not contacting internasal; one (4%) or two (96%) postoculars; one (1%), two (69%), three (27%), four (1%), or five (1%) posterior temporals; 178-198 ventrals in 18 males, 191-210 in 22 females, sexually dimorphic with overlap (t 37 = 4.9581, p <0.00001); 39-48 subcaudals in males, 26-37 in females, sexually dimorphic without overlap; SCR 17.5-20.7% in males, 11.6-15.8% in females, sexually dimorphic without overlap; yellow nuchal collar and yellow markings on prefrontals absent (48%), small or vaguely developed (7%), or present (45%); tail spine paler than remainder of tail; venter uniformly dark brown or dark brown with the posterior of each ventral paler brown or yellowish brown, giving a banded appearance (reported as "blackish brown, edged with light grey" within one year of preservation; Lönnberg 1900).

Comparisons with other species.

Toxicocalamus nymani is distinguished from T. loriae in having a single intergenial (two in T. loriae ) that is widest posteriorly or centrally (widest anteriorly in T. loriae ), by its short, squarish preocular (elongate, more than twice as long as high, in T. loriae ), and by its dark brown or black venter (yellow with or without mid-ventral series of brown spots in T. loriae ); from T. nigrescens by its smaller size (maximum SVL = 540 mm vs. 635 mm in T. nigrescens ), short preocular (elongate, more than three times as long as high, in T. nigrescens ), and in having two postoculars (one in T. nigrescens ) and a dark brown or black venter (grey in T. nigrescens ); and from T. mattisoni in having the prefrontal usually (93%) excluded from contacting the second supralabial (prefrontal and second supralabial in contact in T. mattisoni ), short preocular (elongate, more than twice as long as high, in T. mattisoni ), and in having two postoculars (one in T. mattisoni ) and a dark brown or black venter (yellow or pale grey with grey bands across each ventral in T. mattisoni ).

Description of the lectotype.

An adult female, 435 mm SVL + 42 mm TL = 477 mm TTL. Rostral wider than tall, notched ventromedially; internasals angulate, semi-triangular, wider than long. Prefrontals distinct from preoculars, approximately square but angled posteriorly, slightly longer than wide (Fig. 1B, B '), bordered below by preocular and nasal; preoculars angulate but approximately as high as long, bordered anteriorly by nasal, below by second and third supralabials (Fig. 2C, C ', D, D’). Nasals divided by large nares, with two grooves below and partial groove above naris on right, no grooves on left, both nasals damaged. Postoculars two, irregularly pentagonal in shape, upper larger, slightly smaller than eye. Frontal shield-shaped, not fused with supraoculars, anterior margin extending slightly anterior to remainder of scale medially, lateral margins curved posteriorly; parietals approximately twice as long as wide. One elongate anterior temporal above fifth and sixth supralabials, separating latter from parietal; two posterior temporals, upper twice size of lower, with lower abutting posterodorsal margin of sixth supralabial. Supralabials six on right, seven on left due to division of sixth, third and fourth contacting eye; infralabials six, first three in contact with anterior genial. Mental small, shallow, triangular, wider than long, bordered posteriorly by first supralabials; anterior genials slightly larger than posterior genials but of approximately same length, in medial contact along their entire length; posterior genials completely separated by single elongate intergenial that is widest posteriorly; five gulars separate intergenials from first ventral in the midline; first sublabial separates posterior genial from fifth infralabial (Fig. 3B, B '). Eye relatively small; pupil round.

Dorsal scales smooth, not notched posteriorly, without apical pits, in 15-15-15 rows; ventrals 202, each approximately four times as wide as long; two scales covering vent; subcaudals 28, paired. Tail tipped by a pointed conical spine.

In preservative (114 years after collection), dorsum dark brown dorsally, paler laterally, with brown centre of each scale becoming paler yellow brown at posterior edges. Each ventral scale, including subcaudals, dark brown with paler yellow brown along posterior edges, imparting an overall impression of a dark venter banded with dark brown. Supralabials and rostral largely yellow, all with dark-brown upper margins. Head dark brown with large yellow blotch on each prefrontal and small yellow blotch in anterior part of each preocular. Incomplete yellow nuchal collar, interrupted mid-dorsally by three rows of brown dorsal scales. Chin and throat pale yellow, suffused with brown on mental and anterior portions of anterior gulars and first five infralabials. Tail spine brown, paler than remainder of dorsum. Iris black.

Variation.

Preoculars are not in contact with the nasals in three specimens from Madang Province (AMS R25304, R25752, IRSNB 733678), in which they are separated by contact between prefrontal and second supralabial; in contact with internasals only in MCZ R-76627. Two postoculars, except one on both sides of BPBM 3397 and one on the left side of the paralectotype. Posterior temporals two (in 69% of sides), one (1%), three (27%), four (1%), or five (1%). Six supralabials, except seven on left sides of lectotype and AMS R25608, seven on both sides of BPBM 17173, and five on right side of BPBM 31257 and left side of PNGM 24716; third and fourth supralabials invariably contacting eye. Six infralabials, except five on left side of BPBM 17451 and seven on right side of BPBM 30638. Posterior genials partially (53%) or entirely (45%) separated by single intergenial, except in BPBM 30638, which lacks an intergenial and has irregular scales behind the genials. Intergenials usually widest posteriorly (68%) but may be widest centrally (26%) or occasionally anteriorly (5%).

Dorsal scales invariably in 15-15-15 rows. Ventrals 178-198 (187 ± 7) in 18 males, 191-210 (198 ± 5) in 21 females; subcaudals 39-48 (44 ± 2) in 18 males, 26-37 (30 ± 3) in 20 females; SCR 17.5-20.7% (19.1 ± 0.9%) in males, 11.6-15.8% (13.1 ± 1.1%) in females. Tail tipped by a blunt to pointed conical spine. Maximum SVL in males 422 mm, 540 mm in females, not sexually dimorphic (t 31 = -1.4431, p = 0.0795); TLR sexually dimorphic without overlap, in males 13.3-17.9% (16.1 ± 1.3%), in females 9.2-12.9% (10.5 ± 1.0%).

Variation in head colouration is largely geographical, with yellow-spotted heads largely confined to the Huon Peninsula; the specimen from Garaina (MCZ R-152432) also has some vague pale markings on the head, which are difficult to characterize. All specimens from other areas have uniformly dark heads without yellow markings, as do six specimens from the Huon Peninsula. The Huon populations with spotted heads have a large yellow blotch on each prefrontal, a smaller one on the preocular, and a partial or complete yellow nuchal collar; they may also have yellow marks on the internasals, rostral, posterior portion of the nasal, and anterolateral portion of the parietals. Juveniles of the spotted morph (e.g., BPBM 5442, SVL = 184 mm) also have yellow colouration widely distributed on the parietals, temporals, and frontals (Fig. 4C, D ). These seem to darken and disappear with age in larger animals. In contrast, both adults and the smallest specimens of the dark-headed phenotype (BPBM 23669, SVL = 185 mm; PNGM 24716, SVL = 197 mm) have uniformly dark heads (Fig. 4E, F ), suggesting there is no ontogenetic variation in that morph.

Range.

Known from the Huon Peninsula, Morobe Province, to Karkar Island, Madang Province, in northeastern PNG to the general vicinity of Wau and Garaina in the northern Owen Stanley Range at elevations from 120-1470 m (Fig. 6B ).

Ecological notes.

We have no particular ecological information for this species but assume its habits are similar to the other species in this complex based on similar morphology and ecological conservatism in the genus.

Remarks.

Subsequent to the original description of T. nymani , the German anthropologist and scientific photographer Richard Gustav Neuhauss (1855-1915) collected an additional six specimens at Lialun - 50 km NNW of Sattelberg along the coast - that were shipped to the ZMB as part of three consignments between July 1909 and August 1910. Only the first shipment was large enough to contain these six specimens, so they must have been collected during 1909. These specimens were briefly listed by Vogt (1911), who provided locality data, and then examined by Sternfeld (1913), who provided ventral and subcaudal counts for each but erroneously referred to them as originating from Sattelberg, whence most of Neuhauss’s specimens came ( Vogt 1911). The first person to examine these specimens was certainly Vogt, who would have had the benefit of field labels in the collection jars sent by Neuhauss. It would appear that Vogt dissociated jars and labels while preparing his manuscript, and we conclude that Vogt examined Neuhauss’s six specimens with the benefit of exact locality data but leaving Sternfeld to erroneously infer that they came from Neuhauss’s main reptile collecting site. We consider all six specimens to have originated at Lialun.

These six Lialun specimens are now catalogued as ZMB 24343-44, 78770-71, and MCZ R-76627-28. In the Berlin collection, multiple specimens with the same data were chronologically assigned to numbered lots up until the Second World War. Beginning in 1991, the museum started to divide these early lots and inventory the included specimens individually. As part of this process, one specimen retained the original number and additional specimens were given a number current for that time. Thus, ZMB 24343 was originally a lot with two specimens, one of which retained the original number and the second of which was re-catalogued in April 2013 as ZMB 78770; hence, collection data for ZMB 24343 and ZMB 78770 are identical. Likewise, ZMB 24344 was a lot of two and the second specimen became ZMB 78771. Lastly, there is a note hand-written by Günther Peters from 1963 in the Berlin specimen catalogue explaining that two additional specimens (it is not clear whether both came from lot ZMB 24343, both from lot ZMB 24344, or one came from each) were exchanged in August 1963 with Ernest Williams at the MCZ. These two specimens were catalogued at the MCZ on 24 June 1964 as Pseudapistocalamus nymani (MCZ R-76627-28). The information for these two specimens in the MCZ catalogue is erroneous, however, and the locality is noted as "New Guinea: Neuhauss" with no collector given. McDowell (1969) listed these same specimens as coming from Sattelberg. Both are incorrect listings. It is clear that “Neuhauss” in this case is not a locality but refers to the collector of these specimens, and the locality is Lialun, now Morobe Province, PNG.

This species is unusual within the T. loriae Group for the presence of a distinct head colour-pattern dimorphism, with the heads of most adults from the Huon Peninsula boldly spotted with yellow in both juveniles and adults, whereas a few specimens from the Huon Peninsula and all specimens from other localities have uniformly dark heads as adults and only sparse spotting in some juveniles. Given our observations in other T. loriae Group species, we would expect these variants to represent different species, but we have found no other characters that support such a conclusion. Both morphotypes appear sympatric in at least one location (Masba Creek, Morobe Province), and although the microhabitat of these collection sites differed by 60 m in elevation, we do not consider this to be meaningful. Nonetheless, our hypothesis of conspecificity should be tested with molecular and updated morphological data when additional specimens of both forms become available.

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.