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Narophis bimaculatus (Duméril, Bibron & Duméril, 1854) Black-naped Burrowing Snake
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Holotype. MNHN-RA-0.3942, female, original type locality “Tasmanie” [= Tasmania, Australia] in error, restricted to Perth, Western Australia, fide Storr (1968: 85), obtained from J.P. Verreaux and received in Paris, France in 1844 (G. Shea, pers. comm.).

Furina bi-maculata Duméril, Bibron & Duméril 1854a, Erpétologie Générale ou Histoire Naturelle Complete des Reptiles Tome séptieme: 1240.

Furina bimaculata Duméril, Bibron & Duméril 1854b, Erpétologie Générale ou Histoire Naturelle Complete des Reptiles Tome neuvième: 376.

Brachysoma bimaculatum Günther 1858, Catalogue of Colubrine Snakes in the collection of the British Museum (Natural History): 229.

Pseudelaps bimaculatus Jan 1863, Elenco sistematico degli ofidi descritti e disegnati per l’Iconografia générale: 116.

Vermicella bimaculata Glauert 1950, A Handbook of the Snakes of Western Australia: 37.

Narophis bimaculata Worrell 1961, Western Australian Naturalist: 27.

Neelaps bimaculatus Cogger 1975, Reptiles & Amphibians of Australia: 391.

Simoselaps bimaculata Underwood 1979, Classification and Distribution of Venomous Snakes in the World: 32.

Simoselaps bimaculatus Cogger 1992, Reptiles & Amphibians of Australia: 684.

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Figure 4. Preserved (A) holotype MNHN-RA-0.3942 of Furina bi-maculata and (B) lectotype MNHN-RA-2012.411 of Furina calonotos (photos‒Muséum National ď Histoire naturelle, Paris (France) Collection: Reptiles and Amphibians (RA)).

Figure 5. Adult Narophis bimaculatus photographed in life from (A) 4 km SW of Ulyerra, Eyre Peninsula, South Australia, (B) 14 km E of Zanthus, Western Australia, WAM R112658, (C) 6 km SW of Coorow, Western Australia, WAM R135079, and (D) Cervantes, Western Australia, showing the variation in body colour and intensity of dark posterior margins (photos‒A: B. Schembri, B, C, D: B. Maryan).

Figure 6. Updated distribution maps for (A) Narophis bimaculatus and (B) Neelaps calonotos indicated by solid circles and specimens examined in the phylogenetic (coloured) and morphological (encircled) analyses. Stars indicate reliable photo or field observation. Colours within A represent intraspecific genetic variation in N. bimaculatus. Specimen locality data derived from OZCAM (2023).

Description of holotype (Fig. 4A). Scalation and proportions as in the diagnosis for the genus. Individual characteristics include: internasals much wider than long angled posteriorly; prefrontals about 1 times wide as long, medial suture slightly offline so that right prefrontal contacts left internasal; single preocular, much wider than high, in contact with nasal; two postoculars, the upper larger and lower much smaller; frontal wider anteriorly, gradually narrowing to rounded point between parietals; parietals almost equal size, right parietal slightly wider; temporals above fifth supralabials contacting both postoculars; five supralabials, first and second much the smallest, third higher in contact with orbit, fourth under orbit in contact with lower postocular and fifth very large; mental triangular between first infralabials; seven infralabials; anterior chin shields in broad contact with infralabials one to three; posterior chin shields in narrow contact with third infralabial only; six rows of intergulars between chin shields and anteriormost broad ventral.

After ≥ 167 years in preservative (Fig. 4A), the holotype has faded to white on the dorsal surface, including tip of snout and interspace between dark bands, with indistinct brown apexes extending forward as fine edge on posterior facets of scales along the body, gradually fading posteriorly to tail tip. Head and nape bands have faded to brown.

Colour in life and in preservative. The following description of colour in life is based on Figs. 5A, B, C, D and field observations of N. bimaculatus in Western and South Australia. The descriptions in preservative and of variation are based on all specimens of N. bimaculatus listed in the Appendix.

In life, variable body colour of shades of reddish brown (Figs. 5A, B), yellowish brown (Fig. 5C), orange brown or bright red (Fig. 5D), with paler interspaces on the head or similar to body colour (Figs. 5A, B, C, D), with pale cream to yellowish base or spots along the body, and dark pigment consisting of dark brown or black narrow to wide margins (almost covering ½ of scale) on the posterior facets, grading to reduced margins or smudging on tail. Dark margins varying in their intensity from clearly defined forming a reticulated pattern (Figs. 5A, B), to relatively indistinct with margins mostly restricted to apex, particularly in pale individuals (Fig. 5C). Occasionally without any indication of dark pigment on the body accentuating the pale bases or spots which tend to be larger lateroventrally (Fig. 5D, see also Storr et al. 2002: 130, Plate 30; Allen & Vogel 2019: 92, Plate RS15693‒4). Black tip on the snout present (Figs. 5A, B, D) or absent (Fig. 5C). When present extending back before posterior point of rostral often extending on to ventral edge, and lateral edge partially or narrowly covering internasals ending on nasals. Black band on head commencing on or just before anterior edge of frontal and just forward of eyes, extending back partially or narrowly covering supralabials two to five and ending on or just before posterior edge of parietals. Black band on nape transverse and 3½‒6 scales long, separated from head band by 2‒4½ pale vertebral scales. Occasionally with an indication of a third black blotch on the forebody (Fig. 5A) or reduced black band on the head (Fig. 5C). Eyes are black without discernible pupils. Ventral surface under the head and along the body, including the lateroventral edges, is cream white with glossy shine.

In preservative, body colour fades to white including interspaces on the head. Dark head and nape bands, tips on snouts (when present) and posterior margins on body scales fade to brown. The dark posterior margins on body scales can either be distinct (e.g. SAMA R59348, SAMA R61148, WAM R112153, WAM R118840, WAM R169196) or largely indistinct in the majority of specimens (e.g. SAMA R61174, SAMA R69322, WAM R60628, WAM R123596, WAM R135079), tending to be clear lateroventrally and grading to smudging on tail. The white ventral surface remains glossy.

Storr (1968: 86) noted geographic variation in colour and patterning between coastal and inland groups representing 45 specimens but his observations are incongruent with the intra-specific genetic variation observed (Fig. 2). Storr observed inland specimens differed in the dark tip on the snout being always present, larger head band, smaller nape band and darker body colour. We observed little correlation between these characters and the coastal and inland groups. Using 134 WAM specimens of N. bimaculatus (including those listed in the Appendix), we found the dark tip was present in 42% of individuals from the western and 87% from the eastern parts of range, while the tip was light in 58% of specimens from western parts of the range and only 13% in eastern parts. Five specimens had only a weak indication of a dark spot on the snout. In the South Australian material (see Appendix), a dark tip on the snout is present on all individuals except one specimen (SAMA R61174), the dark head band mostly ended on or just before posterior edge of parietals (e.g. SAMA R61147‒48) and the posterior margins on body scales varied from indistinct smudges (e.g. SAMA R61174, SAMA R69322) to distinct wide margins (e.g. SAMA R59348, SAMA R67492).

The dark head band ending behind posterior edge of parietals was present in 27% of specimens from eastern parts of range including occasional individuals with a third dark blotch on the forebody (e.g. WAM R74220, WAM R147073, WAM R165881, WAM R169106). By contrast, in the western parts of range, we recorded the dark head band typically ending on or just before posterior edge of parietals in 61% of specimens and individuals with a third blotch were rarely seen (WAM R60628, WAM R62367). We observed no appreciable variation in the nature of the nape band between coastal and inland specimens.

Our field observations suggest the degree of the darker body colour in eastern parts of range is affected by the intensity of the posterior margins on scales, and arguably influenced by the subtly different substrates inhabited by these highly fossorial snakes. Individuals from the coastal sands and plains are paler in general appearance when compared to slightly darker individuals occurring on inland sands, a phenomenon regularly observed in other fossorial reptiles (Maryan et al. 2013).

Measurements, counts, and scalation. Table 2 presents the minimum‒maximum range, means and standard deviations of the characters measured and counted (as defined in Table 1) for each sex and all adult specimens of N. bimaculatus listed in the Appendix. Sexual dimorphism is pronounced, where males have longer tails (as found previously by Clarke & How 1995) and higher subcaudal scale counts (Table 2). The widest part of the head is broader in females, indicated by broader total width, broader posterior frontal scale and longer parietal scales (Table 2, Fig. S3).

The head scale configuration in N. bimaculatus display minimal intraspecific variation. For illustrations of head scalation in N. bimaculatus, see Greer (1997: 169) and Storr et al. (2002: 190). All the specimens examined have the nasal contacting the preocular, except for two specimens (WAM R82736, WAM R125965) in which these scales are separated by the prefrontal on both sides and one specimen (SAMA R61147) on the right side. Occasional specimens have the second supralabial contacting the preocular on both sides (WAM R719, WAM R54543, WAM R104313, WAM R172498, WAM R173341). Storr (1968: 85) recorded the typical condition of the upper primary and secondary temporals fused to form a single elongate scale in 92% of specimens. We observed the same typical condition in all specimens on both sides, except in seven specimens: WAM R82736 has 1 primary + 1 secondary temporals on both sides (the secondary is much smaller), SAMA R61148 and WAM R62367 on the left side and WAM R4722, WAM R125965, WAM R135079, WAM R147076 on the right side.

Other variations of scalation include: one specimen (SAMA R61174) has divided prefrontals; one specimen (SAMA R61147) has four supralabials on the left side, caused by fusion between first and second supralabials and the third supralabial is fused with preocular; two specimens (WAM R147073, WAM R147076) have six supralabials on the right side, caused by an extra scale interposed between fourth and fifth supralabials; three specimens (WAM R65292, WAM R123596 on the right side, WAM R172498 on the left side) have one postocular, caused by either fusion of lower with fourth supralabial or upper with parietal; one specimen (WAM R165295) has the temporal fused with fifth supralabial on the right side; two specimens (WAM R125965, WAM R165295) have the temporal reduced in size on the left side, caused by the fifth supralabial contacting parietal; one specimen (WAM R135646) has a combination of both divided and undivided subcaudals and a specimen (WAM R60628) has exceptionally high ventral and vertebral scale counts of 263 and 273, respectively.

Figure 4. Preserved (A) holotype MNHN-RA-0.3942 of Furina bi-maculata and (B) lectotype MNHN-RA-2012.411 of Furina calonotos (photos‒Muséum National ď Histoire naturelle, Paris (France) Collection: Reptiles and Amphibians (RA)).

Figure 5. Adult Narophis bimaculatus photographed in life from (A) 4 km SW of Ulyerra, Eyre Peninsula, South Australia, (B) 14 km E of Zanthus, Western Australia, WAM R112658, (C) 6 km SW of Coorow, Western Australia, WAM R135079, and (D) Cervantes, Western Australia, showing the variation in body colour and intensity of dark posterior margins (photos‒A: B. Schembri, B, C, D: B. Maryan).

Figure 6. Updated distribution maps for (A) Narophis bimaculatus and (B) Neelaps calonotos indicated by solid circles and specimens examined in the phylogenetic (coloured) and morphological (encircled) analyses. Stars indicate reliable photo or field observation. Colours within A represent intraspecific genetic variation in N. bimaculatus. Specimen locality data derived from OZCAM (2023).

Distribution. Essentially unchanged from species maps of Schembri (2017: 26) and Chapple et al. (2019: 537), N. bimaculatus is widespread throughout the temperate, semiarid to arid areas of southern Western Australia and southwestern South Australia (Fig. 6A). In Western Australia, extends north to Denham on the Peron Peninsula at Shark Bay (WAM Photo Database), 22 km SSW of Woodleigh Homestead, 38 km SE of Nerren Nerren Homestead, 29 km SSE of Yalgoo (B. Budrey, pers. comm.) and Lakeside at 48 km SW of Cue, east to 7‒8 km WNW of Point Salvation, Tropicana Mine and 14 km E of Zanthus, and south to Bunbury, Mount Saddleback, Williams, 6 km ENE of Lake Cronin and Peak Charles National Park. There is a disjunct population on the North West Cape, represented by a single specimen (WAM R116665) from 2 km W of Bullara Homestead (Maryan 1995; Fig. 6A). Also on Dirk Hartog Island. In South Australia, extends north to 41 km NNW of Maralinga, Ooldea and Kingoonya, east to Secret Rocks, and south to 4 km SW of Ulyerra on the Eyre Peninsula (Schembri 2017).

The apparent allopatric distribution of this species in the Great Victoria Desert of Western and South Australia (Fig. 6A), with collection records indicating a distance of ~ 688 km between populations, is possibly an artefact of collecting in a poorly surveyed remote region.

Habitat and ecology. Narophis bimaculatus occupies a variety of temperate, semiarid to arid vegetation that grow on sandy to loamy soils, including coastal sand dunes and swales, sandplains, dry sclerophyll woodlands of Banksia L.f., mallee and/or other Eucalyptus L՛Hér., heathlands particularly coastal Acacia Mill. shrublands and inland hummock grassland dominated sand dunes and plains (Wilson & Knowles 1988; Ehmann 1992; How & Shine 1999; Bush et al. 2007, 2010; Schembri 2017; Chapple et al. 2019; Wilson & Swan 2021). At the northern extent of its range, the habitat at 2 km W of Bullara Homestead on the North West Cape consists of a red Triodia R.Br.-covered sand dune with sparse shrubs (B. Maryan, pers. obs.).

Field observations indicate a preference for sandy soils supporting low open vegetation, and a scarcity of observations on harder substrates such as the clay and granitic soils on the Darling Range (Storr et al. 1978; Bush et al. 2010). This preference is exemplified by 156 WAM collection records of which the majority are recorded from the Swan Coastal Plain, Great Victoria Desert and Geraldton Sandplains bioregions (OZCAM 2023). In these vegetation associations, specimens of N. bimaculatus particularly during cooler weather can be raked from the upper layers of soil beneath leaflitter at base of trees and shrubs, piles of dead vegetation, abandoned stick-ant Iridomyrmex conifer Forel, 1902 nests, embedded, decaying logs and stumps and spoil-heaps, consisting of soil, branches and other debris pushed up, usually by machinery, on the side of tracks and firebreaks. Field observations of N. bimaculatus on the surface beneath cover are scarce, apart from one specimen (WAM R64822) found on soil under log.

Additionally, at peak seasonal activity, N. bimaculatus can be funnel or pit-trapped in buckets/pipes and nocturnally observed while driving on roads and tracks (How & Shine 1999; Thompson & Thompson 2007; B. Maryan & B. Bush, pers. obs.). Regarding the usefulness of traps in catching small, fossorial snakes, it is generally considered that capture rates are low for snakes poorly suited to moving long distances across open areas (How & Shine 1999). However, the technique of raking can be productive, allowing targeting of particular microhabitats such as spoil-heaps, when on occasions up to 10‒15 individuals of B. semifasciatus (Günther, 1863), N. bimaculatus, N. calonotos and S. bertholdi (Jan, 1859) can be found in a single day on the Swan Coastal Plain (Maryan 2002; Bush et al. 2007: 33).

Sympatry or close parapatry with other fossorial species. Recorded instances of narrow or broad sympatry, occasionally syntopy, involving N. bimaculatus include with B. approximans (Glauert, 1954) at the southern extremities of its range, B. fasciolatus (Günther, 1872), B. semifasciatus, and S. bertholdi throughout a large area encompassing several bioregions in Western and South Australia, N. calonotos on the Swan Coastal Plain and a small area of the Geraldton Sandplains bioregions and S. littoralis (Storr, 1968) on the west coast between Shark Bay, including Dirk Hartog Island and Cervantes (Chapman & Dell 1985; Storr et al. 2002; Thompson & Thompson 2006; Maryan 2005; Bush et al. 2007, 2010; B. Maryan, pers. obs.). At Bold Park in Western Australia, B. fasciolatus, B. semifasciatus, N. bimaculatus, N. calonotos, and S. bertholdi have all been recorded (How 1998; How & Shine 1999). Based on collection records, B. approximans, S. bertholdi, and S. littoralis are recorded on the North West Cape (Storr et al. 2002). Collection records also indicate N. bimaculatus and S. anomalus occur in sympatry at 7‒8 km WNW of Point Salvation in the Great Victoria Desert bioregion. At the boundary between the eastern Eyre Yorke Block and Gawler bioregions in South Australia, collection records suggest N. bimaculatus and V. annulata (Gray, 1841) occur in close parapatry in the vicinities of Secret Rocks (see Appendix) and Port Augusta (SAMA R2173).

Comparisons with other fossorial species. Diagnostic differences between N. bimaculatus and N. calonotos are listed under the revised diagnosis for Narophis. Narophis bimaculatus will be compared with B. approximans, B. fasciolatus, B. semifasciatus, S. anomalus, S. bertholdi, S. littoralis, and V. annulata using Storr et al. (2002) and Wilson & Swan (2021) with which it occurs in sympatry or close parapatry (see above).

It differs from the listed Brachyurophis species in: larger adult total length to 468 mm (versus to 390 mm), more slender body form (versus moderately robust), higher ventral scale counts of 178‒234 (versus ˂ 190), typically one elongate temporal scale (versus typically 1 + 1, but often fused in B. semifasciatus), 15 midbody scale rows (versus 17), protrusive round-shaped snout without cutting edge (versus protrusive wedge-shaped snout tipped with transverse weak to strong cutting edge) and without bands along the body (versus cross-banded along the body, but bands ragged-edged in B. fasciolatus).

It differs from the listed Simoselaps species in: larger adult total length to 468 mm (versus to 390 mm), more slender body form (versus moderately robust), higher ventral scale counts of 178‒234 (versus ˂ 135), typically one elongate temporal scale (versus typically 1 + 1, but primary temporal fused with second last supralabial in S. anomalus and primary temporal often large contacting oral margin in S. littoralis) and without bands along the body (versus encircled by bands along the body).

It differs from V. annulata in: smaller adult total length to 468 mm (versus to 760 mm), lower ventral scale counts of 178‒234 (versus 195‒257), typically one elongate temporal scale (versus 1 + 1, but primary temporal occasionally fused with second last supralabial), protrusive round-shaped snout (versus bluntly round-shaped snout) and without bands along the body (versus encircled by bands along the body).

Narophis bimaculatus shares a superficial resemblance to the similarly coloured and patterned Furina ornata (Gray, 1842) but differs in numerous ways including the protrusive snout (not protrusive in Furina), black head band extends forward to level of eyes (black head band extends forward to snout in Furina) and one elongate temporal scale (2 + 2 in Furina).

Remarks. Strahan et al. (1998) and How & Shine (1999) present information on the ecology, reproductive biology and diet of N. bimaculatus. In Western Australia, the majority of collection records of N. bimaculatus are from the heavily-cleared and modified Swan Coastal Plain that includes the metropolis of Perth; however this species persists in several remnant bushland areas set aside for the conservation of flora and fauna (e.g. How & Dell 1994; How 1998). Due mainly to paucity of records in South Australia, N. bimaculatus has a conservation status of Near Threatened and is listed as Rare under state legislation, which is likely due to a combination of remoteness of suitable habitat and the cryptic, fossorial nature of the species (Schembri 2017). In general, N. bimaculatus is considered to be Least Concern under the IUCN conservation status owing to its widespread distribution without any perceived threats, occurring in many large, relatively undisturbed nature reserves and national parks (Chapple et al. 2019: 537). We note the distinction between categories Least Concern and Data Deficient referring to “inadequate information” is unclear (International Union for Conservation of Nature 2021), especially when this lack of knowledge of population size and structure, ecology, threats and threat amelioration can be equally applied to many Least Concern listed species such as N. bimaculatus.