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White-lipped snake

The white-lipped snake (Drysdalia coronoides) is a small, slender species of venomous elapid snake endemic to south-eastern mainland Australia and Tasmania, notable for its distinctive white stripe along the upper lip bordered by black, and its adaptation to cold climates as one of Australia's most cold-tolerant reptiles.^[1]^[2] Typically measuring 25–50 cm in length, adults exhibit a body color ranging from dark olive green or green-grey to brown or black, with a pale grey or pinkish underside; juveniles often have a darker appearance with an orange belly.^[1]^[2] The snake's head is narrow and rounded, and its tail tapers to a fine point, contributing to its agile, secretive movement.^[1] It inhabits a variety of environments including heaths, grasslands, open woodlands, sclerophyll forests, from sea level up to 1,300 m elevation, and is particularly active in sunnier spots despite its preference for cooler, southern latitudes.^[1]^[2] Distribution spans across Tasmania and south-eastern states including South Australia, Victoria, and New South Wales, where it shelters under rocks, logs, or debris during inactive periods.^[1]^[2] Active during the day and at night but capable of foraging on mild winter days, the white-lipped snake is a shy predator that primarily feeds on small skinks and their eggs, occasionally consuming frogs.^[1]^[2] It is viviparous, giving birth to 2–10 live young in late summer (March–April) after ovulation in spring, with individuals reaching maturity around three years of age and potentially breeding biennially.^[1] Though venomous with small fangs and glands, its toxin is weakly potent and poses little risk of serious injury to healthy adults, though bites should receive medical attention due to potential sensitivity in some individuals.^[1] Recent proteomic studies have identified novel proteins in its venom, including nicotinic acetylcholine receptor antagonists, highlighting its biochemical complexity despite low human hazard.^[3]^[4] Classified as Least Concern on the IUCN Red List as of the 2018 assessment, the species maintains stable populations with no major threats identified, though habitat preservation in its highland and coastal ranges supports its persistence.^[2]^[5]

Taxonomy

Classification

The white-lipped snake, Drysdalia coronoides, belongs to the family Elapidae, subfamily Hydrophiinae, within the order Squamata.^[6] This placement reflects its status as a venomous elapid snake native to southeastern Australia, characterized by front-fanged dentition typical of the Hydrophiinae.^[7] Originally described by Albert Günther in 1858 as Hoplocephalus coronoides based on specimens from southeastern Australia, the species was later reassigned to the genus Drysdalia.^[8] The genus Drysdalia was established by Eric Worrell in 1961 to group small, terrestrial elapid snakes with similar cranial and dental features, separating them from broader genera like Hoplocephalus.^[9] A key taxonomic revision by A.J. Coventry and P.A. Rawlinson in 1980 confirmed D. coronoides as a distinct species and delineated the genus to include three recognized species: D. coronoides (white-lipped snake), D. mastersii (Masters' snake), and D. rhodogaster (mustard-bellied snake), based on morphological examinations of scale patterns, hemipenial structure, and geographic distribution.^[10] Phylogenetically, D. coronoides forms part of a clade of small, viviparous Australian elapids, with molecular analyses using mitochondrial genes such as cytochrome b indicating close relationships among Drysdalia species, distinguished by sequence divergences of 5-10% in key markers.^[11] Within D. coronoides, studies reveal at least two highly divergent lineages, particularly between Tasmanian and mainland populations, with divergence estimated at 2-3 million years ago during the Plio-Pleistocene, supported by ND4 and 16S rRNA gene data.^[12] Broader elapid phylogenies position Drysdalia near genera like Elapognathus, forming a basal group among terrestrial hydrophiines, as evidenced by bootstrap-supported trees from multi-gene datasets.^[13]

Etymology

The white-lipped snake, scientifically known as Drysdalia coronoides, derives its common name from the distinctive white coloration of its supralabial scales, which form a prominent pale stripe along the upper lip, often bordered above by a narrow black line.^[14]^[15] The species was first described in 1858 by Albert Günther as Hoplocephalus coronoides in his Catalogue of Colubrine Snakes in the Collection of the British Museum, based on specimens from southeastern Australia held in the museum's collection. The specific epithet "coronoides" originates from the Greek words korōnē (κόρωνη), meaning wreath, crown, or diadem, and -eidēs (-εἰδής), meaning resembling or like; it alludes to the snake's head stripes, which Günther noted as resembling a regal diadem or head wreath.^[6]^[16] The genus Drysdalia was established in 1961 by Eric Worrell to accommodate this and related small elapid snakes from southern Australia.^[17] It honors the Australian artist George Russell Drysdale (1912–1981), who accompanied Worrell on collecting expeditions to Tasmanian islands where specimens were obtained.^[16]

Description

Physical characteristics

The white-lipped snake (Drysdalia coronoides) is a small, slender-bodied elapid with smooth dorsal scales arranged in 15 rows at midbody, rarely 17.^[17] Adults typically measure 30-40 cm in total length, with a maximum recorded of approximately 50 cm, consisting of a snout-vent length of up to 40 cm and a tail comprising about 20% of the total length.^[17] The body tapers gently toward the tail, which ends in 38-67 undivided subcaudal scales.^[17] The head is slightly distinct from the neck, featuring 6 supralabial scales, with the third and fourth entering the eye, and 2+2 temporals.^[17] An undivided anal scale precedes the subcaudals.^[17] Ventral scales number 123-152 and are smooth and unkeeled.^[17] Sexual dimorphism is minimal in overall body size, with adult snout-vent lengths averaging around 29 cm in males and 30 cm in females, though males possess significantly longer tails relative to body size than females.^[18] This species' compact size represents a key anatomical adaptation to cold climates, enabling rapid thermoregulation by allowing the snake to warm up quickly during brief periods of suitable weather.^[1]

Coloration and variation

The white-lipped snake exhibits a dorsal coloration that ranges from olive-green to dark brown in adults, while juveniles are typically black. This pattern often includes faint darker bands or spots, particularly more pronounced in younger individuals.^[17] The ventral surface is pale, ranging from salmon pink to orange, providing a subtle contrast to the darker dorsum.^[17]^[19] A defining feature is the prominent white or cream-colored upper lips (supralabials), which form a thin stripe edged above by a narrow black line, sharply contrasting with the dark head and giving the species its common name.^[17]^[1] Occasionally, a second white stripe may appear from the upper temporal area to the neck.^[17] Geographic variation occurs across its range, with Tasmanian populations displaying duller, darker tones—often deep olive-green to green-grey dorsally—likely aiding camouflage in cooler, wetter environments, compared to slightly lighter mainland forms that may include brick-red or grey hues.^[17]^[1]^[19] Ontogenetic changes are evident as the snake matures, with juvenile patterns fading: the black dorsal coloration lightens to olive-green or brown, and the bright orange ventral surface may shift toward paler pink or grey, reducing the distinctiveness of early banding.^[17]^[1]

Distribution and habitat

Geographic range

The white-lipped snake (Drysdalia coronoides) is endemic to southeastern Australia, with its distribution extending from coastal regions of New South Wales southward through the Australian Capital Territory and Victoria to Tasmania.^[2]^[8] The species is notably absent from the arid interior of the continent, reflecting its adaptation to more temperate and mesic environments, but it occupies highland areas up to approximately 1,300 m in Tasmania and higher in mainland Australia, including sites up to nearly 2,000 m in the Snowy Mountains.^[1]^[20]^[21] In Tasmania, the snake is widespread across the island, including offshore populations on islands such as King Island, where it contributes to local reptile diversity.^[1]^[22] Its historical range has shown stability, with no evidence of major contractions prior to the 1900s based on specimen records and early surveys; contemporary distribution is supported by over 2,400 occurrence records from citizen science platforms.^[20]^[23] Within its range, D. coronoides overlaps sympatrically with the congener Drysdalia nasuta in portions of Victoria, where both species inhabit similar southeastern habitats.^[9]^[24]

Habitat preferences

The white-lipped snake (Drysdalia coronoides) primarily inhabits moist temperate forests, woodlands, and heaths, favoring environments with higher humidity and avoiding drier sclerophyll woodlands.^[18]^[1]^[19] These snakes are commonly associated with wetter habitats near water sources and grassy areas, where they utilize rocky outcrops, logs, and leaf litter for shelter and cover.^[24]^[1] Their elevational range extends from sea level to subalpine zones up to approximately 1,300 m in Tasmania and higher on the mainland, allowing occupancy in diverse topographic settings across southeastern Australia.^[1]^[19] Seasonally, these snakes shift habitat use, frequenting aquatic margins and grassy clearings more during summer for foraging, while retreating to upland refugia in winter.^[24]^[1] They exhibit notable cold tolerance, remaining active at temperatures as low as 10°C and foraging on fine winter days due to their small size enabling rapid thermoregulation, making them one of Australia's most cold-adapted snake species.^[18]^[1]^[19] The species often occurs in proximity to skink populations in grassy clearings, reflecting a close ecological association driven by its primary diet of small lizards.^[18]^[1] This distribution in open, prey-rich microhabitats enhances their camouflage through subtle coloration patterns that blend with leaf litter and grasses.^[19]

Behavior and ecology

Activity patterns

The white-lipped snake (Drysdalia coronoides) is primarily diurnal, though it can forage nocturnally, particularly in the evenings, and remains active on mild winter days.^[18]^[1] In temperate regions of its range, it undergoes brumation during winter months, typically from May to September, seeking shelter in burrows, under rocks, logs, or debris to conserve energy amid low temperatures.^[18]^[24] Emergence often follows warming trends, with increased activity post-frost as ambient conditions improve. During active seasons, morning basking aids thermoregulation, particularly in cooler climates where the snake maintains body temperatures around 24–25°C through behavioral adjustments. Heavy rainfall can suppress surface activity, prompting individuals to retreat to cover, though the species tolerates moist environments well.^[18]^[1] Defensive behaviors emphasize evasion over confrontation; when disturbed, the snake tends to hide or flee to nearby cover, though it may assume a defensive posture if cornered.^[1]

Diet and foraging

The white-lipped snake (Drysdalia coronoides) is primarily saurophagous, with approximately 90% of its diet consisting of scincid lizards, particularly small skinks such as those in the genera Lerista (e.g., southeastern sliders in mainland habitats) and Niveoscincus (e.g., in Tasmanian grasslands).^[25] It occasionally preys on frogs, which comprise a minor portion of its overall diet.^[1] These snakes employ a foraging strategy that combines active searching through vegetation, leaf litter, and ground debris with ambush predation, often waiting under cover like rocks, logs, or soil crevices before striking at detected prey.^[24]^[1] Upon envenomating the prey with a quick bite, the snake typically releases it briefly to allow the venom to take effect before tracking and consuming the immobilized lizard or frog head-first.^[25] This method is effective for subduing small, agile prey relative to the snake's slender body, which rarely exceeds 40 cm in total length.^[24] Foraging activity occurs both diurnally and nocturnally, with peaks during warmer months when skinks are more active, though the species' small size enables hunting on mild winter days in cooler regions like Tasmania.^[25] Juveniles focus on smaller lizards to match their limited gape size, while adults target slightly larger individuals, contributing to population control of skinks in grassland and forested ecosystems.^[1]^[24]

Reproduction and life cycle

The white-lipped snake (Drysdalia coronoides) is viviparous, giving birth to live young after internal development of embryos within the female's oviducts.^[18] Ovulation typically occurs in spring (September to November in the Southern Hemisphere), with mating behaviors observed during this period and sometimes extending into autumn.^[18] Males engage in combat rituals to compete for females, involving body twisting and occasional biting while keeping heads raised apart, a common trait in elapid snakes that promotes sexual selection for larger body size.^[18] Gestation lasts approximately 3-4 months, culminating in parturition during late summer to early autumn (January to April).^[18] Litters consist of 2-10 neonates, with an average of 3-5 offspring; larger females tend to produce more young, though fecundity is lower in colder regions like Tasmania where females may reproduce only every 2-3 years.^[18]^[1] Neonates measure 8-12 cm in total length at birth and are fully independent immediately, receiving no parental care from either parent.^[1]^[18] Juveniles exhibit slow growth rates adapted to cold climates, reaching sexual maturity at 2-3 years of age when they attain approximately 20-25 cm in length.^[18]^[1] In the wild, the lifespan is estimated at up to 10 years, though high juvenile mortality from predation contributes to lower overall survival rates.^[26]^[18]

Venom and interactions

Venom composition

The white-lipped snake (Drysdalia coronoides), like other elapids, possesses proteroglyphous fangs—fixed, anteriorly positioned, hollow structures that deliver venom efficiently during envenomation. The venom is primarily neurotoxic, dominated by three-finger toxins (3FTxs) that act as antagonists of nicotinic acetylcholine receptors, inducing paralysis in prey. These include short-chain α-neurotoxins (Type I), long-chain α-neurotoxins (Type II), and κ-bungarotoxins (Type III), with novel isoforms exhibiting unique cysteine arrangements and rapid evolution under positive selection in surface loops for prey-specific adaptations. Myotoxic components, such as phospholipases A2 (PLA₂s), contribute to tissue damage and are present alongside other proteins like snake venom metalloproteases (SVMPs), venom nerve growth factors, serine protease inhibitors, cysteine-rich secretory proteins, vespryns, and a novel phospholipase B (PLB) family.^[27]^[28]^[27] Venom yield is low, typically 2–3 mg per extraction, reflecting the snake's small size, yet it remains potent relative to body mass.^[4] Evolutionary adaptations have simplified the venom composition for targeting small lizards and frogs, differing from the more complex, procoagulant-heavy venoms of larger Australian elapids; this includes functional conservation in 3FTxs despite structural variations like cysteine deletions.^[28]^[27]^[4]

Bites and medical significance

Bites from the white-lipped snake (Drysdalia coronoides) are rare due to the species' shy and reclusive nature, with fewer than 10 documented human cases reported since 1900.^[29] The snake's small size (typically 20-50 cm) and small fangs further reduce the likelihood of significant envenomation in adults, though children, the elderly, or those with compromised health may experience more pronounced effects.^[1] In the limited recorded human envenomations, symptoms typically include local pain and swelling at the bite site, progressing to mild systemic effects such as ptosis (drooping eyelids), nausea, vomiting, abdominal pain, and diarrhea.^[29] Elevated serum creatine kinase (CK) levels, sometimes exceeding 50,000 U/L, and myogloburia have been observed, indicating myotoxic effects, but no coagulopathy or severe neurotoxicity has been reported.^[30] One notable case from 1994 involved facial weakness and ptosis developing 18 hours post-bite, with a serum CK of 57,300 U/L, but the patient made a full recovery without long-term complications.^[30] No human fatalities have been attributed to white-lipped snake bites.^[29] First aid for suspected bites follows standard protocols for Australian elapids: apply a firm pressure immobilization bandage over the bite site and distal to it, immobilize the affected limb, and seek immediate medical attention without inducing panic or unnecessary movement.^[31] In hospital settings, monitoring for myotoxicity and neurotoxic signs is essential; no species-specific antivenom exists, but tiger snake antivenom (1-2 vials) has proven effective in neutralizing effects, as demonstrated in documented cases with full recovery within 48 hours.^[29]^[30] From a public health perspective, the white-lipped snake poses low risk in its range (Tasmania, Victoria, and southeastern New South Wales), with envenomations far less common than those from more aggressive species like the tiger snake.^[1] However, misidentification with deadlier sympatric snakes can lead to overtreatment, emphasizing the need for expert identification in affected areas.

Conservation status

Population trends

The white-lipped snake (Drysdalia coronoides) maintains a stable population trend across its range in south-eastern Australia and Tasmania, with no indications of widespread decline. Classified as Least Concern by the IUCN Red List (as assessed in 2017), the species is locally common in appropriate habitats, particularly alpine and subalpine regions where it represents the dominant snake taxon.^[2]^[32] Population monitoring relies on herpetological field surveys and citizen science contributions, such as those compiled on iNaturalist, which document thousands of observations supporting consistent abundance over time. Australian government datasets, including the Atlas of Living Australia, reveal steady sighting records from the 2000s through the 2020s, with over 2,400 occurrences reported, underscoring the species' persistence without quantitative evidence of reduction.^[15] Regional variations show stability across its range, including in Tasmanian highland populations where the snake remains widespread, based on distributional data from state biodiversity assessments.^[33]

Threats and protection

The white-lipped snake (Drysdalia coronoides) faces limited major threats across its range in south-eastern Australia and Tasmania, primarily due to its adaptable nature and widespread distribution in varied habitats from coastal heaths to alpine areas. Predation by introduced species represents the most significant localized pressure, particularly in Tasmania where domestic and feral cats (Felis catus) actively prey on these small snakes, contributing to mortality rates.^[1] Additionally, the introduced laughing kookaburra (Dacelo novaeguineae) imposes a heavy toll on populations in Tasmania by targeting snakes as prey, exacerbating declines in areas with high predator densities.^[1] Habitat fragmentation from urban expansion and agriculture may indirectly affect some populations by reducing shelter sites under rocks, logs, and debris, though the species' cold tolerance and nocturnal flexibility mitigate broader impacts.^[24] Conservation status for the white-lipped snake is favorable, with the species assessed as Least Concern on the IUCN Red List (as assessed in 2017), indicating no immediate risk of extinction and stable populations overall.^[5] It is not listed under Australia's national Environment Protection and Biodiversity Conservation Act 1999 (EPBC Act) nor on state threatened species lists such as Victoria's Flora and Fauna Guarantee Act 1988, where it is categorized as common.^[2] In Tasmania, populations are considered secure, with no specific decline trends documented.^[1] Protection measures are general rather than species-specific, relying on broader wildlife legislation that prohibits harm, capture, or trade without permits. Under Victoria's Wildlife Act 1975, the snake is fully protected as a native reptile, with rehabilitation restricted to authorized facilities to ensure ethical handling and release into suitable habitats above 20°C.^[24] Similar safeguards apply in New South Wales via the Biodiversity Conservation Act 2016, covering all native reptiles against illegal killing or disturbance.^[34] In Tasmania, while no targeted programs exist, ongoing management of invasive predators like cats and kookaburras through control efforts indirectly supports populations.^[1] Habitat preservation in reserves and promotion of road awareness to reduce vehicle strikes further aid long-term viability, though no dedicated recovery plans are in place due to the species' secure status.^[24]

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