''Toxicocalamus'' is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

snake Snakes are elongated, Limbless vertebrate, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Ma ...

s in the

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

Elapidae Elapidae (, commonly known as elapids ; grc, ἔλλοψ ''éllops'' "sea-fish") is a family of snakes characterized by their permanently erect fangs at the front of the mouth. Most elapids are venomous, with the exception of the genus Emydoce ...

. The genus is

endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...

New Guinea New Guinea (; Hiri Motu Hiri Motu, also known as Police Motu, Pidgin Motu, or just Hiri, is a language of Papua New Guinea, which is spoken in surrounding areas of Port Moresby (Capital of Papua New Guinea). It is a simplified version of ...

Description

Most species of ''Toxicocalamus'' are relatively small, the largest specimen known being the

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...

of the recently described ''Toxicocalamus ernstmayri'', which measures snout-to-vent length (SVL) and in total length (TTL). The second longest is the holotype of ''T. grandis'', which measures SVL, TTL. Most species are under TTL, and several are the thickness of bootlaces. In general females have longer bodies than males, but much shorter tails.

Venom

Members of genus ''Toxicocalamus'' are venomous, with fixed front-fangs (a dental arrangement known as

proteroglyphous A snake skeleton consists primarily of the skull, vertebrae, and ribs, with only vestigial limb, vestigial remnants of the limbs. Skull The skull of a snake is a very complex structure, with numerous joints to allow the snake to swallow prey fa ...

), but are not known to be a threat to humans, being unaggressive, of modest size, and secretive. However, the venom of ''T. longissimus'' is believed to be fairly toxic, since it contains

three-finger toxin Three-finger toxins (abbreviated 3FTx) are a protein superfamily of small toxin proteins found in the venom of snakes. Three-finger toxins are in turn members of a larger superfamily of three-finger protein domains which includes non-toxic prote ...

s (3FTx), Type-I phospholipase A (PLA) and snake venom

metalloproteinase A metalloproteinase, or metalloprotease, is any protease enzyme whose catalytic mechanism involves a metal. An example is ADAM12 which plays a significant role in the fusion of muscle cells during embryo development, in a process known as myo ...

(SVMP), while ''T. buergersi'' possesses long venom glands than extend backwards into the body cavity.

Behaviour

Although most species of ''Toxicocalamous'' are believed to be diurnal, they are fossorial, or semi-fossorial, in habit and rarely encountered. O'Shea, M. (1996). ''A Guide to the Snakes of Papua New Guinea''. Port Moresby, Papua New Guinea: Independent Publishing. vii + 239 pp. ..

Geographic range

Many species of ''Toxicocalamus'' are localised in their distribution and associated with particular islands or mountain ranges. Several species are poorly known, with four known only from their holotypes. ''Toxicocalamus'' is probably not closely related to the Australian Elapidae, being endemic to the island of

, northern coastal offshore islands, i.e. Seleo Is. (

Sandaun Province Sandaun Province (formerly West Sepik Province) is the northwesternmost mainland province of Papua New Guinea. It covers an area of 35,920 km2 (13868 m2) and has a population of 248,411 (2011 census). The capital is Vanimo. In July 1998 the a ...

, PNG); Walis Is. and Tarawai Is. (

East Sepik Province East Sepik is a province in Papua New Guinea. Its capital is Wewak. East Sepik has an estimated population of 433,481 people (2010 census) and is 43,426 km square in size. History Cherubim Dambui was appointed as East Sepik's first premier b ...

, PNG), and Karkar Is. (

Madang Province Madang is a province of Papua New Guinea. The province is on the northern coast of mainland Papua New Guinea and has many of the country's highest peaks, active volcanoes and its biggest mix of languages. The capital is the town of Madang. ...

, PNG), and the archipelagoes of

Milne Bay Province Milne Bay is a province of Papua New Guinea. Its capital is Alotau. The province covers 14,345 km² of land and 252,990 km² of sea, within the province there are more than 600 islands, about 160 of which are inhabited. The province has a ...

to the southeast, i.e.

d'Entrecasteaux Archipelago D'Entrecasteaux Islands () are situated near the eastern tip of New Guinea in the Solomon Sea in Milne Bay Province of Papua New Guinea. The group spans a distance of , has a total land area of approximately and is separated from the Papua New G ...

( Goodenough Is., Fergusson Is., and Normanby Is.), Woodlark Is., and the

Louisiade Archipelago The Louisiade Archipelago is a string of ten larger volcanic islands frequently fringed by coral reefs, and 90 smaller coral islands in Papua New Guinea. It is located 200 km southeast of New Guinea, stretching over more than and spread ...

( Misima Is., Sudest Is., and Rossel Is.).

Diet

The

prey Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...

of snakes in the genus ''Toxicocalamus'' appears to consist almost entirely of

earthworm An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. Th ...

s, particularly the giant earthworms of the

Megascolecidae The Megascolecidae is a taxonomic family of earthworms which is native to Madagascar, Australia, New Zealand and both South East Asia and North America. All species of Megascolecidae belong to the Clitellata class. Megascolecidae are a large fami ...

, hence the adoption of the term "worm-eating snakes" for species within this genus.

Reproduction

In common with other tropical

elapids Elapidae (, commonly known as elapids ; grc, ἔλλοψ ''éllops'' "sea-fish") is a family of snakes characterized by their permanently erect fangs at the front of the mouth. Most elapids are venomous, with the exception of the genus Emydocep ...

, ''Toxicocalamus'' is believed to reproduce by

oviparity Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), and m ...

, with clutch sizes of 3–7 recorded, dependent on species and size of the female.

Natural history

The natural history of many species of ''Toxicocalamus'' is almost entirely undocumented, due to a paucity of specimens and the infrequence of their encounter in the field.

Species

The following 17 species, one of which has two

subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...

, are currently recognised as being valid in the genus ''Toxicocalamus'': * '' Toxicocalamus buergersi'' ( Sternfeld, 1913) (in German). – Buergers' forest snake,

Torricelli Mountains The Torricelli Mountains are a mountain range in Sandaun Province, north-western Papua New Guinea. The highest peak in the range is Mount Sulen at 1650 meters. The Bewani Mountains are located to the west, and the Prince Alexander Mountains ar ...

snake * '' Toxicocalamus cratermontanus''

Kraus Kraus is a German surname meaning "curly". Notable people with the surname include: * Adalbert Kraus (born 1937), German tenor * Adam Kraus (born 1984), American footballer * Adolf Kraus (1850–1928), lawyer and Jewish leader * Adolph Robert ...

, 2017 – Crater Mountain snake * '' Toxicocalamus ernstmayri''

O'Shea O'Shea is a surname and, less often, a given name. It is an anglicized form of the Irish patronymic name Ó Séaghdha or Ó Sé, originating in the Kingdom of Corcu Duibne in County Kerry. Notable people with the name include: Surname *Alicia ...

Parker Parker may refer to: Persons * Parker (given name) * Parker (surname) Places Place names in the United States *Parker, Arizona *Parker, Colorado * Parker, Florida * Parker, Idaho * Parker, Kansas * Parker, Missouri * Parker, North Carolina *Park ...

Kaiser ''Kaiser'' is the German word for "emperor" (female Kaiserin). In general, the German title in principle applies to rulers anywhere in the world above the rank of king (''König''). In English, the (untranslated) word ''Kaiser'' is mainly ap ...

, 2015 –

Star Mountains The Star Mountains ( Dutch ( colonial)'': Sterrengebergte''; Indonesian'': Pegunungan Bintang'') are a mountain range in western Papua New Guinea and the eastern end of Highland Papua, Indonesia, stretching from the eastern end of Indonesia t ...

snake, Star Mountains worm-eating snake * '' Toxicocalamus goodenoughensis'' J.R. Roberts & C. Austin, 2020 * '' Toxicocalamus grandis'' ( Boulenger, 1914) – Setakwa River snake, Setekwa River forest snake * '' Toxicocalamus holopelturus'' McDowell, 1969 – Mt. Rossel forest snake,

Rossel Island Rossel Island (named after de Rossel, a senior officer on the French expedition of d'Entrecasteaux, 1791-1793; also known as Yela) is the easternmost island of the Louisiade Archipelago, within the Milne Bay Province of Papua New Guinea. Tree Isl ...

snake * '' Toxicocalamus longissimus'' Boulenger, 1896 – Fergusson Island forest snake, Woodlark forest snake,

Woodlark Island Woodlark Island, known to its inhabitants simply as Woodlark or Muyua, is the main island of the Woodlark Islands archipelago, located in Milne Bay Province and the Solomon Sea, Papua New Guinea. Although no formal census has been conducted sinc ...

snake * '' Toxicocalamus loriae'' (Boulenger, 1898) – common worm-eating snake, Loria forest snake *'' Toxicocalamus mattisoni'' Kraus, 2020 * '' Toxicocalamus mintoni'' Kraus, 2009 –

Minton's Minton's Playhouse is a jazz club and bar located on the first floor of the Cecil Hotel at 210 West 118th Street in Harlem, Manhattan, New York City. It is a registered trademark of Housing and Services, Inc. a New York City nonprofit provider ...

forest snake, Sudest Island snake * '' Toxicocalamus misimae'' McDowell, 1969 – Misima Island forest snake,

Misima Island Misima (formerly called St. Aignan) is a volcanic island in the northwest of Louisiade Archipelago within Milne Bay Province of Papua New Guinea. History Misima Island was inhabited by Austronesians since about 1500 BC. The island was sig ...

snake * '' Toxicocalamus nigrescens'' Kraus, 2017 –

Fergusson Island Fergusson Island is the largest island of the D'Entrecasteaux Islands, in Papua New Guinea. It has an area of , and mostly consists of mountainous regions, covered by rain forests. There are three large volcanoes on the island. Fergusson Island ...

worm-eating snake * '' Toxicocalamus pachysomus'' Kraus, 2009 – Cloudy Mountains worm-eating snake * '' Toxicocalamus preussi'' (Sternfeld, 1913) – Preuss's forest snake, Preuss' slender worm-eating snake :* '' Toxicocalamus preussi preussi'' (Sternfeld, 1913) – Preuss's Sepek forest snake, Preuss' slender worm-eating snake :* '' Toxicocalamus preussi angusticinctus'' Bogert & Matalas, 1945 –

Fly River The Fly River is the third longest river in the island of New Guinea, after the Sepik River and Mamberamo River, with a total length of and the largest by volume of discharge in Oceania, the largest in the world without a single dam in its cat ...

forest snake, Fly River slender worm-eating snake * '' Toxicocalamus pumehanae'' O'Shea,

Allison Allison may refer to: People * Allison (given name) * Allison (surname) (includes a list of people with this name) * Eugene Allison Smith (1922-1980), American politician and farmer Companies * Allison Engine Company, American aircraft engine ...

& Kaiser, 2018 – Managalas Plateau snake * '' Toxicocalamus spilolepidotus'' McDowell, 1969 – Krakte Mountains spotted snake, spotted forest snake * '' Toxicocalamus stanleyanus'' Boulenger, 1903 –

Owen Stanley Mountains Owen Stanley Range is the south-eastern part of the central mountain-chain in Papua New Guinea. Its highest point is Mount Victoria at , while its most prominent peak is Mount Suckling. History Owen Stanley Range was seen in 1849 by Captain Ow ...

snake, Owen Stanley Range forest snake ''

Nota bene (, or ; plural form ) is a Latin phrase meaning "note well". It is often abbreviated as NB, n.b., or with the ligature and first appeared in English writing . In Modern English, it is used, particularly in legal papers, to draw the atten ...

'': A

binomial authority In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...

in parentheses indicates that the species was originally described in a genus other than ''Toxicocalamus''. These former genera, ''Apistocalamus, Apisthocalamus, Pseudapistocalamus, Pseudapisthocalamus, Ultrocalamus,'' and ''Vanapina'', are now

synonyms A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...

of ''Toxicocalamus''.

Taxonomy

The former species ''Pseudapisthocalamus nymani'' Lönnberg, 1900; ''Apisthocalamus pratti'' Boulenger, 1904; ''A. loennbergii'' Boulenger, 1908; and ''A. lamingtoni''

Kinghorn Kinghorn (; gd, Ceann Gronna) is a town and parish in Fife, Scotland. A seaside resort with two beaches, Kinghorn Beach and Pettycur Bay, plus a fishing port, it stands on the north shore of the Firth of Forth, opposite Edinburgh. Accordin ...

, 1928; are synonyms of ''T. loriae'', ''Vanapina lineata''

De Vis Charles Walter de Vis (Birmingham, England, 9 May 1829 – Brisbane, Queensland, Australia, 30 April 1915),

, 1905 is a synonym of ''T. longissimus'', and ''Ultrocalamus latisquamatus'' Schüz, 1929 is a synonym of ''T. preussi''. Most of the described species are poorly known and rarely encountered. The most widely distributed, and most commonly encountered, species is ''T. loriae'' (itself a possible

species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...

), which accounts for 66% of all ''Toxicocalamus'' specimens in museum collections. ''T. loriae'' is frequently encountered in the

Highlands Highland is a broad term for areas of higher elevation, such as a mountain range or mountainous plateau. Highland, Highlands, or The Highlands, may also refer to: Places Albania * Dukagjin Highlands Armenia * Armenian Highlands Australia *Sou ...

, where large numbers have been collected in village gardens along the Wahgi River valley of

Simbu Province Chimbu, more frequently spelled Simbu, is a province in the Highlands Region of Papua New Guinea. The province has an area of 6,112 km2 and a population of 376,473 (2011 census). The capital of the province is Kundiawa. Mount Wilhelm, the ta ...

, PNG. The next most frequently encountered and widely distributed species are ''T. preussi'' and ''T. stanleyanus''. All the other species are much less well known and localised in distribution. Also on mainland New Guinea, ''T. buergersi'' is known from only six specimens, from the

in the

Sepik The Sepik () is the longest river on the island of New Guinea, and the second largest in Oceania by discharge volume after the Fly River. The majority of the river flows through the Papua New Guinea (PNG) provinces of Sandaun (formerly West Se ...

region (

Sandaun Sandaun Province (formerly West Sepik Province) is the northwesternmost mainland province of Papua New Guinea. It covers an area of 35,920 km2 (13868 m2) and has a population of 248,411 (2011 census). The capital is Vanimo. In July 1998 the a ...

and

s), PNG; ''T. spilolepidotus'' is known from two specimens, from the

Kratke Range Kratke Range (german: Krätkegebirge) is a mountain range in Eastern Highlands, Papua New Guinea. Several rivers, including Aziana, Yaiga, Lamari and Ramu originate in the mountains. Many endemic birds live in the mountains. History In Nove ...

Eastern Highlands Province Eastern Highlands is a highlands province of Papua New Guinea. The provincial capital is Goroka. The province covers an area of 11,157 km², and has a population of 579,825 (2011 census). The province shares a common administrative boundary w ...

, PNG; ''T. pachysomus'' is known from its

, from the Cloudy Mountains,

; PNG, ''T. cratermontanus'' from its holotype, from Crater Mountain,

, PNG, while ''T. ernstmayri'' was only known from its holotype in the

Western Province Western Province or West Province may refer to: * Western Province, Cameroon *Western Province, Rwanda *Western Province (Kenya) *Western Province (Papua New Guinea) *Western Province (Solomon Islands) *Western Province, Sri Lanka *Western Provin ...

, PNG, until a second specimen was observed crawling across mine-workings at the

Ok Tedi Mine The Ok Tedi Mine is an open-pit copper and gold mine in Papua New Guinea located near the headwaters of the Ok Tedi River, in the Star Mountains Rural LLG of the North Fly District of the Western Province of Papua New Guinea. The mine is ...

, in the Star Mountains. ''Toxicocalamus grandis'' is also only known from its holotype, collected on the Setakwa River, western New Guinea, in 1912, (the only species represented by a type specimen west of the WNG/PNG border), and ''T. pumehanae'' is also only known from its holotype, from the Managalas Plateau,

Oro Province Oro Province, formerly (and officially still) Northern Province, is a coastal province of Papua New Guinea. The provincial capital is Popondetta. The province covers 22,800 km2, and has 176,206 inhabitants (2011 census). The province shares la ...

, PNG,. On the islands of Milne Bay, ''T. holopelturus'' is known from 19 specimens from

, also known as Yela; ''T. misimae'' is known from six specimens from

; and ''T. mintoni'' is only known from its holotype, from Sudest Island, also known as Vanantai or Tagula Island, all in the

. ''Toxicocalamus nigrescens'' is only known from its holotype and paratype, from

, in the d'Entercasteaux Archipelago, while ''T. longissimus'' is known from 12 specimens from