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''Toxicocalamus'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
snake Snakes are elongated limbless reptiles of the suborder Serpentes (). Cladistically squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales much like other members of the group. Many species of snakes have s ...
s in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Elapidae Elapidae (, commonly known as elapids , from , variant of "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 '' Emydocephalus ...
. The genus is
endemic Endemism is the state of a species being found only 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 foun ...
to
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
.


Description

Most species of ''Toxicocalamus'' are relatively small, the largest specimen known being the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
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), 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 Phospholipase A can refer to: * Phospholipase A1 * Phospholipase A2 * Outer membrane phospholipase A1 An enzyme that displays both phospholipase A1 and phospholipase A2 activities is called a Phospholipase B (see main article on phospholipase ...
(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 myoge ...
(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 A fossorial animal () is one that is adapted to digging and which lives primarily (but not solely) underground. Examples of fossorial vertebrates are Mole (animal), moles, badgers, naked mole-rats, meerkats, armadillos, wombats, and mole salamand ...
, 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
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
, northern coastal offshore islands, i.e. Seleo Is. (
Sandaun Province Sandaun Province (formerly West Sepik Province) is the northwesternmost mainland Provinces of Papua New Guinea, province of Papua New Guinea (also known as home of the sunset). It covers an area of 35,920 km2 (13868 m2) and has a population ...
, 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 450,530 people (2011 census) and is 43,426 km square in size. Its density is 10.4 people per square kilometer. History Cherubim D ...
, PNG), and Karkar Is. (
Madang Province Madang is a Provinces of Papua New Guinea, 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 capi ...
, 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 ...
to the southeast, i.e. d'Entrecasteaux Archipelago ( 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 o ...
( Misima Is., Sudest Is., and Rossel Is.).


Diet

The
prey Predation is a biological interaction in which 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 ki ...
of snakes in the genus ''Toxicocalamus'' appears to consist almost entirely of
earthworm An earthworm is a soil-dwelling terrestrial invertebrate that belongs to the phylum Annelida. The term is the common name for the largest members of the class (or subclass, depending on the author) Oligochaeta. In classical systems, they we ...
s, particularly the giant earthworms of the
Megascolecidae Megascolecidae is a family of earthworms native to Madagascar, Australia, New Zealand, Asia, and North America. At up to 2 meters in length, their large size distinguishes the Megascolecidae from other earthworm families. They are an essential par ...
, 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 , from , variant of "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 '' Emydocephalus' ...
, ''Toxicocalamus'' is believed to reproduce by
oviparity Oviparous animals are animals that reproduce by depositing fertilized zygotes outside the body (i.e., by laying or spawning) in metabolically independent incubation organs known as eggs, which nurture the embryo into moving offsprings kno ...
, 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 24 species, one of which has two
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
, are currently recognised as being valid in the genus ''Toxicocalamus'': * '' Toxicocalamus atratus'' Kraus, Kaiser,, & O’Shea, 2022 – Black forest snake * '' 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-language surname, meaning 'curly'. In addition to German-speaking countries, the surname is commonly found in the United States, Israel, and the Czech Republic (Czech feminine: Krausová). Notable people with the surname include ...
, 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. Historian C. Thomas Cairney states that the O'Sheas were ...
, Parker &
Kaiser Kaiser ( ; ) is the title historically used by German and Austrian emperors. In German, the title in principle applies to rulers anywhere in the world above the rank of king (). In English, the word ''kaiser'' is mainly applied to the emperors ...
, 2015
Star Mountains The Star Mountains ( Dutch ( colonial)'': Sterrengebergte''; Indonesian'': Pegunungan Bintang'') are a mountain range in eastern end of Highland Papua, Indonesia and the western Papua New Guinea, 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 snake * '' Toxicocalamus lamingtoni'' Kinghorn, 1928 – Mount Lamington forest snake * '' Toxicocalamus loennbergii'' Boulenger, 1908 – Lönnberg’s forest snake * '' Toxicocalamus longhagen'' Roberts, Iova, & Austin, 2022 * '' 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 forest snake, Sudest Island snake * '' Toxicocalamus misimae'' McDowell, 1969 – Misima Island forest snake, Misima Island 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 nymani'' (Lönnberg, 1900) – Loria forest 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 on the island of New Guinea, after the Sepik and Mamberamo, with a total length of . It is the largest by volume of discharge in Oceania, the largest in the world without a single dam in its catchment, an ...
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 * Allison family, a family of RMS Titanic passengers Compan ...
& Kaiser, 2018
– Managalas Plateau snake * '' Toxicocalamus spilolepidotus'' McDowell, 1969 – Krakte Mountains spotted snake, spotted forest snake * '' Toxicocalamus spilorhynchus'' Kraus, Kaiser,, & O’Shea, 2022 * '' Toxicocalamus stanleyanus'' Boulenger, 1903Owen Stanley Mountains snake, Owen Stanley Range forest snake * '' Toxicocalamus vertebralis'' Kraus, Kaiser,, & O’Shea, 2022 – striped forest snake ''
Nota bene ( ; plural: ) is the Latin language, Latin phrase meaning ''note well''. In manuscripts, ''nota bene'' is abbreviated in upper-case as NB and N.B., and in lower-case as n.b. and nb; the editorial usages of ''nota bene'' and ''notate bene'' fi ...
'': A
binomial authority In taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...
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 precisely 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 a ...
of ''Toxicocalamus''.


Taxonomy

The former species ''Pseudapisthocalamus nymani'' Lönnberg, 1900; ''Apisthocalamus pratti'' Boulenger, 1904; ''A. loennbergii'' Boulenger, 1908; and ''A. lamingtoni''
Kinghorn Kinghorn (; ) 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. Known as the place where K ...
, 1928
; are synonyms of ''T. loriae'', ''Vanapina lineata''
De Vis Charles Walter de Vis (9 May 1829 – 30 April 1915)
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 Africa * Highlands, Johannesburg, South Africa * Highlands, Harare, Zimbab ...
, 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 t ...
, 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
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 ...
in the
Sepik The Sepik () is the longest river on the island of New Guinea, and the third largest in Oceania by discharge volume after the Fly River, Fly and Mamberamo River, Mamberamo. The majority of the river flows through the Papua New Guinea (PNG) provi ...
region (
Sandaun Sandaun Province (formerly West Sepik Province) is the northwesternmost mainland province of Papua New Guinea (also known as home of the sunset). It covers an area of 35,920 km2 (13868 m2) and has a population of 248,411 (2011 census). The c ...
and
East Sepik Province East Sepik is a province in Papua New Guinea. Its capital is Wewak. East Sepik has an estimated population of 450,530 people (2011 census) and is 43,426 km square in size. Its density is 10.4 people per square kilometer. History Cherubim D ...
s), PNG; ''T. spilolepidotus'' is known from two specimens, from the Kratke Range,
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
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
, from the Cloudy Mountains,
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 ...
; PNG, ''T. cratermontanus'' from its holotype, from Crater Mountain,
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 t ...
, PNG, while ''T. ernstmayri'' was only known from its holotype in the
Star Mountains The Star Mountains ( Dutch ( colonial)'': Sterrengebergte''; Indonesian'': Pegunungan Bintang'') are a mountain range in eastern end of Highland Papua, Indonesia and the western Papua New Guinea, stretching from the eastern end of Indonesia t ...
of
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 Provinc ...
, PNG, until a second specimen was observed crawling across mine-workings at the
Ok Tedi Mine The Ok Tedi Mine is an Open pit mine, open-pit copper mine, copper and gold mine Mining in Papua New Guinea, 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 ...
, 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 Provinces of Papua New Guinea, province in the Southern Region, Papua New Guinea, Southern Region of Papua New Guinea. The provincial capital is Popondetta. The provinc ...
, PNG,. On the islands of
Milne Bay Milne Bay is a large bay in Milne Bay Province, south-eastern Papua New Guinea. More than long and over wide, Milne Bay is a sheltered deep-water harbor accessible via Ward Hunt Strait. It is surrounded by the heavily wooded Stirling Range (Papu ...
, ''T. holopelturus'' is known from 19 specimens from Rossel Island, also known as Yela; ''T. misimae'' is known from six specimens from Misima Island; and ''T. mintoni'' is only known from its holotype, from Sudest Island, also known as Vanantai or Tagula Island, all in 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 o ...
. ''Toxicocalamus nigrescens'' is only known from its holotype and paratype, from
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 ...
, in the d'Entercasteaux Archipelago, while ''T. longissimus'' is known from 12 specimens from
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 ...
.


References

{{Taxonbar, from=Q195360 Venomous snakes Snake genera Taxa named by George Albert Boulenger