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''Colobopsis schmitzi'', synonym ''Camponotus schmitzi'', is a species of
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
native to
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
, Schuitemaker, J.P. & A. Stärcke 1933. ''Overdruk uit het Natuurhistorisch Maandblad'' 22(3): 29–31. which is commonly known as the diving ant, swimming ant or pitcher-plant ant, due to their habit of diving into the digestive fluids of their plant host ''
Nepenthes bicalcarata ''Nepenthes bicalcarata'' (; from Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now ...
''. They are
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 the island of
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
.


Description

''C. schmitzi'' closely resemble '' Colobopsis ceylonicus'' but are slightly larger. They have rectangular heads about one and a half times as long as they are wide. The
mandibles In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
have five teeth each (except for the minor workers which have four). The eyes are widely separated and located laterally, slightly behind the anterior bulge of the head. The antennae are short, with a scape at in length. The funicules are longer than they are wide proximally and become shorter and wider towards the tip. The body is smooth and shiny. The gaster is small. The legs are relatively large, especially the femurs, and very compressed laterally. Spines are present near the distal tip of the femurs and absent in the tibiae. The tarsi are longer than the tibiae. ''C. schmitzi'' exhibit polymorphism, with three physical castes - minor, median, and major ("soldier") workers. The queen averages at a length of with wings measuring long. The major and median workers are both in length, while the minor workers are long. They are brownish-yellow to reddish-orange in color; the head and gaster are darker than the rest of the body. The
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e are typical of ''Camponotus'' larvae - cylindrical in shape with the head and mouthparts bent at a 90 degree angle from the body.


Distribution and habitat

''Colobopsis schmitzi'' inhabits the hollow
tendril In botany, a tendril is a specialized Plant stem, stem, leaf or Petiole (botany), petiole with a thread-like shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as ''Cuscuta''. There ar ...
s of the plant ''
Nepenthes bicalcarata ''Nepenthes bicalcarata'' (; from Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now ...
''. It is only found in association with ''N. bicalcarata'', which 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 the island of
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
.


Taxonomy

''Camponotus schmitzi'' is classified in the genus ''Colobopsis'', which was formerly treated as a subgenus of ''
Camponotus Carpenter ants (''Camponotus'' spp.) are a genus of large ants (workers ) indigenous to many parts of the world. True carpenter ants build nests inside wood, consisting of galleries chewed out with their mandibles or jaws, preferably in dead, ...
'',
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...
Camponotini Camponotini is a tribe containing 2 extinct ant genera and 8 extant ant genera, including '' Camponotus'' (carpenter ants). Ants in the Camponotini tribe are the primary hosts of ''Ophiocordyceps unilateralis'', also called the zombie-ant fungus ...
, and
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
Formicinae The Formicinae are a subfamily within the Formicidae containing ants of moderate evolutionary development. Formicines retain some primitive features, such as the presence of cocoons around pupae, the presence of ocelli in workers, and littl ...
of the ant family Formicidae. It was first collected by the
botanist Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
Jan Pieter Schuitemaker and described by the
myrmecologist Myrmecology (; from Greek: μύρμηξ, ''myrmex'', "ant" and λόγος, ''logos'', "study") is a branch of entomology focusing on the study of ants. Ants continue to be a model of choice for the study of questions on the evolution of social ...
August Stärcke in 1933 as ''Camponotus schmitzi''.


Myrmecotrophic mutualism

The ant makes its nest in the hollow tendrils of the
pitcher plant Pitcher plants are carnivorous plants known as pitfall traps—a prey-trapping mechanism featuring a deep cavity filled with digestive liquid. The traps of pitcher plant are considered to be "true" pitcher plants and are formed by specialized ...
''
Nepenthes bicalcarata ''Nepenthes bicalcarata'' (; from Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now ...
''. This unique animal-plant interaction was noted by
Frederick William Burbidge Frederick William Thomas Burbidge (1847–1905) was a British explorer who collected many rare tropical plants for the famous Veitch Nurseries. Biography Burbidge was born at Wymeswold, Leicestershire, on 21 March 1847, was son of Thomas Burbid ...
as early as 1880. In 1904,
Odoardo Beccari Odoardo Beccari (16 November 1843 – 25 October 1920) was an Italian botanist famous for his discoveries in Indonesia, New Guinea, and Australia. He has been called the greatest botanist to ever study Malesia. Life Youth and education (18 ...
suggested that the ants feed on insects found on and around the plant, but may fall prey to it themselves. In 1990, B. Hölldobler and
E. O. Wilson Edward Osborne Wilson (June 10, 1929 – December 26, 2021) was an American biologist, naturalist, ecologist, and entomologist known for developing the field of sociobiology. Born in Alabama, Wilson found an early interest in nature and frequ ...
proposed that ''N. bicalcarata'' and ''C. schmitzi'' form a mutually beneficial association. At the time, however, no experimental data existed to support such a hypothesis. A series of observations and experiments carried out in
Brunei Brunei, officially Brunei Darussalam, is a country in Southeast Asia, situated on the northern coast of the island of Borneo. Apart from its coastline on the South China Sea, it is completely surrounded by the Malaysian state of Sarawak, with ...
by
Charles Clarke Charles Rodway Clarke (born 21 September 1950) is a British Labour Party politician who held various Cabinet positions under Prime Minister Tony Blair from 2001 to 2006, lastly as Home Secretary from December 2004 to May 2006. Clarke was th ...
in 1992 and 1998, and by Clarke and Kitching in 1993 and 1995, strongly support the mutualism theory. The ants feed by descending into the pitcher fluid and retrieving
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s caught by the plant. The ants seem to ignore smaller insects and only target larger prey items. Hauling food from the pitcher fluid to the peristome, a distance of no more than , may take up to 12 hours. In this way the contents of ''N. bicalcarata'' pitchers is controlled such that organic matter does not accumulate to the point of
putrefaction Putrefaction is the fifth stage of death, following pallor mortis, livor mortis, algor mortis, and rigor mortis. This process references the breaking down of a body of an animal Post-mortem interval, post-mortem. In broad terms, it can be view ...
, which can lead to the demise of pitcher infauna (which also appear to benefit the plant) and sometimes the pitcher itself. The ants seem to favour upper pitchers and rarely colonise lower pitchers. This is likely because terrestrial traps are periodically submerged in water during heavy rains. Flooding of the ants' nest chamber could result in the death of the developing eggs,
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e, and
pupa A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
e.Clarke, C.M. & C.C. Lee 2004. ''Pitcher Plants of Sarawak''. Natural History Publications (Borneo), Kota Kinabalu. ''C. schmitzi'' nests solely in the tendrils of ''N. bicalcarata'' and rarely ventures onto other plants. The species is completely dependent on ''N. bicalcarata'' for food and
domicile Domicile may refer to: * Home, a place where someone lives * Domicile (astrology), the zodiac sign over which a planet has rulership * Domicile (law), the status or attribution of being a permanent resident in a particular jurisdiction See also * ...
.Clarke, C.M. 1997. ''Nepenthes of Borneo''. Natural History Publications (Borneo), Kota Kinabalu. ''N. bicalcarata'', on the other hand, is able to survive and reproduce without the presence of the ants; it is a facultative mutualist. This being the case, there appear to be few mature plants over in height not colonised by ''C. schmitzi''. John Thompson suggests that ''N. bicalcarata'' may be the only plant species that obtains nutrients through both insect capture and ant-hosting habits. In addition to Clarke & Kitching (1995), three more theories have been investigated so far to explain the symbiotic relationship between ant and plant. In an exclusion experiment it was shown that plants without ''C. schmitzi'' receive greater herbivory damage, and the ants seem to specifically attack a certain weevil ('' Alcidodes'' sp.) that feeds on pitcher plants. Furthermore, data has been collected that hints to an aggressiveness of ''C. schmitzi'' against prey animals that try to escape from the pitchers, thus enhancing the retention rate. Another mechanism appears to be a host-cleaning behaviour of ''C. schmitzi'': the ants remove debris and mold from the peristome, which thus remains smooth and slippery and has a longer operational lifespan.Thornham, D. G.; Smith, J. M.; Ulmar Grafe, T. & Federle, W. (2012). Setting the trap: cleaning behaviour of ''Colobopsis schmitzi'' ants increases long-term capture efficiency of their pitcher plant host, ''Nepenthes bicalcarata''. ''Functional Ecology'' 26, 11–19.


See also

* ''Nepenthes'' infauna *
Myrmecophyte Myrmecophytes (; literally "ant-plant") are plants that live in a mutualistic association with a colony of ants. There are over 100 different genera of myrmecophytes. These plants possess structural adaptations in the form of domatia where ants ...
*
Mutualism (biology) Mutualism describes the ecological Biological interaction, interaction between two or more species where each species has a net benefit. Mutualism is a common type of Ecology, ecological interaction. Prominent examples are: * the nutrient exch ...


References


External links


''Camponotus schmitzi''antbase.net"Setting the trap"
- video by Thornham ''et al.'' of the myrmecotrophic mutualism exhibited by ''C. schmitzi'' from YouTube {{Taxonbar, from=Q1153724 schmitzi Hymenoptera of Asia Insects of Borneo Nepenthes infauna Insects described in 1933 Endemic fauna of Borneo