Colobopsis schmitzi

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Colobopsis schmitzi
Nepenthes bicalcarata and Camponotus schmitzi.png
Colobopsis schmitzi in association with its host plant, Nepenthes bicalcarata
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Formicinae
Genus: Colobopsis
Species:
C. schmitzi
Binomial name
Colobopsis schmitzi
(Stärcke, 1933)
Synonyms
  • Camponotus schmitzi

Colobopsis schmitzi, synonym Camponotus schmitzi, is a species of ant native to Borneo, [1] 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 . They are endemic to the island of Borneo. [2]

Contents

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 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 1 mm (0.039 in) 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. [1]

C. schmitzi exhibit polymorphism, with three physical castes - minor, median, and major ("soldier") workers. The queen averages at a length of 8 mm (0.31 in) with wings measuring 7 mm (0.28 in) long. The major and median workers are both 6.5 mm (0.26 in) in length, while the minor workers are 4 to 5 mm (0.16 to 0.20 in) long. They are brownish-yellow to reddish-orange in color; the head and gaster are darker than the rest of the body. [1]

The larvae are typical of Camponotus larvae - cylindrical in shape with the head and mouthparts bent at a 90 degree angle from the body. [1]

Distribution and habitat

Colobopsis schmitzi inhabits the hollow tendrils of the plant Nepenthes bicalcarata . It is only found in association with N. bicalcarata, which is endemic to the island of Borneo. [2]

Taxonomy

Camponotus schmitzi is classified in the genus Colobopsis, which was formerly treated as a subgenus of Camponotus , tribe Camponotini, and subfamily Formicinae of the ant family Formicidae. It was first collected by the botanist Jan Pieter Schuitemaker and described by the myrmecologist August Stärcke in 1933 as Camponotus schmitzi. [3]

Myrmecotrophic mutualism

Intermediate pitcher of Nepenthes bicalcarata with swollen tendril colonised by Colobopsis schmitzi. The brown scar tissue results from a wound, not from the ants' drilling. Bicalcarata-upperpitcher.jpg
Intermediate pitcher of Nepenthes bicalcarata with swollen tendril colonised by Colobopsis schmitzi. The brown scar tissue results from a wound, not from the ants' drilling.

The ant makes its nest in the hollow tendrils of the pitcher plant Nepenthes bicalcarata . [4] [5] [6] This unique animal-plant interaction was noted by Frederick William Burbidge as early as 1880. [7] In 1904, Odoardo Beccari suggested that the ants feed on insects found on and around the plant, but may fall prey to it themselves. [8] In 1990, B. Hölldobler and E. O. Wilson proposed that N. bicalcarata and C. schmitzi form a mutually beneficial association. [9] At the time, however, no experimental data existed to support such a hypothesis. A series of observations and experiments carried out in Brunei by Charles Clarke in 1992 and 1998, [10] [11] [12] and by Clarke and Kitching in 1993 and 1995, [13] [14] strongly support the mutualism theory.

The ants feed by descending into the pitcher fluid and retrieving arthropods 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 5 cm (2.0 in), may take up to 12 hours. [15] In this way the contents of N. bicalcarata pitchers is controlled such that organic matter does not accumulate to the point of putrefaction, which can lead to the demise of pitcher infauna (which also appear to benefit the plant) and sometimes the pitcher itself. [15]

The ants seem to favour upper pitchers and rarely colonise lower pitchers. [16] 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, larvae, and pupae. [17]

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. [15] 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 2 m (6.6 ft) in height not colonised by C. schmitzi. [15]

John Thompson suggests that N. bicalcarata may be the only plant species that obtains nutrients through both insect capture and ant-hosting habits. [18]

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. [19] 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. [20] 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. [21]

See also

Related Research Articles

<i>Nepenthes</i> Tropical pitcher plants

Nepenthes is a genus of carnivorous plants, also known as tropical pitcher plants, or monkey cups, in the monotypic family Nepenthaceae. The genus includes about 170 species, and numerous natural and many cultivated hybrids. They are mostly liana-forming plants of the Old World tropics, ranging from South China, Indonesia, Malaysia, and the Philippines; westward to Madagascar and the Seychelles (one); southward to Australia (four) and New Caledonia (one); and northward to India (one) and Sri Lanka (one). The greatest diversity occurs on Borneo, Sumatra, and the Philippines, with many endemic species. Many are plants of hot, humid, lowland areas, but the majority are tropical montane plants, receiving warm days but cool to cold, humid nights year round. A few are considered tropical alpine, with cool days and nights near freezing. The name "monkey cups" refers to the fact that monkeys were once thought to drink rainwater from the pitchers.

<i>Nepenthes albomarginata</i> Species of pitcher plant

Nepenthes albomarginata is a tropical pitcher plant native to Borneo, Peninsular Malaysia, and Sumatra.

<i>Nepenthes rajah</i> Species of pitcher plant from Borneo

Nepenthes rajah is a carnivorous pitcher plant species of the family Nepenthaceae. It is endemic to Mount Kinabalu and neighbouring Mount Tambuyukon in Sabah, Malaysian Borneo. Nepenthes rajah grows exclusively on serpentine substrates, particularly in areas of seeping ground water where the soil is loose and permanently moist. The species has an altitudinal range of 1,500–2,650 m (4,920–8,690 ft) above sea level and is thus considered a highland or sub-alpine plant. Due to its localised distribution, N. rajah is classified as an endangered species by the IUCN and listed on CITES Appendix I.

<i>Nepenthes rafflesiana</i> Species of pitcher plant from Southeast Asia

Nepenthes rafflesiana, or Raffles' pitcher-plant, is a species of tropical pitcher plant. It has a very wide distribution covering Borneo, Sumatra, Peninsular Malaysia, and Singapore. Nepenthes rafflesiana is extremely variable, with numerous forms and varieties described. In Borneo alone, there are at least three distinct varieties. The giant form of this species produces enormous pitchers rivalling those of N. rajah in size.

<i>Nepenthes ampullaria</i> Species of pitcher plant

Nepenthes ampullaria is a very distinctive and widespread species of tropical pitcher plant, present in Borneo, the Maluku Islands, New Guinea, Peninsular Malaysia, Singapore, Sumatra, and Thailand.

<i>Nepenthes gracilis</i> Species of pitcher plant from Southeast Asia

Nepenthes gracilis, or the slender pitcher-plant, is a common lowland pitcher plant that is widespread in the Sunda region. It has been recorded from Borneo, Cambodia, Peninsular Malaysia, Singapore, Sulawesi, Sumatra, and Thailand. The species has a wide altitudinal distribution of 0 to 1100 m above sea level, although most populations are found below 100 m and plants are rare above 1000 m. Despite being a widespread plant, natural hybrids between N. gracilis and other species are quite rare.

<i>Nepenthes reinwardtiana</i> Species of pitcher plant from Southeast Asia

Nepenthes reinwardtiana is a tropical pitcher plant native to Borneo and Sumatra and to a number of smaller surrounding islands including Bangka, Natuna, Nias, and Siberut. Although some sources have included Peninsular Malaysia and Singapore within the range of this species, these records appear to be erroneous.

<i>Nepenthes bicalcarata</i> Species of pitcher plant from Borneo

Nepenthes bicalcarata, also known as the fanged pitcher-plant, is a tropical pitcher plant endemic to northwestern Borneo, Indonesia. It is a myrmecophyte noted for its mutualistic association with a species of ant, Camponotus schmitzi. As an ant-fed plant it lacks many of the features that characterise the carnivorous syndrome in Nepenthes, including viscoelastic and highly acidic pitcher fluid, the waxy zone of the pitcher interior, and possibly even functional digestive enzymes.

<i>Nepenthes <span style="font-style:normal;">×</span> kinabaluensis</i> Species of pitcher plant from Borneo

Nepenthes × kinabaluensis, or the Kinabalu pitcher-plant, is the natural hybrid between N. rajah and N. villosa. It was first collected near Kambarangoh on Mount Kinabalu, Borneo by Lilian Gibbs in 1910 and later mentioned by John Muirhead Macfarlane as "Nepenthes sp." in 1914. Although Macfarlane did not formally name the plant, he noted that "[a]ll available morphological details suggest that this is a hybrid between N. villosa and N. rajah". It was finally described in 1976 by Shigeo Kurata as N. × kinabaluensis. The name was first published in Nepenthes of Mount Kinabalu, but was a nomen nudum at the time as it lacked an adequate description and information on the type specimen. The name was subsequently published validly by Kurata in 1984.

<i>Nepenthes lowii</i> Species of pitcher plant from Borneo

Nepenthes lowii, or Low's pitcher-plant, is a tropical pitcher plant endemic to Borneo. It is named after Hugh Low, who discovered it on Mount Kinabalu. This species is perhaps the most unusual in the genus, being characterised by its strongly constricted upper pitchers, which bear a greatly reduced peristome and a reflexed lid with numerous bristles on its lower surface.

<i>Nepenthes stenophylla</i> Species of pitcher plant from Borneo

Nepenthes stenophylla, or the narrow-leaved pitcher-plant, is a tropical pitcher plant endemic to Borneo. The species produces attractive funnel-shaped pitchers up to 25 cm high. It is listed as Least Concern on the IUCN Red List. Nepenthes stenophylla belongs to the loosely defined "N. maxima complex", which also includes, among other species, N. boschiana, N. chaniana, N. epiphytica, N. eymae, N. faizaliana, N. fusca, N. klossii, N. maxima, N. platychila, and N. vogelii.

Dr. Charles M. Clarke is an ecologist and botanist specialising in the carnivorous plant genus Nepenthes, for which he is regarded as a world authority. Clarke has an honours degree in Botany from Monash University in Melbourne, and a Ph.D. in Ecosystem management at the University of New England, in Armidale, New South Wales.

<i>Nepenthes clipeata</i> Species of pitcher plant from Borneo

Nepenthes clipeata, or the shield-leaved pitcher-plant, is a tropical pitcher plant known only from the near-vertical granite cliff faces of Mount Kelam in West Kalimantan, Indonesia. It has an elevational distribution between approximately 600 and 800 m.

<i>Nepenthes ephippiata</i> Species of pitcher plant from Borneo

Nepenthes ephippiata, or the saddle-leaved pitcher-plant, is a tropical pitcher plant endemic to Borneo. It occurs in the Hose Mountains of central Sarawak, as well as Mount Raya and Bukit Lesung in Kalimantan. It grows in montane forest from 1,000 to 1,900 meters elevation.

<i>Nepenthes hispida</i> Species of pitcher plant from Borneo

Nepenthes hispida is a tropical pitcher plant species native to Borneo. It grows at elevations of 100 to 800 m in kerangas forest. It is known with certainty only from Lambir Hills National Park and surrounding areas.

<i>Nepenthes campanulata</i> Species of pitcher plant from Borneo and the Philippines

Nepenthes campanulata, the bell-shaped pitcher-plant, is a tropical pitcher plant native to Borneo. It has also been reported from Palawan, the Philippines, though further field work is needed to confirm this identification.

<i>Nepenthes mapuluensis</i> Tropical pitcher plant endemic to Borneo

Nepenthes mapuluensis, the Mapulu pitcher-plant, is a species of tropical pitcher plant native to East Kalimantan, Borneo. It is known only from a restricted geographical range and is listed as Endangered on the IUCN Red List.

<i>Nepenthes inermis</i> Species of pitcher plant from Sumatra

Nepenthes inermis is a tropical pitcher plant endemic to the Indonesian island of Sumatra. The specific epithet inermis is Latin for "unarmed" and refers to the upper pitchers of this species, which are unique in that they completely lack a peristome.

<i>Nepenthes <span style="font-style:normal;">×</span> trichocarpa</i> Species of pitcher plant from Southeast Asia

Nepenthes × trichocarpa, the dainty pitcher-Plant, is a common natural hybrid involving N. ampullaria and N. gracilis. It was originally thought to be a distinct species and was described as such.

<i>Nepenthes chaniana</i> Species of pitcher plant from Borneo

Nepenthes chaniana is a tropical pitcher plant species belonging to the genus Nepenthes. It is characterised by a dense indumentum of long, white hairs. Pitchers are cylindrical and mostly white to yellow in colouration. Nepenthes chaniana belongs to the loosely defined "N. maxima complex", which also includes, among other species, N. boschiana, N. epiphytica, N. eymae, N. faizaliana, N. fusca, N. klossii, N. maxima, N. platychila, N. stenophylla, and N. vogelii.

References

  1. 1 2 3 4 (in French) Schuitemaker, J.P. & A. Stärcke 1933. "Contribution a l'étude de la faune Népenthicole. Art. III. Un nouveau Camponotus de Bornéo, habitant les tiges creuses de Nepenthes, récolté par J. P. Schuitemaker et décrit par A. Stärcke, den Dolder" (PDF).[ permanent dead link ]Overdruk uit het Natuurhistorisch Maandblad22(3): 29–31.
  2. 1 2 Eric Hansen (October 1, 2001). "Flesh-eating Plants". Discover Magazine. Retrieved January 12, 2012.
  3. ↑ R.L. Kitching (2000). Food Webs and Container Habitats: The natural history and ecology of phytotelmata. Cambridge University Press. p. 371. ISBN   978-0-521-77316-4.
  4. ↑ Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  5. ↑ Shelford, R. 1916. A Naturalist in Borneo. T. Fisher Unwin, London.
  6. ↑ Cresswell, J.E. 2000. Resource input and the community structure of larval infaunas of an eastern tropical pitcher plant Nepenthes bicalcarata. Ecological Entomology25(3): 362–366. doi : 10.1046/j.1365-2311.2000.00267.x
  7. ↑ Burbidge, F.W. 1880. The Gardens of the Sun: or, a Naturalist's Journal on the Mountains and in the Forests and Swamps of Borneo and the Sulu Archipelago. John Murray, London.
  8. ↑ Beccari, O. 1904. Wanderings in the great forests of Borneo. Archibald and Constable, London.
  9. ↑ Hölldobler, B. & E.O. Wilson 1990. The Ants. Springer-Verlag, Berlin, Germany.
  10. ↑ Clarke, C.M. 1992. The ecology of metazoan communities in Nepenthes pitcher plants in Borneo, with special reference to the community of Nepenthes bicalcarata Hook.f. Ph.D. thesis, University of New England, Armidale, New South Wales. 269 pp.
  11. ↑ Clarke, C.M. 1998. Initial colonisation and prey capture in Nepenthes bicalcarata (Nepenthaceae) pitchers in Brunei. Sandakania12: 27–36.
  12. ↑ Clarke, C.M. 1998. The aquatic arthropod community of the pitcher plant, Nepenthes bicalcarata (Nepenthaceae) in Brunei. Sandakania11: 55–60.
  13. ↑ Clarke, C.M. & R.L. Kitching 1993. The Metazoan Food Webs from Six Bornean Nepenthes Species. Ecological Entomology18: 7–16.
  14. ↑ Clarke, C.M. & R.L. Kitching 1995. Swimming Ants and Pitcher Plants: a Unique Ant-Plant Interaction from Borneo. Journal of Tropical Ecology11(4): 589–602.
  15. 1 2 3 4 Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  16. ↑ Clarke, C.M. 1997. The effects of pitcher dimorphism on the metazoan community of the carnivorous pitcher plant Nepenthes bicalcarata Hook.f.. Malayan Nature Journal50: 149–157.
  17. ↑ Clarke, C.M. & C.C. Lee 2004. Pitcher Plants of Sarawak. Natural History Publications (Borneo), Kota Kinabalu.
  18. ↑ Thompson, J.H. 1981. Reverse animal-plant interactions: the evolution of insectivorous and ant-fed plants. Biological Journal of the Linnean Society16: 147–155.
  19. ↑ M. A. Merbach, G. Zizka, B. Fiala, D. J. Merbach, W. E. Booth, U. Maschwitz (2007). Why a carnivorous plant cooperates with an ant. Selective defense against pitcher-destroying weevils in the myrmecophytic pitcher plant Nepenthes bicalcarata. Ecotropica 13, 45–56.
  20. ↑ Bonhomme, V.; Gounand, I.; Alaux, C.; Jousselin, E.; Barthelemy, D. & Gaume, L. (2011). The plant-ant Colobopsis schmitzi helps its carnivorous host-plant Nepenthes bicalcarata to catch its prey. Journal of Tropical Ecology 27, 15–24.
  21. ↑ 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.
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