Colobopsis schmitzi | |
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Colobopsis schmitzi in association with its host plant, Nepenthes bicalcarata | |
Scientific classification | |
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 | |
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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]
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]
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]
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]
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]
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.
Nepenthes albomarginata is a tropical pitcher plant native to Borneo, Peninsular Malaysia, and Sumatra.
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.
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.
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.
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.
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.
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.
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.
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.
Nepenthes macrophylla, the large-leaved pitcher-plant, is a tropical pitcher plant known only from a very restrictive elevation on Mount Trusmadi in Sabah, Malaysian Borneo.
Nepenthes hurrelliana is a tropical pitcher plant endemic to Borneo, where it has been recorded from northern Sarawak, southwestern Sabah, and Brunei. It is of putative hybrid origin; its two original parent species are thought to be N. fusca and N. veitchii. A thick indumentum of rusty-brown hairs covers the entire plant, a characteristic presumably inherited from the latter.
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.
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.
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.
Nepenthes beccariana is a tropical pitcher plant. The species was described in 1908 by John Muirhead Macfarlane based on a specimen collected from the island of Nias, which lies off the western coast of Sumatra. It appears to be closely related to both N. longifolia and N. sumatrana, and the former is possibly a heterotypic synonym of this taxon.
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.
Nepenthes attenboroughii, or Attenborough's pitcher plant, is a montane species of carnivorous pitcher plant of the genus Nepenthes. It is named after the celebrated broadcaster and naturalist Sir David Attenborough, who is a keen enthusiast of the genus. The species is characterised by its large and distinctive bell-shaped lower and upper pitchers and narrow, upright lid. The type specimen of N. attenboroughii was collected on the summit of Mount Victoria, an ultramafic mountain in central Palawan, the Philippines.
Nepenthes of Borneo is a monograph by Charles Clarke on the tropical pitcher plants of Borneo. It was first published in 1997 by Natural History Publications (Borneo), and reprinted in 2006. Clarke describes it as "primarily an ecological monograph".