''Nepenthes'' infauna are the organisms that inhabit the pitchers of ''

Nepenthes ''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 mos ...

'' plants. These include

fly Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwing ...

and

midge A midge is any small fly, including species in several families of non-mosquito Nematoceran Diptera. Midges are found (seasonally or otherwise) on practically every land area outside permanently arid deserts and the frigid zones. Some mid ...

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...

spiders Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species dive ...

mites Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...

ants Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22,00 ...

, and even a species of

crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all the ...

, ''

Geosesarma malayanum ''Geosesarma malayanum'' is a species of small red crab found in Malaysia. It is famous for its relationship with pitcher plants; as such, it is classified as a nepenthephile. ''G. malayanum'' is known to visit ''Nepenthes ampullaria'' plants a ...

''. The most common and conspicuous predators found in pitchers are

mosquito Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning " gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "li ...

e, which consume large numbers of other larvae during their development. Many of these animals are so specialised that they cannot survive anywhere else, and are referred to as ''nepenthebionts''. The complex relationships between these various organisms are not yet fully understood. The question of whether infaunal animals "steal" food from their hosts, or whether they are involved in a mutually beneficial (

symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...

) association has yet to be investigated experimentally and is the source of considerable debate.

Charles Clarke Charles Rodway Clarke (born 21 September 1950) is a British Labour Party politician, who was the Member of Parliament (MP) for Norwich South from 1997 until 2010, and served as Home Secretary from December 2004 until May 2006. Early life Th ...

suggests that mutualism is a "likely situation", whereby "the infauna receives

domicile Domicile may refer to: * Home, a place where someone lives * Domicile (astrology), the zodiac sign over which a planet has rulership * Domicile (law) Domicile is relevant to an individual's "personal law," which includes the law that governs a p ...

, protection and food from the plant, while in return, the infauna helps to break down the prey, increase the rate of digestion and keep bacterial numbers low".

Classification

Nepenthebionts

Nepenthebionts are animals which are specialised to live inside ''Nepenthes'' pitchers and are totally dependent on them at least at some stage of their lives. Many species of

e fall into this category. Examples include ''

Culex rajah ''Culex rajah'' is a species of mosquito in the genus ''Culex''. It is endemic to Sabah, Malaysian Borneo. ''C. rajah'' is placed in the subgenus ''Culiciomyia''. In its larval stage, ''C. rajah'' is found exclusively in the pitchers of ''Nepe ...

'' and ''

Toxorhynchites rajah ''Toxorhynchites rajah'' is a species of mosquito in the genus '' Toxorhynchites''. It is endemic to Sabah, Malaysian Borneo. In its larval stage, ''T. rajah'' is found exclusively in the pitchers of ''Nepenthes rajah'' (hence the name), a spec ...

'', which are named after ''

Nepenthes rajah ''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.Clarke 1997, p. 123. ''Nepenthes rajah'' grows exclusively on ...

'', as well as the related species ''

Culex jenseni ''Culex jenseni'' is a species of mosquito in the genus '' Culex''. It is endemic to Sabah, Malaysian Borneo. ''C. jenseni'' is placed in the subgenus ''Lophoceraomyia''. In its larval stage, ''C. jenseni'' develops in the pitchers of ''Nepen ...

'' and ''

Uranotaenia moultoni ''Uranotaenia moultoni'' is a species of mosquito in the genus ''Uranotaenia''. It is endemic to Sabah, Malaysian Borneo. ''U. moultoni'' is placed in the subgenus ''Pseudoficalbia''. In its larval stage, ''U. moultoni'' develops in the pitch ...

''.

Nepenthephiles

Nepenthephiles are organisms that are frequently found in ''Nepenthes'' pitchers, but which are not completely dependent on them at any stage of their lives. Most animals that coexist with ''Nepenthes'' fall into this category. Examples include '' Misumenops nepenthicola'', a

crab spider The Thomisidae are a family of spiders, including about 170 genera and over 2,100 species. The common name crab spider is often linked to species in this family, but is also applied loosely to many other families of spiders. Many members of th ...

that feeds on flies caught by ''Nepenthes'', and ''

'', a species of

Nepenthexenes

Nepenthexenes are animals which are not normally associated with pitchers, but which are occasionally encountered in them. These are usually found if a pitcher becomes overloaded with

putrefying Putrefaction is the fifth stage of death, following pallor mortis, algor mortis, rigor mortis, and livor mortis. This process references the breaking down of a body of an animal, such as a human, post-mortem. In broad terms, it can be viewed ...

prey, when it may be colonised by the larvae of various

species. These simply feed on rotting matter regardless of whether it is found in pitchers.

Diversity

In a 1991 study,Mogi, M. & H. S. Yong. 1992. Aquatic arthropod communities in ''Nepenthes'' pitchers: the role of niche differentiation, aggregation, predation and competition in community organization. ''

Oecologia ''Oecologia'' is an international peer-reviewed English-language journal published by Springer since 1968 (some articles were published in German or French until 1976). The journal publishes original research in a range of topics related to plant ...

'' 90(2): 172-184
Abstract
- a wide diversity of animals was found in pitchers of ''

Nepenthes ampullaria ''Nepenthes ampullaria'' (; Latin ''ampulla'' meaning "flask") 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.McPh ...

'': Carrion feeders: * '' Endonepenthia'' (

Phoridae The Phoridae are a family of small, hump-backed flies resembling fruit flies. Phorid flies can often be identified by their escape habit of running rapidly across a surface rather than taking to the wing. This behaviour is a source of one of thei ...

) * Syrphidae Filter feeders: * ''

Aedes albopictus ''Aedes albopictus'' (''Stegomyia albopicta''), from the mosquito (Culicidae) family, also known as the (Asian) tiger mosquito or forest mosquito, is a mosquito native to the tropical and subtropical areas of Southeast Asia. In the past few ce ...

'', ''

Culex coerulescens ''Culex'' is a genus of mosquitoes, several species of which serve as vectors of one or more important diseases of birds, humans, and other animals. The diseases they vector include arbovirus infections such as West Nile virus, Japanese encep ...

'', '' Culex hewitti'', '' Culex navalis'', '' Tripteroides nepenthis'', '' Tripteroides tenax'', ''

'' (

Culicidae Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning "gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "litt ...

) Detritus feeders: * several Anoetidae * ''

Dasyhelea ''Dasyhelea'' is a genus of biting midges and the only genus of its subfamily, the Dasyheleinae. Larvae of species in this genus are characterized by an anal segment with retractile posterior prolegs. Larvae are aquatic animal, aquatic and adults ...

'', ''

Forcipomyia ''Forcipomyia'' is a genus of biting midges in the subfamily Forcipomyiinae. Species of the subgenus '' Lasiohelea'' suck vertebrate blood. Some species are ectoparasites on larger insects. Other species in the genus are important pollinators of t ...

'' (

Ceratopogonidae Ceratopogonidae is a family of flies commonly known as no-see-ums, or biting midges, generally in length. The family includes more than 5,000 species, distributed worldwide, apart from the Antarctic and the Arctic. Ceratopogonidae are holomet ...

) * several

Chironomidae The Chironomidae (informally known as chironomids, nonbiting midges, or lake flies) comprise a family of nematoceran flies with a global distribution. They are closely related to the Ceratopogonidae, Simuliidae, and Thaumaleidae. Many species ...

Harpacticoida Harpacticoida is an order of copepods, in the subphylum Crustacea. This order comprises 463 genera and about 3,000 species; its members are benthic copepods found throughout the world in the marine environment (most families) and in fresh water ...

Sciaridae The Sciaridae are a family of flies, commonly known as dark-winged fungus gnats. Commonly found in moist environments, they are known to be a pest of mushroom farms and are commonly found in household plant pots. This is one of the least studie ...

Nipping predators (can swim in the fluid,

mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...

s move against one another in a horizontal plane): * '' Aedes brevitibia'', '' Toxorhynchites indicus'' (Culicidae) * ''

Corethrella calathicola ''Corethrella'' is a genus of midges that are classified in the family Corethrellidae Corethrellidae are a family of biting midges, small flying insects belonging to the order Diptera, females of which feed on the blood of frogs. The members o ...

'' (

Chaoboridae Chaoboridae, commonly known as phantom midges or glassworms, is a family of fairly common midges with a cosmopolitan distribution. They are closely related to the Corethrellidae and Chironomidae; the adults are differentiated through peculiarit ...

) Hooking predators (can only crawl, mandibles move parallel to one another in a vertical plane): *

Dolichopodidae Dolichopodidae, the long-legged flies, are a large, cosmopolitan family of true flies with more than 7,000 described species in about 230 genera. The genus ''Dolichopus'' is the most speciose, with some 600 species. Dolichopodidae generally are ...

* '' Listodiplosis'' (

Cecidomyiidae Cecidomyiidae is a family of flies known as gall midges or gall gnats. As the name implies, the larvae of most gall midges feed within plant tissue, creating abnormal plant growths called galls. Cecidomyiidae are very fragile small insects usu ...

) * '' Nepenthosyrphus'' (Syrphidae) Terrestrial predators: * ''

Misumenops ''Misumenops'' is a common genus of crab spider with more than 50 described species. The majority of the species Mesumenops, more than 80, have been transferred to 13 genera: '' Ansiea'', '' Demogenes'', '' Diaea'', '' Ebelingia'', '' Ebrechtel ...

'' (

Thomisidae The Thomisidae are a family of spiders, including about 170 genera and over 2,100 species. The common name crab spider is often linked to species in this family, but is also applied loosely to many other families of spiders. Many members of thi ...

) * '' Xenoplatyura'' (

Mycetophilidae The Mycetophilidae are a family of small flies, forming the bulk of those species known as fungus gnats. About 3000 described species are placed in 150 genera, but the true number of species is undoubtedly much higher. They are generally found ...

) Nepenthebiont

Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...

: * '' Aedes brevitibia'' * several

* ''

'' * '' Culex hewitti'' * ''

'' * '' Culex navalis'' * several ''

'' species ('' D. ampullariae'', '' D. biseriata'' and others) *

* '' Endonepenthia'' * '' Nepenthosyrphus'' * '' Toxorhynchites indicus'' * '' Tripteroides nepenthis'' * '' Tripteroides tenax'' * ''

References

{{Nepenthes

Classification

Nepenthebionts

Nepenthephiles

Nepenthexenes

Diversity

See also

References