Nepenthes infauna
Encyclopedia
Nepenthes infauna are the organisms that inhabit the pitchers of Nepenthes
Nepenthes
The Nepenthes , popularly known as tropical pitcher plants or monkey cups, are a genus of carnivorous plants in the monotypic family Nepenthaceae. The genus comprises roughly 130 species, numerous natural and many cultivated hybrids...

plants. These include fly
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...

 and midge
Midge (insect)
Midges comprise many kinds of very small two-winged flies found world-wide. The term does not encapsulate a well-defined taxonomic group, but includes animals in several families of Nematoceran Diptera. While some midges are vectors for disease, many others play useful roles as prey items for...

 larva
Larva
A larva 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...

e, spiders, mites
MITES
MITES, or Minority Introduction to Engineering and Science, is a highly selective six-week summer program for rising high school seniors held at the Massachusetts Institute of Technology. Its purpose is to expose students from minority, or otherwise disadvantaged backgrounds, to the fields of...

, ants, and even a species of crab
Crab
True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...

, Geosesarma malayanum
Geosesarma malayanum
Geosesarma malayanum is a species of small red crab which is 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 and raid the pitchers of their contents. It uses its claws...

. The most common and conspicuous predators found in pitchers are mosquito
Mosquito
Mosquitoes are members of a family of nematocerid flies: the Culicidae . The word Mosquito is from the Spanish and Portuguese for little fly...

 larva
Larva
A larva 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...

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
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...

) association has yet to be investigated experimentally and is the source of considerable debate. Charles Clarke
Charles Clarke (botanist)
Dr. Charles M. Clarke is a botanist and taxonomist specialising in the carnivorous plant genus Nepenthes. 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.Clarke first...

 suggests that mutualism is a "likely situation", whereby "the infauna receives domicile
Domicile
*In architecture, a general term for a place of residence or "permanent residence" in legal terms*Domicile , the zodiac sign over which a planet has rulership...

, 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".

Nepenthebionts

Nepenthebionts are animals which are specialised to life inside Nepenthes pitchers and are totally dependent on them at least at some stage of their lives. Many species of mosquito
Mosquito
Mosquitoes are members of a family of nematocerid flies: the Culicidae . The word Mosquito is from the Spanish and Portuguese for little fly...

 larva
Larva
A larva 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...

e fall into this category. Examples include Culex rajah
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 Nepenthes rajah , a species of pitcher plant. As such, it is considered a...

and Toxorhynchites rajah
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 , a species of pitcher plant. As such, it is considered a nepenthebiont.-References:* *...

, which are named after Nepenthes rajah
Nepenthes rajah
Nepenthes rajah is an insectivorous pitcher plant species of the Nepenthaceae family. 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...

, as well as the related species Culex jenseni
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 Nepenthes species, especially N. rajah. As such, it is considered a...

and Uranotaenia moultoni
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 pitchers of Nepenthes species, especially N. rajah. As such, it is considered a...

.

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
Misumenops nepenthicola
Misumenops nepenthicola is a species of crab spider. It lives inside the pitchers of a number of lowland Nepenthes pitcher plants in Malaysia, Indonesia and Singapore. As such, it is classified as a nepenthephile. They are slow-moving spiders which do not actively hunt. Males and females both...

, a crab spider
Crab spider
Crab spider is a common name applied loosely to many species of spiders, but most nearly consistently to members of the family Thomisidae...

 that feeds on flies caught by Nepenthes, and Geosesarma malayanum
Geosesarma malayanum
Geosesarma malayanum is a species of small red crab which is 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 and raid the pitchers of their contents. It uses its claws...

, a species of crab
Crab
True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...

.

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 prey, when it may be colonised by the larvae of various fly
Fly
True flies are insects of the order Diptera . They possess a pair of wings on the mesothorax and a pair of halteres, derived from the hind wings, on the metathorax...

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

Diversity

In a 1991 study, a wide diversity of animals was found in pitchers of Nepenthes ampullaria
Nepenthes ampullaria
Nepenthes ampullaria , the Flask-Shaped Pitcher-Plant, is a very distinctive and widespread species of Nepenthes, present in Borneo, Sumatra, Thailand, Peninsular Malaysia, Singapore, the Maluku Islands, and New Guinea...

:

Carrion feeders:
  • Endonepenthia (Phoridae)
  • Syrphidae


Filter feeders:
  • Aedes albopictus, Culex coerulescens, Culex hewitti, Culex navalis, Tripteroides nepenthis, Tripteroides tenax, Uranotaenia moultoni
    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 pitchers of Nepenthes species, especially N. rajah. As such, it is considered a...

    (Culicidae)


Detritus feeders:
  • several Anoetidae
  • Dasyhelea
    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 and adults do not feed on vertebrate blood, nor do they prey on other insects...

    , Forcipomyia (Ceratopogonidae
    Ceratopogonidae
    Ceratopogonidae, or biting midges , are a family of small flies in the order Diptera...

    )
  • several Chironomidae
    Chironomidae
    Chironomidae are a family of nematoceran flies with a global distribution. They are closely related to the Ceratopogonidae, Simuliidae, and Thaumaleidae...

  • Harpacticoida
    Harpacticoida
    Harpacticoida is an order of copepods, in the Subphylum Crustacea. This order comprises 463 genera and about 3,000 species. Members of it are benthic copepods found throughout the world in the marine environment and in fresh water...

  • Sciaridae
    Sciaridae
    Sciaridae is 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 studied of the large Diptera families, probably due to the small...



Nipping predators (can swim in the fluid, mandibles move against one another in a horizontal plane):
  • Aedes brevitibia, Toxorhynchites indicus (Culicidae)
  • Corethrella calathicola (Chaoboridae
    Chaoboridae
    Chaoboridae, commonly known as phantom midges, are a family of fairly common midges with a cosmopolitan distribution. They are closely related to Corethrellidae and Chironomidae; the adults are differentiated through peculiarities in wing venation. If they eat at all, the adults feed on nectar. The...

    )


Hooking predators (can only crawl, mandibles move parallel to one another in a vertical plane):
  • Dolichopodidae
    Dolichopodidae
    Dolichopodidae, the long-legged flies, make up a large family of true flies with more than 7,000 described species in about 230 genera distributed worldwide. The genus Dolichopus is the most speciose, with some 600 species. They are generally small flies with large, prominent eyes and a metallic...

  • Listodiplosis (Cecidomyiidae
    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.These are very fragile small insects usually only 2–3 mm in length and many are less than...

    )
  • Nepenthosyrphus (Syrphidae)


Terrestrial predators:
  • Misumenops
    Misumenops
    Misumenops is a common genus of crab spider with more than 100 described species.-Distribution:Most species of this genus occur in the Americas from Canada to Argentina, with only a few exceptions:...

    (Thomisidae)
  • Xenoplatyura (Mycetophilidae
    Mycetophilidae
    Mycetophilidae is a family of small flies, forming the bulk of those species known as fungus gnats. There are approximately 3000 described species in 150 genera but the true number of species is undoubtedly much higher...

    )


Nepenthebiont Diptera
Diptera
Diptera , or true flies, is the order of insects possessing only a single pair of wings on the mesothorax; the metathorax bears a pair of drumstick like structures called the halteres, the remnants of the hind wings. It is a large order, containing an estimated 240,000 species, although under half...

:
  • Aedes brevitibia
  • several Chironomidae
    Chironomidae
    Chironomidae are a family of nematoceran flies with a global distribution. They are closely related to the Ceratopogonidae, Simuliidae, and Thaumaleidae...

  • Corethrella calathicola
  • Culex hewitti
  • Culex coerulescens
  • Culex navalis
  • several Dasyhelea
    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 and adults do not feed on vertebrate blood, nor do they prey on other insects...

    species (D. ampullariae, D. biseriata and others)
  • Dolichopodidae
    Dolichopodidae
    Dolichopodidae, the long-legged flies, make up a large family of true flies with more than 7,000 described species in about 230 genera distributed worldwide. The genus Dolichopus is the most speciose, with some 600 species. They are generally small flies with large, prominent eyes and a metallic...

  • Endonepenthia
  • Nepenthosyrphus
  • Toxorhynchites indicus
  • Tripteroides nepenthis
  • Tripteroides tenax
  • Uranotaenia moultoni
    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 pitchers of Nepenthes species, especially N. rajah. As such, it is considered a...


See also

  • Tripteroides sp. No. 2
    Tripteroides sp. No. 2
    Tripteroides sp. No. 2 is a species of mosquito in the genus Tripteroides which has not yet been formally named. It is endemic to Sabah, Malaysian Borneo. T. sp. No. 2 is placed in the subgenus Rachionotomyia. In its larval stage, T. sp. No. 2 develops in the pitchers of Nepenthes species,...

  • Phytotelmata
    Phytotelmata
    Phytotelma is a term for water bodies held by plants. The water accumulated on these plants may serve as substratum for associated fauna, and often the fauna associated with phytotelmata is unique....

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