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The pitchers of N. rajah are some of the largest Nepenthes phytotelmata, capable of holding 2.5 litres of fluid[1]

This list of Nepenthes pitcher inhabitants and visitors is a listing of organisms recorded from the traps of Nepenthes pitcher plants. It includes true infaunal organisms—those that inhabit the pitcher fluid as part of their life cycles or enter it to forage—as well as organisms that are found inside or on the pitchers but above the fluid level. It excludes pitcher visitors that typically fall prey to the plant, as well as those that feed on the pitchers (such as leaf miners and weevils). That is, only examples of putative ecological facilitation are listed, where the interspecific relationship is thought to be either commensal or mutualistic.

Around 100 different species have been recorded from Nepenthes pitchers. Some of these are specialised to life inside Nepenthes pitchers, while others are generalists that colonise any suitable body of water.

Species are ordered alphabetically by genus, with indeterminate identifications included at the end. The following information is included in brackets after the taxon name:

  • Host Nepenthes taxa from which the pitcher associate has been recorded.
  • Known geographical extent of this association. For non-nepenthebionts, this may be smaller than their entire natural range. Generally follows Roger A. Beaver (1983),[2] with additional categories added to accommodate new records.
  • Ecological group to which the pitcher associate belongs, if known. This classification follows August Thienemann, who divided Nepenthes infauna into three groups based on the intimacy and necessity of their associations:[3][4]
    • Nepenthebionts — organisms which are specialised to life inside Nepenthes pitchers and are totally dependent on them at least at some stage of their lives. Obligate pitcher dwellers.
    • Nepenthephiles — organisms which are frequently found in Nepenthes pitchers, but which are not completely dependent on them at any stage of their lives. Facultative pitcher dwellers.
    • Nepenthexenes — organisms which are not normally associated with pitchers, but which are occasionally encountered in them. Accidental pitcher associates.
  • Additional information on the nature of the organism's association with Nepenthes, including relevant behaviour.

In cases where the originally published name of an infaunal or host species differs from that currently recognised, the former is indicated in square brackets following the accepted name. Similarly, where a misidentification has taken place with respect to the infaunal or host species, the originally reported name is included in square brackets following the one currently accepted. Homotypic (objective) synonyms are indicated by a triple bar (≡) and heterotypic (subjective) synonyms by an equals sign (=); a lack of either sign indicates a misidentification.

The list largely follows similar compilations of Nepenthes infauna by Beaver (1983)[2] and Adlassnig et al. (2011).[5]

Fluid in unopened pitchers has antimicrobial properties and has long been thought to be sterile[6] (but see [7]).

Mammals[edit]

Kerivoula hardwickii bats roost in the expanded upper portion of N. hemsleyana aerial pitchers
Rattus baluensis feeding on nectar from a pitcher of N. rajah

Bats[edit]

Vespertilionidae (vesper bats, evening bats, common bats)
  • Kerivoula
    • Kerivoula hardwickii (N. bicalcarata,[8] N. hemsleyana[9] [previously called N. baramensis and "N. rafflesiana var. elongata"];[10] roost in upper pitchers of N. hemsleyana, with faeces accounting for an estimated 33.8% of total foliar nitrogen;[11] vocalisations of K. hardwickii (highest frequency known among bat species) are likely adaptation to locate pitchers among dense surrounding vegetation;[12] highly echo-reflective parabolic structure in rear wall of N. hemsleyana upper pitchers may further aid bats in finding pitchers;[12] N. hemsleyana pitchers are less abundant than those of N. bicalcarata but provide higher quality roosts (bats have higher body condition and fewer parasites), and are occupied longer and chosen preferentially;[8] relationship appears to be mutualistic in N. hemsleyana and commensal or parasitic in N. bicalcarata[8])[13][14]
Incertae familiae

Primates[edit]

Tarsiidae (tarsiers)

Rodents[edit]

Muridae

Tree shrews[edit]

Tupaiidae

Birds[edit]

Nectariniidae (sunbirds, spiderhunters)
Zosteropidae (white-eyes)

Reptiles[edit]

Nepenthes madagascariensis with Phelsuma quadriocellata

Lizards[edit]

Scincidae (skinks)

Amphibians[edit]

Heterixalus boettgeri in a dried upper pitcher of N. madagascariensis
A damaged lower pitcher of N. tentaculata containing frog spawn
Kalophrynus pleurostigma has been recorded from N. ampullaria in Singapore
Bufonidae (true toads)
Dendrobatidae (poison dart frogs)
Hyperoliidae
Microhylidae (narrow-mouthed frogs)
Rhacophoridae (shrub frogs)
Incertae familiae

Insects[edit]

  • Undetermined pupae (N. pitopangii; Sulawesi; attached to upper portion of inner pitcher surface; only observed in lower pitchers; may belong to leaf miner that is also found only in lower pitchers of N. pitopangii)[55]

Dragonflies[edit]

Libellulidae (skimmers, perchers)

Flies[edit]

Calliphoridae (blow flies, cluster flies, bluebottles, greenbottles)
Cecidomyiidae (gall midges, gall gnats)
Ceratopogonidae (biting midges)
Chironomidae (non-biting midges)
Chloropidae (frit flies, grass flies)
Corethrellidae (frog-biting midges)
A lower pitcher of N. attenboroughii supporting a population of mosquito larvae. The upright lid of this species means that its pitchers are open to the elements and thus often completely filled with fluid.[71]
Nepenthes bicalcarata (A) with associated C. schmitzi ants (B) and various infaunal fly larvae (C), including Culex, Polypedilum, Toxorhynchites, Tripteroides, Uranotaenia, and Wilhelmina nepenthicola
Camponotus schmitzi ants hunting mosquito pupae in a pitcher of Nepenthes bicalcarata
Culicidae (mosquitoes)
Dolichopodidae (long-legged flies)
Keroplatidae / Mycetophilidae (fungus gnats)
Lauxaniidae
  • Undetermined Lauxaniidae (N. sp.; Java; ecological group unknown; family identification doubtful[2])[96]
Muscidae (stable flies, house flies, face flies)
Phoridae (scuttle flies)
Sarcophagidae (flesh flies)
Sciaridae (dark-winged fungus gnats)
Camponotus schmitzi ant hunting a large fly larva of the genus Eristalis
Syrphidae (hoverflies, flower flies)
Tachinidae (tachina flies)








Some species of mosquito[70][104] are only known from their pitcher plant hosts and have been named after them.




Infaunal mosquito larvae in the digestive fluid of N. rajah. Culex rajah and Toxorhynchites rajah were described from the pitchers of this species and are named after it.[77]


At least in some genera, the eggs of mosquitoes that breed in Nepenthes pitchers differs from those that colonise artificial containers.[86][105]

Larvae of the syrphid fly Nepenthosyrphus, which are hooking predators, have been recorded from N. ampullaria[63] and N. bicalcarata pitchers, although they are considered relatively rare pitcher inhabitants.[64]

Frit fly larvae of the family Chloropidae.[35] Syrphid fly larvae of the genus Nepenthosyrphus are subaquatic predators in Nepenthes pitchers, feeding on mosquito larvae and other insects. They are thought to have evolved from detritivorous ancestors.[106] The phorid fly Megaselia campylonympha feeds on recently drowned insects.[15]


Flies of the genus Endonepenthia (family Phoridae) are carrion feeders.[63] Biting midges of the genera Dasyhelea (D. ampullariae, D. biseriata, and others) and Forcipomyia are detritivorous members of the pitcher infauna.[63] 5 species of Dasyhelea are known to breed in Nepenthes pitchers.[107] The non-biting midge

Gall midges of the genus Listodiplosis are known to be hooking predators in the infaunal foodweb of N. ampullaria.[63]

The larva of the fungus gnat Xenoplatyura beaveri (family Mycetophilidae) is a terrestrial predator in N. ampullaria pitchers.[94] It forms a web across the mouth of the pitcher which may completely block the opening. Insects caught in this web are killed by the highly acidic sticky droplets that the larva secrets onto it. Infaunal insects emerging from the pitcher fluid are likely to constitute an important source of food.[45] The larva has also been seen fishing out small insects from the surface of the fluid.[108] A species of Xenoplatyura has also been recorded from N. bicalcarata, although the gnat is apparently only present in pitchers that have not been colonised by the ant Camponotus schmitzi, another terrestrial predator of the pitcher foodweb.[64]

Hymenopterans[edit]

An intermediate pitcher of N. bicalcarata with a swollen tendril colonised by Camponotus schmitzi
Camponotus schmitzi and Alcidodes sp. in association with N. bicalcarata
Nepenthes gracilis upper pitcher with Polyrhachis pruinosa
Diapriidae
Elachertidae
Encyrtidae
Formicidae (ants)
Ichneumonidae (ichneumon wasps, scorpion wasps)
Proctotrupidae

Mantises[edit]

Nepenthes madagascariensis with undetermined Mantidae
Nepenthes madagascariensis with undetermined Mantidae
Empusidae
Mantidae

Moths[edit]

Nepenthes madagascariensis with undetermined Arctiidae caterpillar
Gracillariidae
Noctuidae (owlet moths)
Psychidae (bagworm moths)

Springtails[edit]

Podura aquatica on the surface of a pool
Poduridae

Arachnids[edit]

Mites[edit]

Acaridae
Histiostomatidae


[157][158][159]


Some mite species inhabit a number of different Nepenthes species, while others are restricted to only one.[160]

Spiders[edit]

Nepenthes madagascariensis with Peucetia madagascariensis
Nepenthes madagascariensis with Peucetia madagascariensis
Nepenthes madagascariensis with undetermined Thomisidae
Nepenthes madagascariensis with undetermined Thomisidae
Araneidae (orb-weaver spiders)
Lycosidae (wolf spiders)
Oxyopidae (lynx spiders)
  • Peucetia
    • Peucetia madagascariensis (N. madagascariensis; Madagascar; third most common spider associate of N. madagascariensis after Synema and ?Salticus; observed on pitchers, laminae, and inflorescences; one plant found to harbour an adult with at least 20 juveniles; prey (including pollinators) partly caught by means of spider web, which may cover entire inflorescence; one older juvenile found to have spun web around orifice of pitcher inhabited by Synema sp.; also found on other plants)[140]
    • Peucetia sp. (N. madagascariensis; Madagascar)[35]
Pholcidae (cellar spiders)
Salticidae (jumping spiders)
Tetragnathidae (long-jawed orb weavers, long jawed spiders)
Theridiidae (tangle-web spiders, cobweb spiders, comb-footed spiders)
Thomisidae (crab spiders)
Incertae familiae
  • Undetermined "large spiders" (N. sp. Anipahan; Mount Anipahan, Philippines; observed inside several upper pitchers, perched near opening;[92] climbed out of pitchers when disturbed[171])
  • Undetermined small spiders (N. leonardoi; Philippines)[93]
  • Undetermined spiders (not Misumenops) (N. mirabilis; New Guinea; build webs in upper portion of damaged and drained pitchers; prey on insects that fall in; utilise basal holes chewed by undetermined phytophagous insects for escape; "not normal inhabitants of Nepenthes pitchers")[16]



Misumenops nepenthicola N. albomarginata N. gracilis N. gymnamphora N. rafflesiana N. reinwardtiana N. tobaica Beaver, 1983 Juniper et al., 1989 Choo et al., 1997


Undetermined spiders (N. mirabilis) Hua, 2004

Hua YJ. 2004. Nepenthes and its food web. In: Society ICP ed. 5th I. C. P. S. Congress. Lyon, International Carnivorous Plant Society.

Crustaceans[edit]

Branchiopods[edit]

Incertae familiae

Copepods[edit]

Canthocamptidae
Parastenocarididae
Phyllognathopodidae
Incertae familiae

Decapods[edit]

Sesarmidae
Incertae familiae

Molluscs[edit]

Incertae familiae

Annelids[edit]

Incertae familiae

Nematodes[edit]

The most comprehensive study of infaunal nematodes was carried out in 1922 based on the pitchers of N. gymnamphora[note a]
Anguinidae
Diplogastridae
Dorylaimidae
Panagrolaimidae
Plectidae
Rhabditidae
Incertae familiae

Rotifers[edit]

Incertae familiae

Protozoans[edit]

Amoebozoans[edit]

Amoebidae
Arcellidae
Centropyxidae
Cochliopodiidae
Difflugiidae
Lesquereusiidae
Microcoryciidae
Incertae familiae

Ciliates[edit]

Incertae familiae

Fungi[edit]

Higher fungi[edit]

Entylomataceae
Sporidiobolaceae
Tremellaceae
Trichosporonaceae

Pin molds[edit]

Mucoraceae

Sac fungi[edit]

Dothioraceae
Saccharomycetaceae
Trichocomaceae

Algae[edit]

Diatoms[edit]

Achnanthaceae
Cocconeidaceae
Naviculaceae
Rhopalodiaceae

Golden algae[edit]

Incertae familiae

Green algae[edit]

Desmidiaceae

Bacteria[edit]

Cyanobacteria[edit]

Merismopediaceae

Firmicutes[edit]

Bacillaceae
  • Morohoshi, T., M. Oikawa, S. Sato, N. Kikuchi, N. Kato & T. Ikeda 2011. Isolation and characterization of novel lipases from a metagenomic library of the microbial community in the pitcher fluid of the carnivorous plant Nepenthes hybrida. Journal of Bioscience and Bioengineering 112(4): 315–320. doi:10.1016/j.jbiosc.2011.06.010
  • Uncultured bacterium lip1 genes for lipase, putative ABC transporter, partial and complete cds, clone: pNEL01
  • Uncultured bacterium lip2 gene for lipase, complete cds, clone: pNEL02
  • Priliani, L., A. Siegara & Yogiara N.d. Genetic diversity of bacterial community associated with two pitcher plant fluid (Nepenthes spp.) grown in nursery. Unpublished. [sequences published on GenBank]

Bacteria are by far the most numerous inhabitants of Nepenthes pitchers. One study found 26 strains of bacteria in only 4 pitchers. Of these, 10 strains were gram-positive and 16 were gram-negative, and 10 showed casein hydrolase activity.[189]

One study found between 48,000 and 8,000,000 rod-shaped bacteria per cubic centimetre of pitcher fluid. They form a symbiotic relationship with Nepenthes, gaining nutrients from captured prey and in return assisting, to a limited degree, in their digestion.[190]

see also [7]

Notes[edit]

a.^ Endonepenthia was formally synonymised with Megaselia in 1981,[98] but some authors choose to treat them separately since Endonepenthia differs ecologically from Megaselia sensu stricto in that it concentrates in young pitchers.[63][61]
b.^ In his 1988 book Stranger in the Forest, Eric Hansen attributes a similar ant–plant relationship to N. lowii:[191]

Weng [a Penan guide from the upper Malinau River] told me the story of a diving ant that launches itself from the rim of Lowes [sic] pitcher plant (Nepenthes lowii) and plunges into the insect-eating reservoir of digestive fluid contained within the body of the plant. The diving ant rescues some of the insects by "swimming" them to the edge of the reservoir like a miniature lifesaver. Then the ant eats the insect.

c.^ The upper pitchers of N. lowii secrete a sugary, white substance that accumulates among the bristles on the exposed lower surface of the lid. In the 1960s, J. Harrison assumed that these white beads were snail eggs.[192] E. J. H. Corner, who led the 1961 and 1964 Royal Society Expeditions to Mount Kinabalu, wrote:[193][194]

In the early morning there is a ringing gonging which we traced to tupaias (tree-shrews) scampering over the pitchers of N. lowii and banging the old, empty and resonant, pitchers together. The late Professor J. Harrison, of Singapore, discovered that a snail laid its eggs in the hairs under the lid and that the tupaias came to eat them.

It is now known that these white beads are of the plant's own production and serve to attract tree shrews, from whose droppings they derive most of their foliar nitrogen.[22]

References[edit]

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Further reading[edit]

  • Adam, J.H., C.C. Wilcock & M.D. Swaine 1994. Short notes on the ecology of Bornean Nepenthes. Sumber 8: 99–101.
  • Bordoloi, R.P.M. 1977. The Pitcher Plant: Nepenthes khasiana. Carnivorous plants of North East India I. Dutta Baruah, Gauhati.
  • Clarke, C. 1998. A re-examination of geographical variation in Nepenthes food webs. Ecography 21(4): 430–436. doi:10.1111/j.1600-0587.1998.tb00408.x
  • Clarke, C. 2002. Twelve years of ecological research on Nepenthes in Southeast Asia - some personal highlights. Proceedings of the 4th International Carnivorous Plant Conference, Hiroshima University, Tokyo: 1–7.
  • Clarke, C.M. & J. Moran 2001. The Inhabitants of Nepenthes Pitchers. In: C.M. Clarke Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu. pp. 61–68.
  • Coesel, P.F.M. 2002. Taxonomic and biogeographical notes on Malagassy desmids (Chlorophyta, Desmidiaceae). Nordic Journal of Botany 22(2): 239–256. doi:10.1111/j.1756-1051.2002.tb01372.x
  • Edwards, F.W. 1925. Mosquito notes—V. Bulletin of Entomological Research 15: 257–270. doi:10.1017/S0007485300046204
  • Fish, D. & R.A. Beaver 1979. A bibliography of the aquatic fauna inhabiting bromeliads (Bromeliaceae) and pitcher plants (Nepenthaceae and Sarraceniaceae). Proceedings of the Florida Anti-Mosquito Association (19th meeting, April 1978) 49: 11–19.
  • (in German) Guenther, K. 1913. Die lebenden Bewohner der Kannen der insektenfressenden Pflanze Nepenthes destillatoria auf Ceylon. Zeitschrift für wissenschaftliche Insektenbiologie 9: 122–130, 156–160, 198–207.
  • Jensen, H. 1910. Nepenthes-Tiere II. Biologische Notizen. Annales du Jardin Botanique de Buitenzorg 3(supplement): 941–946.
  • Kitching, R.L. & C.J. Schofield 1986. Every pitcher tells a story. New Scientist 109(1492): 48–50.
  • Mey, F.S., L.H. Truong, D.V. Dai & A.S. Robinson 2011. Nepenthes thorelii, an emended description and novel ecological data resulting from its rediscovery in Tay Ninh, Vietnam. McPherson, S.R. New Nepenthes: Volume One. Redfern Natural History Productions, Poole. pp. 104–131.
  • Mogi, M. & K.L. Chan 1996. "Predatory habits of dipteran larvae inhabiting Nepenthes pitchers" (PDF). The Raffles Bulletin of Zoology 44(1): 233–245.
  • Paulian, R. 1950. Les moustiques des Nepenthes à Madagascar. Naturaliste Malgache 2: 119–121.
  • Pimm, S.L., J.H. Lawton & J.E. Cohen 1991. "Food web patterns and their consequences" (PDF). Nature 350(6320): 669–674. doi:10.1038/350669a0
  • Pollard, S.D. 2005. Bugs in jugs. Nature Australia 28: 68–69.
  • Rembold, K., E. Fischer, B.F. Striffler & W. Barthlott 2013. Crab spider association with the Malagasy pitcher plant Nepenthes madagascariensis. African Journal of Ecology 51(1): 188–191. doi:10.1111/aje.12037
  • (in German) Schmid-Hollinger, R. 2010. Nepenthes macfarlanei: Ameisennest in alter Kanne. Das Taublatt 66: 39–42.
  • Striffler, B.F. & K. Rembold 2009. Henriksenia labuanica nom. nov., a replacement name for Misumenops nepenthicola Bristowe (1930) and clarification of the current taxonomic status of Misumenops nepenthicola Fage (1928) (Arachnida, Araneae, Thomisidae). In: S.R. McPherson Pitcher Plants of the Old World. Volume 2. Redfern Natural History Productions, Poole. pp. 1334–1337.

[[:Category:Nepenthes infauna| *]] [[:Category:Lists of biota]]

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