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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.
- Borneo — encompasses all records from Brunei, the Malaysian states of Sabah and Sarawak, and the Indonesian portion of the island (Kalimantan).
- Java
- Madagascar
- Malaya — includes Peninsular Malaysia and Singapore
- New Guinea — includes Papua New Guinea and Indonesian-administered Western New Guinea
- Seychelles
- Sri Lanka
- Sumatra
- Thailand
- 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]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
- Undetermined bats (N. bicalcarata and N. rafflesiana, found roosting in pitchers; unknown whether this behaviour is common)[15]
Primates
[edit]- Tarsiidae (tarsiers)
- Tarsius
- "Tarsius spectrum" (various N. spp.; Sabah, Borneo; observed sitting on peristome and feeding on trapped insects;[16] T. spectrum is now considered to be a junior synonym of T. tarsier, which is absent from Borneo[17])[18]
Rodents
[edit]- Muridae
- Rattus
- Rattus baluensis (N. rajah; Borneo)[19][20]
Tree shrews
[edit]- Tupaiidae
- Tupaia
- Tupaia montana (N. lowii, N. macrophylla, N. rajah, N. ephippiata? [suspected]; Borneo; colour of lower lid surface, particularly that of N. rajah, corresponds to T. montana visual sensitivity maxima in the green and blue wavebands[21])[22][23][24][25][19][20][26]
Birds
[edit]- Nectariniidae (sunbirds, spiderhunters)
- Aethopyga
- Aethopyga sp. (N. faizaliana; Mount Api, Borneo; observed sitting on peristome of upper pitcher, feeding on nectar from underside of lid; not known to contribute to pitcher contents)[15][27]
- Zosteropidae (white-eyes)
- Chlorocharis
- Chlorocharis emiliae (N. lowii; Borneo; observed sitting on rim of upper pitcher, feeding on nectar from underside of lid;[28] occasional visitors; not known to contribute to pitcher contents[29])
Reptiles
[edit]Lizards
[edit]- Scincidae (skinks)
- Lioscincus
- Lioscincus tillieri (N. vieillardii; New Caledonia; observed lying in wait for prey on pitcher lid)[30]
Amphibians
[edit]- Bufonidae (true toads)
- Bufo
- ?Bufo melanostictus/Bufo sp.[31] (N. ampullaria in Singapore;[31] doubtful identification[32])
- Pelophryne
- Pelophryne linanitensis (N. muluensis, N. murudensis; Mount Murud, Borneo) - suspected[33]
- Pelophryne murudensis (N. muluensis, N. murudensis; Mount Murud, Borneo) - suspected[33]
- Dendrobatidae (poison dart frogs)
- Dendrobates
- Dendrobates spp. (N. spp.)[34]
- Hyperoliidae
- Heterixalus
- Heterixalus tricolor (N. madagascariensis; Madagascar)[35]
- Microhylidae (narrow-mouthed frogs)
- Kalophrynus
- Kalophrynus pleurostigma (N. ampullaria; Singapore, Malaya)[32][36][37]
- Kalophrynus yongi (N. macfarlanei; Mount Brinchang, Borneo; eggs and larvae found in large pitcher; eggs attached to inner surface above fluid level)[38]
- ?Kalophrynus sp. (N. sp.; Kuching, Borneo; tadpoles observed in pitcher; originally identified as Microhyla borneensis by Parker (1934);[39][32] later recognised as an undescribed species of Kalophrynus by Inger (1966),[40] but Matsui (2009) wrote that this "generic identification remained questionable"[38])
- Microhyla
- Microhyla borneensis — see Kalophrynus sp.
- Microhyla nepenthicola (N. ampullaria; Matang Range, Sarawak, Borneo)[41]
- Rhacophoridae (shrub frogs)
- Philautus
- ?Philautus aurantium (N. fusca; Mount Trusmadi, Borneo; found inside partly-dried pitcher)[42]
- ?Philautus aurifasciatus (N. sanguinea; Genting Highlands, Peninsular Malaysia)[43][32]
- Philautus mjobergi (N. villosa; Borneo)[44][45]
- Philautus saueri (N. villosa; Borneo; occasionally lay eggs in dead pitchers)[46]
- Philautus small golden-coloured species (N. hurrelliana; Mount Lumarku, Borneo; observed perched on peristome)[28][47]
- Philautus sp. (N. bicalcarata; Borneo; lay eggs in pitchers)[48]
- Philautus sp. (N. × harryana; Mount Kinabalu, Borneo; observed emerging from pitcher)[49]
- Incertae familiae
- Undetermined tadpoles (N. ampullaria, N. bicalcarata)[50][51]
- Undetermined tree frogs (N. mirabilis; southern China; feed on insects attracted to pitchers; retreat into pitcher fluid when disturbed)[52][53]
- Undetermined tree frogs (cultivated N. alata, other N. spp.; Florida, United States; retreat into pitcher when disturbed)[54]
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)
- Nepenthomyia
- Nepenthomyia malayana[58] (N. ampullaria; Malaya; nepenthebiont;[2] larvae move along inner pitcher surface, destroying webs of Xenoplatyura beaveri; rarely found in the same pitchers as X. beaveri[59])[60][16]
- Nepenthomyia sp. (N. ampullaria; Singapore)[61]
- Wilhelmina
- Wilhelmina nepenthicola (N. ampullaria,[2] N. eustachya [N. alata],[62] N. rafflesiana; Borneo, Sumatra; nepenthebiont[2])
- Cecidomyiidae (gall midges, gall gnats)
- Lestodiplosis
- Lestodiplosis syringopais (N. albomarginata, N. gracilis, N. tobaica; Malaya, Sumatra; nepenthebiont)[2]
- Lestodiplosis spp. (N. ampullaria, N. eustachya [N. alata],[62] N. gracilis, N. mirabilis, N. rafflesiana; aquatic predator)[63][64][65][66]
- Ceratopogonidae (biting midges)
- Culicoides
- Dasyhelea
- Dasyhelea ampullariae (N. ampullaria, N. gracilis; Malaya; nepenthebiont)[2]
- Dasyhelea biseriata (N. ampullaria; Malaya; nepenthebiont)[2]
- Dasyhelea confinis (N. mirabilis; Sumatra; nepenthebiont)[2]
- Dasyhelea nepenthicola (N. albomarginata, N. ampullaria, N. gracilis; Malaya; nepenthebiont)[2]
- Dasyhelea subgrata (N. mirabilis; New Guinea; nepenthebiont)[2]
- Dasyhelea spp. (N. albomarginata, N. ampullaria, N. bicalcarata, N. eustachya [N. alata],[62] N. gracilis,[66] N. mirabilis, N. rafflesiana; detritus feeder)[67][63][50][51][34]
- Forcipomyia
- Forcipomyia sp. (N. ampullaria)[63]
- Undetermined Ceratopogonidae (N. mirabilis; Hong Kong; ecological group unknown)[2]
- Undetermined Ceratopogonidae, Phoridae, and Syrphidae (N. albomarginata, N. ampullaria, ?N. eustachya [N. alata], N. gracilis, N. mirabilis, N. reinwardtiana, N. spathulata, "N. sp. A", "N. sp. B"; West Sumatra, Sumatra; detritus feeders)[68]
- Chironomidae (non-biting midges)
- Metriocnemus
- Metriocnemus sp. (N. ampullaria, N. tentaculata, N. cf. villosa,[34] N. sp.; Borneo; ecological group unknown[2])
- Polypedilum
- Polypedilum convexum (N. ampullaria, N. bicalcarata; detritus feeder[15])[34]
- Polypedilum sp. (N. ampullaria, N. bicalcarata, N. cf. villosa, N. spp.)[65][50][51][34]
- Undetermined Chironomidae (N. mirabilis; Hong Kong; ecological group unknown)[2]
- Undetermined Chironomidae (N. ampullaria)[63]
- Chloropidae (frit flies, grass flies)
- Undetermined Chloropidae (N. madagascariensis; Madagascar; ecological group unknown)[69][2][35]
- Corethrellidae (frog-biting midges)
- Corethrella
- Corethrella calathicola (N. ampullaria; Borneo, Malaya; nepenthebiont[2])[63][70]
- Corethrella spp. (N. ampullaria, N. bicalcarata)[50][51]
- Culicidae (mosquitoes)
- Aedes
- Aedes albopictus (N. ampullaria;[67][70][2][63] Malaya;[2] cultivated N. ventricosa in Virginia, United States;[72] nepenthexene;[2] only medically important mosquito recorded from Nepenthes[16][59])
- Aedes brevitibia (N. ampullaria, N. bicalcarata, N. rafflesiana; Borneo; nepenthebiont[2])[63]
- Aedes dybasi (N. mirabilis; Palau; nepenthephile[2])[73][74][75]
- Aedes gani (N. sp.; New Guinea; nepenthebiont)[2]
- Aedes medialis (N. sp.; New Guinea; nepenthebiont)[2]
- Aedes maehleri (N. mirabilis; Yap, Micronesia; nepenthebiont[2])[73][74][75]
- Aedes treubi (N. gymnamphora; Java; nepenthebiont)[2]
- Anopheles
- Armigeres
- Armigeres conjugens (N. sp.; Malaya; nepenthexene)[2]
- Armigeres durhami (N. ampullaria; Malaya; nepenthexene;[2] accidentals in old pitchers in Singapore[70])
- Armigeres flavus (N. sp.; Malaya; nepenthexene)[2]
- Armigeres giveni (N. ampullaria, N. rafflesiana; Malaya; nepenthebiont;[2] commonly encountered in living pitchers in Singapore[70])[67]
- Armigeres hybridus (N. sp.; Malaya; nepenthexene)[2]
- Armigeres kuchingensis[76] (N. ampullaria; Malaya; nepenthexene;[2] accidentals in old pitchers in Singapore[70])
- Armigeres magnus (N. mirabilis; Hong Kong; nepenthephile?)[2]
- Armigeres malayi (N. sp.; Malaya; nepenthexene)[2]
- Culex
- Culex acutipalus (N. ampullaria; Malaya; nepenthebiont[2])[63]
- Culex brevipalpus (N. sp.; Malaya; nepenthebiont)[2]
- Culex coerulescens (N. ampullaria, N. bicalcarata, N. spp.;[70][63][65][50][51] Borneo, Malaya; nepenthebiont[2])
- Culex curtipalpis (N. albomarginata,[2] N. eustachya [N. alata],[62] N. gracilis, N. mirabilis; Borneo, Malaya, Sumatra, Thailand; nepenthebiont[2])[70]
- Culex eminentia (N. ampullaria, N. gracilis, N. rafflesiana; Borneo, Malaya; nepenthebiont;[2] filter feeder[61])[70]
- Culex hewitti (N. ampullaria; Borneo, Malaya; nepenthebiont[2])[70][63]
- Culex jenseni (N. gymnamphora, N. reinwardtiana; Java, Malaya, Sumatra; nepenthebiont[2])[77]
- Culex lucaris (N. albomarginata; Malaya, Thailand; nepenthebiont)[2]
- Culex navalis (N. ampullaria, N. bicalcarata,[2] N. eustachya [N. alata],[62] N. rafflesiana; Borneo, Malaya; nepenthebiont[2])[67][70][63]
- Culex rajah (N. rajah)[77]
- Culex shebbearei (N. rajah;[78][2] Borneo;[2] nepenthephile?;[2] doubtful record - probably a misidentified C. rajah[77])
- Culex sumatranus (N. mirabilis; Hong Kong; nepenthephile?)[2]
- Culex spp. (N. gracilis; Singapore, Malaya; filter feeder)[66]
- Culex spp. (N. stenophylla; Sarawak, Borneo; found in association with Topomyia nepenthicola)[79]
- Megarhinus — see Toxorhynchites
- Mimomyia
- Mimomyia jeansottei (N. madagascariensis; Madagascar; nepenthexene)[2]
- Topomyia
- Topomyia nepenthicola (N. stenophylla; Sarawak, Borneo; nepenthebiont?; aquatic predator; not found in sympatric N. gracilis and N. veitchii; found in association with Culex spp., Toxorhynchites sp., Tripteroides spp. and Uranotaenia spp.; extended aquatic cycle under laboratory conditions)[79]
- Toxorhynchites
- Toxorhynchites acaudatus [≡Megarhinus acaudatus] (N. ampullaria, N. rafflesiana; Malaya; nepenthebiont?;[2] larvae prey on larvae of other species and cannibalise each other, leaving only one larva per pitcher[60][59])[67][70][61]
- Toxorhynchites ater (N. rafflesiana; Malaya; nepenthebiont?[2])[80]
- Toxorhynchites aurifluus (N. sp.; Sumatra; nepenthexene?; may be misidentification[2])[16]
- Toxorhynchites coeruleus (N. sp.; Sumatra; nepenthebiont?)[2][16]
- Toxorhynchites indicus (N. ampullaria)[63]
- Toxorhynchites klossi (N. albomarginata; Malaya; nepenthebiont)[2]
- Toxorhynchites metallicus [≡Megarhinus metallicus][81][16] (N. sanguinea; Malaya; nepenthexene?; may be misidentification)[2]
- Toxorhynchites nepenthicola (N. sp.)[82][2][16]
- Toxorhynchites nepenthis (N. sp.; Philippines; nepenthebiont[2])[83][16]
- Toxorhynchites nigripes (?N. sp.; Borneo; ecological group unknown[2])[80][16]
- Toxorhynchites pendleburyi (N. sp.; Borneo, Malaya; nepenthebiont?[2])[16]
- Toxorhynchites quasiferox (N. sp.; Malaya; nepenthexene[2])[16]
- Toxorhynchites rajah (N. rajah)[77]
- Toxorhynchites splendens (N. ampullaria, N. rafflesiana; Malaya, New Guinea; nepenthexene)[2]
- Toxorhynchites sumatranus (N. sp.; Sumatra; nepenthebiont?[2])[16]
- Toxorhynchites sp. (N. ampullaria, N. bicalcarata, N. eustachya [N. alata],[62] N. rafflesiana)[65][50][51]
- Toxorhynchites sp. (N. campanulata; Gunung Mulu National Park, Borneo; forms simple food web with Zwickia sp. and Tripteroides nepenthis, and preys on latter; unknown whether regular coloniser or accidental)[84]
- Toxorhynchites sp. (N. stenophylla; Sarawak, Borneo; found in association with Topomyia nepenthicola)[79]
- Toxorhynchites spp. (N. albomarginata, N. ampullaria, ?N. eustachya [N. alata], N. gracilis, N. reinwardtiana, N. spathulata, "N. sp. B"; West Sumatra, Sumatra; aquatic predators)[68]
- Tripteroides
- Tripteroides adentata (N. sp.; New Guinea; nepenthebiont?)[2]
- Tripteroides apoensis (N. sp.; Philippines; nepenthebiont)[2]
- Tripteroides aranoides [≡Rachionotomyia aranoides] (N. sanguinea,[81][16] N. sp. (Malaya); nepenthexene[2])[67][85]
- Tripteroides bambusa (N. albomarginata, N. ampullaria, N. gracilis; Malaya; nepenthephile)[2]
- Tripteroides barraudi (N. sp.; Philippines; nepenthebiont)[2]
- Tripteroides belkini (N. sp.; Philippines; nepenthebiont)[2]
- Tripteroides bimaculipes (N. sp.; New Guinea; nepenthephile)[2]
- Tripteroides bisquamatus (N. mirabilis; New Guinea; nepenthephile[2])[16]
- Tripteroides brevirhynchus (N. mirabilis; Australia, New Guinea; nepenthebiont)[2][16]
- Tripteroides caledonicus (N. vieillardii [N. mirabilis];[2][16] New Caledonia; nepenthebiont;[2] elongate-ovoid eggs are possible adaptation to Nepenthes pitchers[86])
- Tripteroides christophersi (N. sp.; Philippines; nepenthebiont)[2]
- Tripteroides cuttsi (N. sp.; New Guinea; nepenthebiont?)[2]
- Tripteroides delpilari (N. sp.; Philippines; nepenthebiont)[2]
- Tripteroides digoelensis (N. sp.; New Guinea; nepenthebiont)[2]
- Tripteroides dofleini (N. distillatoria; Sri Lanka; nepenthebiont)[2]
- Tripteroides dyari (N. sp.; Philippines; nepenthephile?)[2]
- Tripteroides elegans (N. sp.; New Guinea; nepenthebiont?)[2]
- Tripteroides filipes (N. mirabilis; Australia, New Guinea; nepenthebiont[2])[16]
- Tripteroides flabelliger (N. sp.; New Guinea; nepenthebiont?)[2]
- Tripteroides intermediatus (N. sp.; Philippines; nepenthebiont?)[2]
- Tripteroides kingi (N. mirabilis; New Guinea; nepenthebiont?[2])[16]
- Tripteroides longipalpatus (N. sp.; New Guinea; nepenthexene?)[2]
- Tripteroides malvari (N. sp.; Philippines; nepenthebiont?)[2]
- Tripteroides sp. nr. mathesoni (N. sp.; New Guinea; ecological group unknown)[2]
- Tripteroides mendacis (N. sp.; Malaya; nepenthebiont?)[2]
- Tripteroides microcala (N. sp.; Philippines; nepenthebiont)[2]
- Tripteroides cf. microcala (N. sp., Philippines; elongate-ovoid eggs are possible adaptation to Nepenthes pitchers)[87][86]
- Tripteroides microlepis (N. sp.; New Guinea; nepenthebiont?)[2]
- Tripteroides nepenthicola (N. alata; Philippines; nepenthebiont)[2]
- Tripteroides nepenthis [≡Rachionotomyia nepenthis][76] (N. ampullaria, N. bicalcarata, N. campanulata,[84] N. rafflesiana;[70][63][50][51] Borneo, Malaya; nepenthebiont;[2] filter feeder;[61] in N. campanulata in Gunung Mulu National Park, Borneo, forms simple food web with Zwickia sp. and Toxorhynchites sp., and is preyed on by latter[84])[67]
- Tripteroides nepenthisimilis (N. ampullaria, ?N. rafflesiana; Borneo, Malaya; nepenthebiont)[2]
- Tripteroides obscurus (N. sp.; New Guinea; nepenthebiont)[2]
- Tripteroides pallidus (N. sp.; New Guinea; nepenthebiont?)[2]
- Tripteroides papua (N. sp.; New Guinea; nepenthebiont)[2]
- Tripteroides pilosus (N. sp.; New Guinea; nepenthebiont?)[2]
- Tripteroides reiseni (N. sp.; Philippines; nepenthebiont)[2]
- Tripteroides roxasi (N. sp.; Philippines; nepenthebiont)[2]
- Tripteroides simplex (N. sp.; New Guinea; nepenthebiont)[2]
- Tripteroides simulatus (N. sp.; Philippines; nepenthebiont?)[2]
- Tripteroides subobscurus (N. mirabilis; Australia, New Guinea; nepenthebiont[2])[16]
- Tripteroides tenax (N. albomarginata, N. ampullaria, N. gracilis, N. gymnamphora, N. mirabilis, N. rafflesiana, N. reinwardtiana, N. sanguinea, N. tobaica; Borneo, Java, Malaya, Sumatra, Thailand; nepenthebiont;[2] filter feeder[61])[63]
- Tripteroides vicinus (N. gracilis, N. lowii, N. mirabilis, N. sanguinea; Borneo, Malaya, Sumatra; nepenthebiont[2])[81][70][16]
- Tripteroides werneri (N. sp.; Philippines; nepenthebiont)[2]
- Tripteroides sp. 1 (N. sp.; Borneo; ecological group unknown)[2][88]
- Tripteroides sp. 2 (N. sp.; Borneo; ecological group unknown)[2][88]
- Tripteroides sp. No. 2 (N. rajah)[77]
- Tripteroides spp. (N. ampullaria, N. eustachya [N. alata][62])[63]
- Tripteroides spp. (N. gracilis; Singapore, Malaya; filter feeder)[66]
- Tripteroides spp. (N. stenophylla; Sarawak, Borneo; found in association with Topomyia nepenthicola)[79]
- ?Tripteroides sp. (N. sp., Philippines; elongate-ovoid eggs are possible adaptation to Nepenthes pitchers)[87][86]
- Uranotaenia
- Uranotaenia ascidiicola (N. gymnamphora; Java; nepenthebiont)[2]
- Uranotaenia belkini (N. madagascariensis; Madagascar; nepenthebiont[2])[89][35]
- Uranotaenia bosseri (N. madagascariensis; Madagascar; nepenthebiont[2])[89][35]
- Uranotaenia brevirostris[76] (N. ampullaria,[70] N. gracilis[67])
- Uranotaenia brunhesi (N. madagascariensis; Madagascar; nepenthebiont[2])[89][35]
- Uranotaenia damasii (N. madagascariensis; Madagascar; nepenthebiont?[2])[89][35]
- Uranotaenia gigantea (N. eustachya [N. alata],[62] N. tobaica; Sumatra; nepenthebiont[2])
- Uranotaenia moultoni (N. alata, N. ampullaria, N. bicalcarata, N. gracilis, N. rafflesiana, N. rajah;[77] Borneo, Malaya; nepenthebiont;[2] filter feeder[61])[63][50][51]
- Uranotaenia nepenthes (N. pervillei; Seychelles; nepenthebiont[2])[16]
- Uranotaenia nivipleura (N. distillatoria; Malaya, Sri Lanka; nepenthebiont?[2])[16]
- Uranotaenia xanthomelaena (N. gracilis; Malaya; nepenthebiont[2])[70]
- Uranotaenia sp. (N. ampullaria)[63]
- Uranotaenia spp. (N. stenophylla; Sarawak, Borneo; found in association with Topomyia nepenthicola)[79]
- Undetermined Culicidae (not Toxorhynchites) (N. albomarginata, N. ampullaria, ?N. eustachya [N. alata], N. gracilis, N. mirabilis, N. reinwardtiana, N. spathulata, "N. sp. A", "N. sp. B"; West Sumatra, Sumatra; filter feeders)[68]
- Undetermined Culicidae (N. attenboroughii, N. deaniana, N. gantungensis, N. leonardoi, N. mantalingajanensis, N. mira, N. palawanensis, N. philippinensis, N. sp. Anipahan; Palawan, Philippines; recorded from all known Palaweño Nepenthes;[90] very common and present in large numbers in at least N. attenboroughii[90] and N. leonardoi[91])[71][92][93]
- Dolichopodidae (long-legged flies)
- Systenus
- Undetermined Dolichopodidae (N. ampullaria)[34]
- Keroplatidae / Mycetophilidae (fungus gnats)
- Xenoplatyura
- Xenoplatyura beaveri[94] (N. ampullaria; Malaya; nepenthebiont;[2] terrestrial predator; larvae build webs above fluid level onto which they secrete highly acidic droplets (pH 1–2); catch arthropods leaving fluid as well as prey that falls in from above; one larva per pitcher; webs destroyed by larvae of Nepenthomyia malayana and therefore rarely found in the same pitchers as that species[59])[60][95][58][65]
- Lauxaniidae
- Undetermined Lauxaniidae (N. sp.; Java; ecological group unknown; family identification doubtful[2])[96]
- Muscidae (stable flies, house flies, face flies)
- Phaonia
- Phaonia nepenthincola (N. gymnamphora; Java; nepenthebiont?[2])[97]
- Phaonia sp. (N. eustachya [N. alata])[62]
- Phoridae (scuttle flies)
- Endonepenthia — see Megaselia[a]
- Megaselia[a]
- Megaselia bivesicata (N. ampullaria; Borneo, Java, ?Malaya; nepenthexene)[2]
- Megaselia cambodiae [≡Endonepenthia cambodiae][98] (N. sp.; Cambodia; nepenthebiont?[2])[34]
- Megaselia campylonympha [≡Endonepenthia campylonympha][98] (N. ampullaria,[34] N. mirabilis; Borneo, Sumatra; nepenthebiont[2])
- Megaselia corkerae (N. mirabilis)[16]
- Megaselia decipiens (N. gymnamphora; Java; nepenthebiont)[2]
- Megaselia deningsi (N. distillatoria; Sri Lanka; ecological group unknown[2])[16]
- Megaselia gregalis [≡Endonepenthia gregalis][98] (N. distillatoria, N. gymnamphora; Java, Sri Lanka; nepenthebiont; parasitised by undetermined ?Proctotrupidae[2][34])
- Megaselia meningi (N. distillatoria)[2][16]
- Megaselia nepenthina (N. albomarginata, N. gracilis, N. mirabilis,[2] N. sp.;[34] Borneo, ?Malaya, Sumatra; nepenthebiont[2])
- Megaselia schuitemakeri [≡Endonepenthia schuitemakeri][98] (N. albomarginata,[2][16] N. gracilis,[2][16][65][66] N. mirabilis,[2] N. rafflesiana;[65] Borneo, Malaya, Sumatra; nepenthebiont; carrion feeder; parasitised by undetermined genus nr. Tachinaephagus[2])[99][95][59]
- Megaselia tobaica [≡Endonepenthia tobaica][98] (N. tobaica; Sumatra; nepenthebiont; parasitised by undetermined Encyrtidae[2])[34]
- Megaselia sp. (N. tobaica; Sumatra; ecological group unknown)[3][2]
- Megaselia sp. n. (N. mirabilis; Hong Kong; ecological group unknown)[2]
- Megaselia spp. [=Endonepenthia spp.][98] (N. ampullaria, N. eustachya [N. alata],[62] N. spp.)[16][63]
- Undetermined Phoridae (N. gracilis, N. spathulata, "N. sp. A"; West Sumatra, Sumatra; pitcher visitors; "stayed inside the upper part of pitchers probably to mate and/or to oviposit")[68]
- Sarcophagidae (flesh flies)
- Pierretia — see Sarcophaga
- Sarcophaga
- Sarcophaga carolinensis [≡Sarcosolomonia carolinensis][34] (N. sp.)[34]
- Sarcophaga papuensis [≡Sarcosolomonia papuensis][100] (N. mirabilis in Australia; larvae have "prominent creeping welts" on body that facilitate movement in pitcher fluid; saprophagous; apparently leave pitcher to pupate[100])[34]
- Sarcophaga urceola [≡Pierretia urceola][101][2] (N. albomarginata, N. gracilis; Malaya; nepenthebiont[2])[16]
- Sarcophaga sp. (N. sanguinea; Malaya; ecological group unknown)[102][2]
- Sarcophaga sp. [=Pierretia sp.] (N. rafflesiana; Singapore)[61]
- Sarcosolomonia — see Sarcophaga
- Undetermined Sarcophagidae (N. maxima)[34]
- Sciaridae (dark-winged fungus gnats)
- Pnyxiopalpus
- Pnyxiopalpus nepenthophilus (N. sp.?; Malay Peninsula, Malaya; type description does not mention relationship with Nepenthes)[103]
- Syrphidae (hoverflies, flower flies)
- Nepenthosyrphus
- Nepenthosyrphus capitatus (N. reinwardtiana, N. tobaica; ?Malaya, Sumatra; nepenthebiont)[2]
- Nepenthosyrphus malayanus (N. sp.; Malaya; nepenthebiont)[2]
- Nepenthosyrphus oudemansi (N. ampullaria, N. rafflesiana; Borneo; nepenthebiont)[2]
- Nepenthosyrphus venustus (?N. sp.; Philippines; nepenthebiont?)[2]
- Nepenthosyrphus spp. (N. ampullaria, N. bicalcarata; Singapore[61])[63][50][51]
- Xylota
- Xylota sp. (N. ampullaria; Malaya; nepenthexene)[60][2]
- Undetermined genus nr. Pipizella (N. sanguinea; Malaya; ecological group unknown)[102][2]
- Tachinidae (tachina flies)
- Succingulum
- Succingulum fransseni (N. mirabilis; Borneo; ecological group unknown)[2]
- Undetermined Anoetidae (N. albomarginata, N. ampullaria)[2][63]
- Undetermined Harpacticoidae (N. ampullaria)[63]
- Undetermined Muscomorphae (N. eustachya [N. alata])[62]
- Undetermined Sciarida (N. ampullaria, N. eustachya [N. alata][62])[63]
Some species of mosquito[70][104] are only known from their pitcher plant hosts and have been named after them.
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]- Diapriidae
- Trichopria
- Trichopria sp. (N. ampullaria; Malaya; parasitoid wasps; Megaselia sp. recorded as host)[2]
- Elachertidae
- Undetermined Elachertidae (N. tobaica; Sumatra; parasitoid wasps; Phyllocnistis nepenthae recorded as host)[2]
- Encyrtidae
- Undetermined genus n. nr. Tachinaephagus (N. albomarginata, N. gracilis; Malaya; parasitoid wasps; Megaselia schuitemakeri [Endonepenthia schuitemakeri] recorded as host[2])[95][59]
- Undetermined Encyrtidae (N. tobaica; Sumatra; parasitoid wasps; Megaselia tobaica [Endonepenthia tobaica] recorded as host)[2]
- Formicidae (ants)
- Camponotus
- Camponotus schmitzi [identified as "Camponotus sp." in some early literature] (predominantly N. bicalcarata, but also N. gracilis and N. mirabilis; Borneo;[2] nepenthebiont[59] [not nepenthexene as reported by Beaver];[2] terrestrial predator and aquatic forager; colonises chamber in swollen tendril of N. bicalcarata; often remains hidden under inner peristome fold, but may attack newly caught insects and thereby prevent prey escape;[109][110][111] able to swim in pitcher fluid using tripod-like leg coordination,[112][113] remaining submerged for up to 30 seconds; targets large prey items only, cooperatively retrieving them from the fluid; prey remains and ant egesta drop back into pitcher, making nutrients more readily available to the plant;[59] plants colonised by C. schmitzi have more nitrogen available to them, and a higher proportion of it is insect-derived;[114] ant remains and egesta account for an estimated 42% of total foliar nitrogen on average (76% in plants with ant occupancy rates >75%);[110] ants increase nutrient retention by predating kleptoparasitic dipteran infauna, this nutrients later becoming available to the plant through the ants' waste;[114] breakup of prey keeps ammonium levels low, preventing putrefaction and therefore infaunal demise and premature pitcher death;[59] ant-inhabited plants have more leaves and a greater total leaf area; plants not inhabited by C. schmitzi do not appear to benefit significantly from carnivory; ant presence is associated with lower pitcher abortion rates and more voluminous pitchers (and consequently greater prey biomass);[110] C. schmitzi cleans peristome of fungal hyphae and other contaminants, maintaining high trapping efficiency over pitcher lifespan;[115][116] protects plant from pitcher-destroying weevils of the genus Alcidodes;[117][118][110] deutonymphs of the mite Naiadacarus nepenthicola are dispersed through phoresy on C. schmitzi;[119] C. schmitzi is an obligate mutualist, while N. bicalcarata is a facultative mutualist[59])[120][121][122][123][124][125][126][127][128][64][129][15][130][131][132][133][51][134][108][48][135][136][137][b]
- Camponotus sp. (N. bicalcarata; Borneo; nepenthexene; listed by Beaver as a previously unpublished record separate from Camponotus schmitzi)[2]
- Crematogaster
- Crematogaster sp. "small black" (N. bicalcarata, others; feed on nectaries on pitcher, pitcher lid, and laminar midrib;[28] undetermined Crematogaster sp. found as prey in N. bicalcarata[110])
- Dolichoderus
- Dolichoderus bituberculatus (N. gracilis, N. mirabilis; Borneo; nepenthexene)[2]
- Oecophylla
- Oecophylla smaragdina (N. ampullaria, N. rafflesiana; observed cooperatively retrieving prey from N. ampullaria pitcher and catching live mosquito larvae from N. rafflesiana pitcher)[28]
- Polyrhachis
- Polyrhachis pruinosa [originally identified as "Polyrachis" sp.[45] and later as "Polyrachis pruinosa"[28]] (N. gracilis, N. mirabilis, N. rafflesiana,[28] N. stenophylla?;[138] feeds on peristome nectar; apparently able to escape from pitcher fluid,[139] but also found as prey in N. bicalcarata[110] where it has been observed being attacked by Camponotus schmitzi;[111] 'drums' on pitcher lid as territorial threat display[45][28])
- Polyrhachis sp. (N. bicalcarata; Borneo; observed visiting nectaries on pitcher thorns)[59]
- Undetermined "small black" Dolichoderinae [originally identified as Crematogaster sp.][45] (N. gracilis and N. rafflesiana, "small, thick, web-like nests" attached to upper portion of live pitchers)[45][28]
- Undetermined ant (N. sibuyanensis; Philippines; nests in lower pitchers, also visiting upper pitchers; "offers little benefit to the plant")[59]
- Ichneumonidae (ichneumon wasps, scorpion wasps)
- Allocata
- ?Allocata sp. n. (N. tobaica; Sumatra; parasitoid wasps; Thomisus nepenthiphilus egg cocoon recorded as host[2])[3][59]
- Proctotrupidae
- Undetermined Proctotrupidae (N. gymnamphora; Java; parasitoid wasps; Megaselia gregalis [Endonepenthia gregalis] recorded as host; "[f]amily identification doubtful"[2])[96]
Mantises
[edit]- Empusidae
- Idolomorpha
- Idolomorpha madagascariensis (N. madagascariensis; Sainte-Luce, Madagascar; observed motionless on plant, but not near pitchers or flowers; member of same family found as prey of N. madagascariensis on east coast; rare)[140]
- Mantidae
- Undetermined Mantidae (N. madagascariensis; Madagascar; found on both pitchers and inflorescences; observed to move between plants; eggs found attached to plants; not particularly common)[140]
- Undetermined ?Mantidae (N. rafflesiana; Bako National Park, Borneo; observed perched under lid of upper pitcher, placing head inside trap)[141]
Moths
[edit]- Gracillariidae
- Phyllocnistis
- Phyllocnistis nepenthae (N. tobaica; ?Malaya, Sumatra; nepenthebiont?;[2] larvae mine pitcher wall parenchyma;[59] parasitised by undetermined Elachertidae[2])[142][3]
- Undetermined Gracillariidae (?N. eustachya [N. alata], N. inermis [N. bongso], "N. sp. B"; West Sumatra, Sumatra; larvae mine pitcher wall parenchyma; also found as leaf miners in laminae[68])[59]
- Noctuidae (owlet moths)
- Eublemma
- Eublemma radda (N. albomarginata,[45] N. bicalcarata, N. mirabilis, N. rafflesiana; Borneo, Malaya, Sumatra, Thailand; nepenthebiont;[2] larvae pupate in upper portion of N. albomarginata and N. rafflesiana pitchers in Peninsular Malaysia, Sarawak, and Singapore;[45] feeds on inner pitcher walls prior to pupation, causing conspicuous damage to interior surface[59])[143][130]
- Psychidae (bagworm moths)
- Nepenthophilus
- Nepenthophilus tigrinus (N. distillatoria; Sri Lanka; ecological group unknown; "[p]robably not a genuine inhabitant";[2][144] do not harm pitchers[59])[34][145]
Springtails
[edit]- Poduridae
- Podura
- Podura aquatica (N. gymnamphora [=N. melamphora])[146]
Arachnids
[edit]Mites
[edit]- Acaridae
- Naiadacarus
- Naiadacarus nepenthicola (N. bicalcarata; Brunei, Borneo; deutonymphs dispersed through phoresy on Camponotus schmitzi)[119]
- Histiostomatidae
- Anoetus — see Zwickia
- Creutzeria
- Creutzeria seychellensis (N. pervillei; Seychelles; nepenthebiont[2])[147][16]
- Creutzeria nr. seychellensis (N. pervillei ["N. perviellei"]; Seychelles)[148]
- Creutzeria tobaica[149] (N. madagascariensis, N. mirabilis, "N. tobaica";[2][16] Java, New Guinea, Madagascar; nepenthebiont; "[r]ecords probably include misidentifications";[2] originally described from a Javanese Nepenthes identified as N. tobaica,[149][150] although N. tobaica has not been recorded outside of Sumatra;[151] Juniper et al. (1989) wrote that N. reinwardtiana had been misidentified as N. tobaica,[16] but this species is also absent from Java[151])
- Creutzeria n. sp. (N. mirabilis; Australia)[148]
- Creutzeria n. sp. (N. madagascariensis; Madagascar)[148]
- Creutzeria sp. (N. madagascariensis)[152][35]
- Creutzeria spp. (N. albomarginata (Brunei), N. ampullaria (Brunei), N. bicalcarata (Brunei), N. gracilis (Brunei, Singapore, southern Thailand), N. mirabilis (Brunei))[148]
- Nepenthacarus
- Zwickia
- Zwickia guentheri[154] [≡Anoetus guentheri][154][150][16] (N. albomarginata, N. ampullaria, N. distillatoria, N. gracilis, N. gymnamphora, N. mirabilis; Java, Malaya, Sri Lanka, Sumatra; nepenthebiont; "[r]ecords probably include misidentifications"[2])[65]
- Zwickia nepenthesiana[155] [≡Anoetus nepenthesiana][2] (N. albomarginata, N. ampullaria, N. gracilis, N. gymnamphora, N. mirabilis, N. tobaica; Borneo, Java, Malaya, Sumatra; nepenthebiont; "[r]ecords probably include misidentifications"[2])[155][65]
- Zwickia sp. ["Zwickea sp."][84] (N. campanulata; Gunung Mulu National Park, Borneo; forms simple food web with Toxorhynchites sp. and Tripteroides nepenthis; 2 of 17 examined pitchers were colonised solely by this mite)[84]
- Undetermined Histiostomatidae [=Anoetidae] (N. eustachya [N. alata])[62]
- Undetermined mites (N. ampullaria; Singapore; several species(?) found)[156]
Some mite species inhabit a number of different Nepenthes species, while others are restricted to only one.[160]
Spiders
[edit]- Araneidae (orb-weaver spiders)
- Undetermined Araneidae (N. madagascariensis; Sainte-Luce, Madagascar; observed only once)[140]
- Lycosidae (wolf spiders)
- Undetermined Lycosidae (N. madagascariensis; Sainte-Luce, Madagascar; observed only once)[140]
- 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)
- Leptopholcus sp. (N. madagascariensis; Sainte-Luce, Madagascar; observed only once)[140]
- Salticidae (jumping spiders)
- Myrmarachne
- Myrmarachne sp. (N. madagascariensis; Madagascar; collected among Tetraponera ants, which it appears to mimic)[140]
- Salticus
- ?Salticus sp. [striped black and white species] (N. madagascariensis; Madagascar; second most common spider associate of N. madagascariensis after Synema; found on all plant parts including pitcher exterior; move between different pitchers and plants; flee pitcher when disturbed)[140]
- Thyene
- Thyene sp. ["Thyena sp."][35] (N. madagascariensis; Madagascar)[35]
- Undetermined Salticidae (2 species) (N. madagascariensis; Madagascar; found on all plant parts including pitcher exterior; move between different pitchers and plants; flee pitcher when disturbed; different species to ?Salticus sp.)[140]
- Tetragnathidae (long-jawed orb weavers, long jawed spiders)
- Undetermined Tetragnathidae (N. madagascariensis; Sainte-Luce, Madagascar; observed only once)[140]
- Theridiidae (tangle-web spiders, cobweb spiders, comb-footed spiders)
- Theridion
- Theridion decaryi (N. madagascariensis; Madagascar; nepenthephile?[2])[35][161]
- Theridion sp. n. (?N. stenophylla; Borneo; ecological group unknown)[2]
- Undetermined Theridiidae [white to colourless species 3–5 mm long] (N. madagascariensis; Madagascar; far less common associates of N. madagascariensis than Salticidae and Thomisidae; partially block pitcher opening with web, which stretches from lid to peristome; usually flee to peristome edge upon disturbance, but some do not exhibit fleeing behaviour; often observed carrying round egg sacs)[140]
- Thomisidae (crab spiders)
- Misumenops
- Misumenops nepenthicola[162] (N. albomarginata, N. gracilis, N. rafflesiana, N. reinwardtiana, N. tobaica; Borneo, Malaya, Sumatra; nepenthebiont;[2] lowland Nepenthes in Malaysia, Indonesia, Brunei, and Singapore; terrestrial predator and aquatic forager; drops into pitcher fluid when disturbed; can remain submerged for several minutes thanks to air bubble held in pit on ventral surface of abdomen; only present in larger pitchers of N. gracilis[66])[163][161][45][164][66]
- Misumenops thienemanni[165] (N. tobaica; Sumatra; nepenthebiont)[2]
- Synema
- Synema sp. (N. madagascariensis; Madagascar; )[140]
- Thomisus
- Thomisus callidus (N. tobaica; Sumatra; nepenthephile)[2]
- Thomisus nepenthiphilus (N. gracilis,[66] N. mirabilis, N. rafflesiana, N. reinwardtiana, N. tobaica,[2] N. sp.;[65] Sumatra; nepenthephile?; egg cocoon parasitised by ?Allocata sp.;[2] apparently incapable of aquatic foraging[66])[166]
- Thomisus sp. (N. eustachya; West Sumatra, Sumatra; observed waiting for prey just below the peristome on the outer surface of a pitcher)[59][167]
- Undetermined Thomisidae (?N. eustachya [N. alata], N. gracilis, N. spathulata, "N. sp. A", "N. sp. B"; West Sumatra, Sumatra; terrestrial predator)[68]
- Undetermined Thomisidae (N. mirabilis; southern Cambodia; drops into pitcher fluid when disturbed; appeared to be common; one per pitcher)[168]
- Undetermined Thomisidae (N. paniculata; New Guinea; observed on peristome of upper pitcher)[169]
- Undetermined "yellow crab spiders" (N. philippinensis; Mount Victoria, Philippines; drop into pitcher fluid when disturbed)[90][170]
- 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.
- Pollard, S.D. 2012. Fishing spiders in the hanging stomachs of Borneo. Carnivorous Plant Newsletter 41(1): 20–23.
Crustaceans
[edit]Branchiopods
[edit]- Incertae familiae
- Undetermined Cladocera sp. (N. ampullaria; Singapore)[156]
Copepods
[edit]- Canthocamptidae
- Parastenocarididae
- Phyllognathopodidae
- Incertae familiae
- Undetermined Harpacticoida (N. ampullaria, detritus feeders)[63]
- Undetermined copepods (N. ampullaria; Singapore; two species found)[156]
Decapods
[edit]- Sesarmidae
- Incertae familiae
- Undetermined crabs (N. attenboroughii; Mount Sagpaw, Philippines; at least two individuals found as prey in pitchers, including one female surrounded by possible eggs; may use pitchers to breed)[90]
- Undetermined Decapoda [Dutch: Garneeltjes] (N. cf. mirabilis)[177][178][179][59]
Molluscs
[edit]- Incertae familiae
- Undetermined snails (N. lowii; Borneo; occasional visitors; not known to contribute to pitcher contents)[29][c]
- Undetermined snail eggs (N. edwardsiana, N. villosa; Borneo; eggs laid on waxy inner surface of fresh pitchers)[45]
Annelids
[edit]- Incertae familiae
- Undetermined Oligochaeta (N. sp.)[34]
Nematodes
[edit]- Anguinidae
- Subanguina
- Subanguina spp. (N. gracilis,[66] N. spp.; detritus feeder)[65]
- Diplogastridae
- Diplogaster
- Diplogaster sp. (N. gymnamphora, accidental)[180]
- Dorylaimidae
- Dorylaimus
- Dorylaimus sp. (N. gymnamphora, accidental;[180] N. spp.[65])
- Panagrolaimidae
- Baujardia
- Baujardia mirabilis (N. mirabilis in Thailand, nepenthebiont?)[181]
- Panagrellus
- Panagrellus nepenthicola (N. gymnamphora, infaunal)[180][182]
- Plectidae
- Plectus
- Plectus sp. (N. gymnamphora, accidental)[180]
- Rhabditidae
- Rhabditis
- Rhabditis sp. (N. gymnamphora, accidental)[180]
- Incertae familiae
- Undetermined nematodes (N. eustachya [N. alata])[62]
Rotifers
[edit]- Incertae familiae
- Undetermined rotifers (N. ampullaria; Singapore; several species found)[156]
Protozoans
[edit]Amoebozoans
[edit]- Amoebidae
- Amoeba
- Amoeba guttula (N. gymnamphora [=N. melamphora])[146][156]
- Amoeba nepenthesi (N. gymnamphora [=N. melamphora])[146][156]
- Amoeba verrucosa (N. ampullaria; Singapore; single specimen found)[156]
- Arcellidae
- Arcella
- Arcella hemispherica (N. ampullaria; Singapore)[156]
- Arcella singaporensis[156] (N. ampullaria; Singapore; numerous specimens found)[156]
- Arcella subconica[156] (N. ampullaria; Singapore; several specimens found)[156]
- Arcella subhemispherica[156] (N. ampullaria; Singapore)[156]
- Arcella triangularis[156] (N. ampullaria; Singapore)[156]
- Arcella vulgaris (N. gymnamphora [=N. melamphora])[146][156]
- Centropyxidae
- Centropyxis
- Centropyxis aculeata (N. gymnamphora [=N. melamphora])[146][156]
- Cochliopodiidae
- Cochliopodium
- Cochliopodium bilimbosum (N. gymnamphora [=N. melamphora])[146][156]
- Difflugiidae
- Difflugia
- Difflugia constricta (N. gymnamphora [=N. melamphora])[146][156]
- Difflugia cf. pyriformis (N. ampullaria; Singapore; may represent new species)[156]
- Difflugia singaporensis[156] (N. ampullaria; Singapore)[156]
- Difflugia subpyriformis[156] (N. ampullaria; Singapore)[156]
- Difflugia suburceolata[156] (N. ampullaria; Singapore)[156]
- Lesquereusiidae
- Lesquereusia
- Lesquereusia epistomium (N. gymnamphora [=N. melamphora])[146][156]
- Microcoryciidae
- Diplochlamys
- Diplochlamys sp. (N. ampullaria; Singapore; single specimen found)[156]
- Incertae familiae
- Pyridicula
- Pyridicula klossi[156] (N. ampullaria; Singapore)[156]
Ciliates
[edit]- Incertae familiae
- Undetermined Vorticella-like protozoans have been recorded from the body surfaces of some infaunal mosquito larvae.[77]
Fungi
[edit]- Undetermined mycelial fungus (N. mirabilis; along Jardine River, Australia; observerd as both free-living in the trap's fluid and attached to chitinous insect remains)[183][184]
- Undetermined yeasts (N. madagascariensis; Madagascar)[35]
Higher fungi
[edit]- Entylomataceae
- Tilletiopsis
- Tilletiopsis sp. (West Malaysia)[185][186]
- Sporidiobolaceae
- Rhodotorula
- Rhodotorula rubra (West Malaysia)[185]
- Sporobolomyces
- Sporobolomyces roseus (West Malaysia)[185]
- Tremellaceae
- Bullera
- Bullera alba (West Malaysia)[185]
- Cryptococcus
- Cryptococcus albidus (West Malaysia)[185][186]
- Cryptococcus laurentii (West Malaysia)[185]
- Trichosporonaceae
- Trichosporon
- Trichosporon pullulans (West Malaysia)[185]
Pin molds
[edit]- Mucoraceae
Sac fungi
[edit]- Dothioraceae
- Aureobasidium
- Aureobasidium pullulans (West Malaysia)[185]
- Saccharomycetaceae
- Candida
- Candida diffluens (West Malaysia)[185]
- Trichocomaceae
Algae
[edit]Diatoms
[edit]- Achnanthaceae
- Achnanthes
- Achnanthes lanceolata (N. gymnamphora [=N. melamphora])[146]
- Achnanthes minutissima (N. gymnamphora [=N. melamphora])[146]
- Cocconeidaceae
- Cocconeis
- Cocconeis placentula var. lineata (N. gymnamphora [=N. melamphora])[146]
- Naviculaceae
- Navicula
- Navicula elliptica (N. gymnamphora [=N. melamphora])[146]
- Navicula viridis (N. gymnamphora [=N. melamphora])[146]
- Rhopalodiaceae
- Epithemia
- Epithemia sorex (N. gymnamphora [=N. melamphora])[146]
Golden algae
[edit]- Incertae familiae
- "minute simple flagellates of monad-type" (N. ampullaria; Singapore; identification based on "small bodies with indications of flagella")[156]
Green algae
[edit]- Desmidiaceae
- Euastrum
- Euastrum sp. (N. gymnamphora [=N. melamphora])[146]
Bacteria
[edit]Cyanobacteria
[edit]- Merismopediaceae
- Merismopedia
- Merismopedia glauca ["Merismopedium glaucum"] (N. gymnamphora [=N. melamphora])[146]
Firmicutes
[edit]- Bacillaceae
- Bacillus
- Bacillus cereus (N. sp[p].; secretes amylases, lipases, and proteases; may play role in digestion)[188]
- 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
- Yiung, C.L. 2012. A novel analysis of bioactive compounds in pitcher fluids of Nepenthes pitcher plants. Ph.D. thesis, Monash University Malaysia, Bandar Sunway.
- 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]
- (in Indonesian) Yogiara 2004. "Analisis komunitas bakteri cairan kantung semar (Nepenthes spp.) menggunakan teknik terminal restriction fragment length polymorphism (T-RFLP) dan amplified ribosomul DNA restriction analysis (ARDRA)" (PDF). M.Sc. thesis, Bogor Agricultural University, Bogor.
- Yogiara, A. Suwanto & M.T. Suhartono 2006. A complex bacterial community living in pitcher plant fluid. Jurnal Mikrobiologi Indonesia 11(1): 9–14.
- Siegara, A. & Yogiara 2009. Bacterial community profiles in the fluid of four pitcher plant species (Nepenthes spp.) grown in a nursery. Microbiology Indonesia 3(3): 109–114.
- Takeuchi, Y., M.M. Salcher, M. Ushio, R. Shimizu-Inatsugi, M.J. Kobayashi, B. Diway, C. von Mering, J. Pernthaler & K.K. Shimizu 2011. In situ enzyme activity in the dissolved and particulate fraction of the fluid from four pitcher plant species of the genus Nepenthes. PLoS ONE 6(9): e25144. doi:10.1371/journal.pone.0025144
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
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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]]