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Nephilengys malabarensis (Walckenaer, 1842)
The malabar spider. Eunuch phenomenon.

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Nephilengys malabarensis (Photo by Marcus Ng (C))

Biology

Web

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Typical web construction of an orb-web spider (Picture taken from Spiders, The Ultimate Predators)

N. malabarensis construct large aerial orb webs that can measure up to a metre high depending on the size of the spider. The webs have a tubular retreat connected to the hub so as to allow the spider to escape when faced with threats. Unlike Nephila, Nephilengys do not have golden webs and are always built against substrates such as tree trunks, rock outcrops and mainly against house walls and roofs. Immature spiders tend to have symmetrical webs while mature females have large webs that are disproportionately closer to the top frame. Unlike most of its orb web relatives that repair the entire web, N. malabarensis only repairs the damaged parts of the webs [1] .

Feeding habits

N._malabarensis_feeding.png
Female Nephilengys malabarensis with cicada and waiting Argyrodens flavescens (Photo by Marcus Ng (C))


N. malabarensis is observed to feed on small insects in the wild. In laboratory, they grow well with a diet of fruit flies (Drosophila melanogaster), house flies (Musca domestica), mealworm (Tenebrio molitor), superworm (Zophobus morio).

Generally, it is observed that the spider rushes to the prey upon it being trapped in the web. The spider then delivers a bite, holds the prey to immobilize it or break up the prey with its powerful jaws. In some instances, the spider will wrap the prey up using the silk from their spinnerets then start feeding or, just consume the prey without wrapping [2] [3]

 .

External digestion

Powerful enzymes are released into the prey via the spider's maxillary glands, digesting the prey and its internal organs gradually. The spider will then suck up the partially digested food source into its alimentary canal. This action of its jaws and digestion process results in the prey becoming a dry and indigestible mass. In the event that the prey has a hard exosleleton, the spider will pierce a hole and inject enzymes to digest the prey. After the digestion is complete, it will then suck up the content and discard the empty exoskeleton.

The internal process

The partially digested liquid will first pass through the spider's oesophagus into the sucking stomach. The walls of the sucking stomach is attached to strong muscles that allows strong sucction force to take in the liquidified food. There is also a filter system formed by a system of hairs along the mouth and throat to prevent the intake of any undigested food particles. Particles that are trapped in the system will be washed out of the mouth by the secretion of digestive enzymes from the maxillary glands. The liquid food will enter the intestines by the action of circular muscles surrounding the sucking stomach. The actual digestion is increased by a series of sacs or diverticula that branch off from the midgut. At the last part of the alimentary canal, the cloaca stores the waste food and this will be passed out of the osy as faeces. Nitrogenous waste in the body is produced by malphigian tubules (a kind of kidney). The malphigian tubules secrete a substance called guanates which will be passed into the cloaca to be passed out of the body together with the other waste materials.


Reproduction


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Mature female with male palp insertion (Photo by Lee Qi Qi (C))
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Male preparing to climb onto female's abdomen
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Male on female's abdomen
Mating_3.png
Successful copulation and palp insertion

N. malabarensis reproduces sexually with the insertion of the male palps into the complementary female's epigynum. Extreme sexual dimorphism is prevalent in

N. malabarensis where the females are much larger than the males. A mature female measures approximately 20-30mm while the males are approximately 3-5mm. The males are opportunistic and tend to mate with the females when they are occupied with their meals or are distracted to prevent being cannibalized by the females.

During mating, the male typically wave its legs and shake the female's web as it approach the female. Upon close proximity, the male will touch the female's legs and/or abdomen and climb onto the female when the opportunity arises. When the male is on the dorsum, it will pause and stroke the female and approach the epigynum laterally or via the abdomen apex. The male will then attempt to initiate copulation as it searches for the copulatory opening on the epigynum with its palps so as to insert the palp into the female's epigynum. Upon the insertion of the palp, the male will draw its body backwards to facilitate palp removal by self mutilation. In the case of the female, it will either be receptive, unreceptive or aggressive. A receptive female will move its legs and orient itself towards the male as it moved out of the retreat. In the event that the female is unreceptive, it will just remain motionless. Females may also display aggression by shaking its body and web and also chasing the males away. During copulation, the females may partake sexual cannibalism of her mating partner [4] .

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Egg sac of Nephilengys malabarensis (Photo by Lee Qi Qi (C))



Eunuch Phenomenon

One interesting point about these spiders is the epigynum plugging by males as seen in the screenshots above. In order to increase the paternity of the males, the males can undergo self-mutilation to result in entire palp loss so as to prolong the sperm transfer time to allow more sperms transfer to the females. This is hypothesised to ensure paternity and also to escape cannibalism. This is known as the eunuch phenomenon. Males with both palps are known as intact males; males with one palp are known as half-eunuchs and males without palps are known as eunuchs.

After plugging the female with its palp, the male will normally stay on the female web in an attempt to guard off other males from removing his palp and mating with the female. It is also hypothesised that males that has been emasculated tend to be more aggressive; eunuchs are more aggressive than half eunuchs and half eunuchs are more aggressive than intact males.

There are ongoing research about this phenomenon and more can be known from the paper "Eunuchs are Better Fighters"
[5] .

More images of intact males, half eunuchs and eunuchs

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Lateral view of a eunuch spider (Photo by Lee Qi Qi (C))


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Ventral view of a half-eunuch (Photo by Lee Qi Qi (C))


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Anterior view of intact male (Photo by Lee Qi Qi (C))
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Eunuch male spider with no palps. (Photo by Lee Qi Qi (C))


Description

General

Nephilengys_cephalothorax_-_dorsal.2.jpg

Eyes

  • Eight eyes
  • Position of the lateral eyes differentiates araneidae from nephilidae
  • Eyes are widely separated in Nephila and Nephilengys [6]

  • Almost equidistant from one eye to another
  • Eyes 2 and 3 are the main eyes which possess retina and lens
  • The rest of the six eyes are secondary eyes with lens, but the retina differs from the main eyes
  • N. malabarensis do not depend much on their eyes as they are not hunters


Chelicerae

  • Used for feeding
  • Large, black mouthpart
Nephilengys_male_-_palp.p.jpg

Palps (male)

  • Cockscrew, hook-like embolic conductor in adult males
  • Inserted into the female's epigynum during mating
  • Can be self-emasculated during mating so as to increase sperm transfer [7]

Epigynum (female)

  • Absent (undeveloped) in juvenile female
  • Will only be developed after approximately six times of moulting
  • Distinctively opened and black in colour
  • Males will insert palp and plug its palp into the epigynum opening during copulation (mating)

Book lungs

  • Respiratory organ of the spider
  • Resembles pages in a book
  • Consist of pile of hollow plates
  • Blood passing through the plates in the booklungs on the way back to the heart picks up oxygen which diffuses from the outiside into the hollo within the plates abd then through the plate wall
Nephilengys_spinnerets_1a_copy.jpgSpinnerets

  • Presence of cribellum that produces a special type of trapping silk called "hackled band"
  • Cribellum is formed from a modified pair of spinnerets
  • Multi-stranded spider silk produced from cribellum and combed with special hairs on the spider's hind legs
  • Only present in seven families, the cribellate


Anal tubercle

  • A small lobe anterior to the spinnerets bearing the anal opening
Nephilengys_claws.jpgClaws

  • Three claws
  • Typical in web-weaving spiders
  • For handling silk and locomotion on web
illustrated1.jpg


illustrated2.jpg


Male

FeatureJuvenileAdult

Juvenile_male_dorsal_view.pngAdult_male.jpg
PalpsNephilengys_male_sub_adult_-_both_palps_.jpg
Brown-black colour

Undeveloped

Short and swollen embolic conductor with distal bulge
Nephilengys_male_-_palp.p.jpg
Orange-black colour

Developed

With long and slender embolic conductor with distal bulge
Size1mm - 5mm3mm - 5mm
CephalothoraxDark brown black with medially white sternumDark reddish orange with orange sternum
AbdomenPatterned grey-black with 6 white spots at venter, brown scutumBlack and grey with orange scutum
VenterLight brown grey with two pairs of lighter spotsOrange
AppendagesAlternating yellow and black
Dense hair on appendages
Generally solid grey-black except coxae, trochanters and proxiaml femora that are yellow
Dense hair on appendages

More pictures of male spiders
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Dorsal view of adult male (Photo by Rayson Lim (C))
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Ventral view of adult male (Photo by Rayson Lim (C))
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Juvenile male (Photo by Lee Qi Qi (C))



Female

FeaturesJuvenileAdult

Juvenile_female_front.jpgPicture_5.png
EpigynumNot developed, closed, translucentDeveloped, open, black coloured Approximately as long as wide, with (inverted T-shaped) epigynal septum, medial copulatory openings and anterior rim
Inner epigynum with sclerotized arch and round spermathecae juxtaposed medially
Copulatory ducts are round and heavily sclerotized and the fertilization ducts are long
Epigynal shape may vary within populations
Size10mm to 30mm30mm - 50mm
CephalothoraxDark brown black
With medially white sternum
Sternum colour varies from yellow to bright orange
With broad lateral brown bands
AbdomenPatterned brown-black dorsal with yellow spots at venter
Yellow sternum
Patterned black and brown dorsal with bright orange patches on venter
Orange sternum
Width is variable depending on food intake
AppendagesAlternating yellow and blackSolid brown-black with yellow parts

Black: coxae, trochanters, distal femora, patellae, distal tibiae, metatarsi, tarsi
Yellow: proximal femora, tibiae
First femur with sparse and long prolateral spines


More photos of female N. malabarensis

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Ventral view of adult female (Photo by Rayson Lim (C))
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Lateral view of adult female (Photo by Rayson Lim (C))
Anterior_view.jpg
Juvenile female anterior view (Photo by Marcus Ng (C))

Original description

Epéire ANAMA. Long.
Corselet brun rougeâtre ; yeux antérieurs du carré intermédiaire plus rapprochés entre eux que les postérieurs ; yeux latéraux rapprochés au niveau de ceux d'enhaut. Sternum d'un jaune vif boidé de noir. Abdomen d'un brun paie, moucheté de jaune sur le dos, avec des raies arquées jaunes sur les côtés; qui se prolongent sous les côtés du ventre plus clairs et plus larges; deux croissants jaunes opposés sous le ventre entourant la vulve.Pattes rouges avec des anneaux bruns aux articulations , fines et pas très-allongées. (M.)
Ancien-Monde— Asie—Cochinchine. (Walckenaer, 1842)

Translated from Google translate:
Spider ANAMA. Long.
Thorax reddish-brown eyes past the square through closer together than the posterior, lateral eyes close to the level of the right side up. Sternum bright yellow black. Abdomen brown, speckled with yellow on the back, with arched yellow stripes on the sides, which extend under the belly side clearer and broader; two yellow crescent opposite the belly around the vulva. Legs red with brown rings in the joints, and thin not very elongated. (M.)
Old-World-Asia-Cochin.


Distribution


N. malabarensis is first discovered in South India on the Malabar coast and found all across South, South-East and East Asia: from India and Sri Lanka to the Philippines, north to China (Yunnan: Yin et al., 1990), north-east to Japan (Saga, Kompira: Bösenberg & Strand, 1906: 192), east to Ambon (Thorell, 1878: 123; 1881: 157).

This species is synanthropical; dwelling in human habitats in South and South-East Asia, and less common in rainforest (Murphy & Murphy, 2000). Some populations like the Niah (Borneo) population are also known to inhabit cave entrances (Deeleman-Reinhold, 1989).

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Distribution of Nephilengys malabarensis (Taken from Wikipedia)




Where is it found in singapore?

These are some of the sites that N. malabarensis can be found. These spiders are synantropic, you may be able to find them in any town parks or housing areas [8].



Taxonavigation

Juvenile_male.jpgKingdom: Animalia

Phylum: Arthropoda

Class: Arachnida

Order: Araneae

Suborder: Araneomorphae

Family: Nephilidae

Genus: Nephilengys

Species: Nephilengys malabarensis




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Simplified phylogenetic tree showing phylogenetic relationships of Nephilengys malabarensis



Belonging to the family Nephilidae, N. malabarensis has relatively large orb webs like the spiders from its spiderrs sister genus, Nephila. This simplified tree shows the relative phylogenetic relations between the species in the Nephilengys species and the other genus. The review of the species status of Nephilengys borbonica has also been noted and changed accordingly.  [9] [10] [11]

Etymology

The etymology of the genus name comes about from the proximity of relationship to the sister genus Nephila where Nephilengys = Nephila + Ancient greek -engy- which means "near to" or "close to". The spider is named after the place it was discovered, the Malabar coast in India.

Synonyms

SynonymAuthor, year, detailsMuseumDescription

Epeira anama [12]

Walckenaer 1842: 102-Description of female (from Anam or Cochinchina, today’s Vietnam); type(s) not found

Epeira malabarica [13]

Doleschall, 1857: 420--

Epeira rhodosternon [14]

Doleschall, 1859: 40, pl. 12, f. 6RMNHDescription of female (from Java)

Nephila rivulata [15]

O. P.-Cambridge, 1871: 618, pl. 49, f. 1–2OUMNHDescription of male and female (from Labuan, China Sea)

Nephilengys schmeltzii [16]

L. Koch, 1872: 144, pl. 11, f. 7,-Description of female (from the Philippines)

Nephilengys hofmanni [17]

L. Koch, 1872: 145, pl. 11, f. 8-Description of female (from Borneo)

**Nephilengys malabarensis [18] [19] [20] [21] [22] [23] [24]

Thorell, 1878: 123
Dahl, 1912: 49
Roewer, 1942: 933
Deeleman-Reinhold, 1989: 624, f. 11–14
Barrion & Litsinger, 1995: 565, f. 350a–c
Song et al., 1999: 217, f. 125F–I
Platnick, 2005
--

Nephila urna [25]

van Hasselt, 1882: 28, pl. 4, f. 12–14-Description of male (from Sumatra); type(s) not found

Nephilengys malabarensis var. g, annulipes [26]

Thorell, 1890: 188-Description of male and immature female (from Sumatra)

Nephila malabarensis [27] [28] [29] [30] [31] [32] [33] [34]

Simon, 1894: 745, f. 827
Pocock, 1900: 217, 219
Bosenberg & Strand, 1906: 192, pl. 11, f. 216
Bonnet, 1958: 3079
Wiehle, 1967: 195, f. 48
Tikader, 1982: 95, f. 183–186
Millidge, 1988: 258, f. 24
Yin et al., 1990: 3, 141, f. 6–9
--

Metepeira andamanensis [35]

Tikader, 1977: 181, f. 12A–C-Description of female

Nephilengys niahensis [36]

Deeleman-Reinhold, 1989: 626, f. 15–16MHNGDescription of female (from Borneo); syn.n. Female holotype, three female and an immature paratype from Niah Cave, Sarawak, East Malaysia

Abbreviation
MHNG: Muséum d’Histoire Naturelle de la Ville de Genève, Geneva, Switzerland
OUMNH: Oxford University Museum of Natural History, Oxford, U.K.Abbreviations
RMNH: Rijksmuseum van Natuurlijke Historie, Leiden, the Netherlands

Type Information


The holotype for Nephilengys malabarensis is stored in Muséum National d’Histoire Naturelle, Paris, France.

A holotype is a physical (or illustrated) specimen used in the naming and original description of a species. There is only one holotype and it is important to take note of the original description and holotype so that one can refer back to the original specimen should there be any confusion in identifying a specimen or to review the species, genus, or family status. In the event that a holotype is destroyed or lost, another specimen will be selected and this is known as a neotype. Click on this webpage for more information on the different types.


Diagnosis


As spiders are specific in their mating, successful mating can only occur when the palp (male) is complementary to the epigyne (female) like those of the "lock and key" system. Each species of spider have its unique palpal and epigynal structure, rendering these features important diagnostic tools.

N. nephilengys differs from N. cruentata and N. borbonica by the presence of a well-defined anterior rim and the epigynal septum. The epigynum septum separates two chambers with medially orientated copulatory openings, by the presence of a sclerotized arch in the inner epigynum and by the absence of the epigynal copulatory groove. They differ from N. papuana by the combination of the following: sternum orange, rarely with a thin dark brown lateral edge, venter with two conspicuous pairs of orange dots, first femur with sparse prolateral spines of normal length, epigynum with a narrow septum and both copulatory and fertilization ducts longer than the spermatheca radius.

N. malabarensis males has a long and slender embolic conductor and distal modification of the embolus in the form of a bulge, differentiating itself from N. cruentata and N. borbonica. N. malabarensis can be differentiated from N. papuana by the size where the prosoma length 2.5–3.1; total length 5.0–5.9) as well as the embolic conductor shape and sclerotization. The distal EC has a longer and spiralling tip causing it to appear hooked when viewed from and mesal sides. N. malabarensis' proximal embolic conductor is long, straight and sclerotized.

Illustrations of epigynum and palps can be found here.



Taxonomic history

  • Walckenaer (1842) described Epeira anama from Vietnam and E. malabarensis from India’s Malabar Coast.
  • Dahl (1912: 46) pointed out their synonymy and chose to keep the name Nephilengys malabarensis.
  • Walckenaer’s type series in MNHN is scattered and partly lost, examination the holotype of E. malabarensis but not the type(s) of E. anama was conducted by Kuntner (2007), but the material at hand, coupled with Walckenaer’s description, support the synonymy.
  • The first record of N. malabarensis in Java was by Doleschall (1857), who, perhaps unintentionally (cf. Dahl,1912: 46), used the incorrect name Epeira malabarica (Walck.). He later described another species from Java, Epeira rhodosternon Doleschall, 1859.
  • This was synonymized with N. malabarensis by Thorell (1881), and confirmed as such by Dahl (1912), also in this study. In 1871, O. P.-Cambridge described Nephila rivulata from Labuan, in the China Sea. Koch (1872: 143) and Dahl (1912: 47) cited the type locality of this species as Ceylon (Sri Lanka). The exact type locality(ies) of N. rivulata remain(s) dubious; see type label data, where Ceylon [Sri Lanka] and Amboina [Ambon] are cited as the type localities. ‘Labuan’ could be in the Philippines, Malaysia or Indonesia (hence, the broad geographical term China Sea). As O. P.-Cambridge noted (p. 619), his N. rivulata is ‘nearly allied to Epeira malabarensis’. In fact, it is synonymous, as pointed out by Dahl (1912: 47). O. P.-Cambridge’s description of both sexes is detailed enough and sufficiently illustrated to confirm that N. rivulata is a synonym of N. malabarensis. Kuntner (2007)'s examination of syntypes supports the synonymy.
  • However, O. P.-Cambridge has to be given credit for recognizing the importance of male genitalic characters in spider taxonomy. His vivid description of the male palpal ‘corkscrew-spine’ is in fact the first detailed description of the male (embolic) conductor of this species. Koch (1872: 143) described two new species, Nephilengys schmeltzii L. Koch, 1872 and Nephilengys hofmanni L. Koch, 1872, from the Philippines and Borneo, respectively.
  • Dahl (1912) synonymized both with N. malabarensis and declared that the epigynum figure of N. schmeltzii in Koch (1872: pl. 11, f. 7) depicts an immature animal. Koch’s illustrations of the epigyna are not detailed enough for a reliable identification, but both do show adult epigyna.
  • Kuntner (2007)'s examination of Nephilengys material from the Philippines and Borneo and the study of epigynal variation support the synonymy with N. malabarensis.
  • Thorell (1878: 123) recorded N. malabarensis from Amboina (Ambon, Maluku, Indonesia), possibly the species’ easternmost locality.
  • From Sumatra two new names were introduced. van Hasselt, (1882: 28) described a male of Nephila urna, and Thorell (1890: 188) described a new subspecies, N. malabarensis annulipes.
  • Dahl (1912: 49) synonymized both with N. malabarensis and redescribed the species (without illustrations).
  • In 1977, Tikader described the female of a new species, Metepeira andamanensis, from Andaman Islands, which he later (Tikader, 1982) correctly synonymized with Nephila malabarensis (1/4 Nephilengys m).
  • Deeleman-Reinhold (1989: 626) described a new species, Nephilengys niahensis, from Borneo. After Kunter (2007) examined the holotype and paratype females, as well as additional male and female material from Borneo, including a male from the type locality, he proposes that N. niahensis is a junior synonym of N. malabarensis.
  • Most authors (Chrysanthus, 1959, 1971; Davies, 1988) have treated all Australasian Nephilengys as incorrectly as N. malabarensis.
  • The populations from New Guinea and Australia can be diagnosed as a distinct species, N. papuana. Kuntner (2007) delimit the geographical distribution of N. malabarensis to include South and South-East Asia, but not New Guinea and Australia.


Literature and References


  1. ^ Kuntner, M., 2007. A monograph of Nephilengys, the pantropical ‘hermit spiders’ (Araneae, Nephilidae, Nephilinae). Systematic Entomology. 32: 95–135.

  2. ^ Kuntner, M., 2007. A monograph of Nephilengys, the pantropical ‘hermit spiders’ (Araneae, Nephilidae, Nephilinae). Systematic Entomology. 32: 95–135.

  3. ^ Preston-Mafham, R., 1991. Spiders, An Illustrated Guide. Blandford, Great Britain. 144pp.

  4. ^ Kuntner, M., 2007. A monograph of Nephilengys, the pantropical ‘hermit spiders’ (Araneae, Nephilidae, Nephilinae). Systematic Entomology. 32: 95–135.

  5. ^ Kralj-Fišer, S., M. Gregorič, S. Zhang, D. Li & M. Kuntner, 2011. Eunuchs are better fighters. Animal Behaviour.81: 933-939

  6. ^ Kuntner, M., J. A. Coddington & G. Hormiga, 2008. Phylogeny of extant nephilid orb-weaving spiders (Araneae, Nephilidae): testing morphological and ethological homologies. Cladistics. 24:147–217.

  7. ^ Oh, J., 2010. The Adaptive Significance of Entire Palp Removal in the Orb-Weaving Spider, Nephilengys malabarensis (Araneae: Nephilidae).

  8. ^ http://static.stomp.com.sg/stomp/sgseen/what_bugs_me/771374/creepy_spiders_at_bus_stop_make_commuters_skin_crawl.html?commentId=810630&action=stomp_club_report_comment_offensive&tokenKey=stomp_club_report_comment_offensive_810630&token=&article=contentbean:771374&video=

  9. ^ Kuntner, M. & I. Agnarsson, 2011. Biogeography and diversification of hermit spiders on Indian Ocean islands (Nephilidae: Nephilengys). Molecular Phylogenetics and Evolution. 59:477–488

  10. ^ Kuntner, M., S. Kralj-Fišer & M, Gregorič, 2010. Ladder webs in orb-web spiders: ontogenetic and evolutionary patterns in Nephilidae. Biological Journal of the Linnean Society. 99: 849–866.

  11. ^ Kuntner, M., J. A. Coddington & G. Hormiga, 2008. Phylogeny of extant nephilid orb-weaving spiders (Araneae, Nephilidae): testing morphological and ethological homologies. Cladistics. 24:147–217.

  12. ^ Walckenaer, C.A.,1842. Apte`res. Histoire Naturelle des Insects Vol.2//. Roret, Paris. Pp. 1–549.

  13. ^ Doleschall, C.L.,1857. Bijdrage tot de Kennis der Arachniden van den Indischen Archipel. Natuurkundig Tijdschrift voor Nederlandsch
    Indie. 13:399–434.

  14. ^ Doleschall, C.L., 1859. Tweede Bijdrage tot de Kennis der Arachniden van den Indischen Archipel. Verhandelingen der Natuurkundige
    Vereeniging in Nederlandsch Indie. 5:1–60.

  15. ^ Cambridge, O.P., 1871. Notes on some Arachnida collected by Cuthbert Collingwood, Esq., M.D., during rambles in the China Sea//. Proceedings of the Zoological Society of London. Pp. 617–622.

  16. ^ Koch, L., 1872. Die Arachniden Australiens//. Nürnberg.

  17. ^ Koch, L., 1872. Die Arachniden Australiens//. Nürnberg.

  18. ^ Thorell, T., 1878. Studi sui ragni Malesi e Papuani. II. Ragni di Amboina raccolti Prof. O. Beccari. Annali del Museo Civico di Storia Naturale di Genova. 13:1–317.

  19. ^ Dahl, F.,1912. Seidenspinne und Spinnenseide. Mitteilungen aus dem Zoologischen Museum in Berlin. 6:1–90.

  20. ^ Roewer, C.F., 1942. Katalog der Araneae von 1758 bis 1940, bzw 1954 Vol. 1. P. Budy, Bremen.

  21. ^ Deeleman-Reinhold, C.L.,1989. Spiders from Niah Cave, Sarawak, East Malaysia, collected by P. Strinati. Revue Suisse de Zoologie. 96:619–627.

  22. ^ Barrion, A.T. & Litsinger, J.A.,1995. Riceland Spiders of South and Southeast Asia. CAB, Wallingford, Oxfordshire.

  23. ^ Song, D., Zhu, M. & Chen, J.,1999. The Spiders of China. Hebei Science and Technology Publishing House, Shijiazhuang.

  24. ^ Platnick, N.I.,2011. The World Spider Catalog, Version 6.0. URL http://research.amnh.org/entomology/spiders/catalog/ [accessed 1 November 2011].

  25. ^ van Hasselt, A.W.M., 1882. Araneae//. Midden Sumatra (ed. by P.J. Veth), Leiden. Pp. 1–56.

  26. ^ Thorell, T., 1890. Studi sui ragni Malesi e Papuani. IV, 1. Annali del Museo Civico di Storia Naturale di Genova. 28:1–419.

  27. ^ Simon, E., 1894. Histoire Naturelle des Araigne´es Vol. 1, Paris. Pp. 489–760.

  28. ^ Pocock, R.I., 1900. Arachnida. The Fauna of British India, Including Ceylon and Burma. Taylor & Francis, London.

  29. ^ Bösenberg, W. & Strand, E.,1906. Japanische Spinnen. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft. 30: 93–422.

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This page was authored by Lee Qi Qi

Last curated in 2011

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