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'''''Nephila pilipes''''' ('''northern golden orb weaver''' or '''giant golden orb weaver'''<ref name=AU>[http://www.arachne.org.au/01_cms/details.asp?ID=1627 ''Nephila pilipes''], Arachne.org.au</ref>) is a species of [[golden orb-web spider]]. It can be found in [[Japan]], [[China]], [[Vietnam]], [[Cambodia]], [[Taiwan]], [[Malaysia]], [[Singapore]], [[Myanmar]], [[Indonesia]], [[Thailand]], [[Laos]], [[Philippines]], [[Sri Lanka]], [[India]], [[Nepal]], [[Papua New Guinea]], and [[Australia]]. It is commonly found in [[Primary forest|primary]] and [[Secondary forest|secondary forests]] and gardens. Females are large and grow to a body size of 30–50&nbsp;mm (overall size up to 20&nbsp;cm), with males growing to 5–6&nbsp;mm. It is the second largest of the orb-weaving spiders apart from the recently discovered ''[[Nephila komaci]]''. The first, second, and fourth pairs of legs of juvenile females have dense hairy brushes, but these brushes disappear as the spider matures.
'''''Nephila pilipes''''' ('''northern golden orb weaver''' or '''giant golden orb weaver'''<ref name=AU>[http://www.arachne.org.au/01_cms/details.asp?ID=1627 ''Nephila pilipes''], Arachne.org.au</ref>) is a species of [[golden orb-web spider]]. It resides all over countries in [[East Asia|East]] and [[Southeast Asia]] as well as [[Oceania]]. It is commonly found in [[Primary forest|primary]] and [[Secondary forest|secondary forests]] and gardens. Females are large and grow to a body size of 30–50&nbsp;mm (overall size up to 20&nbsp;cm), with males growing to 5–6&nbsp;mm. It is the second largest of the orb-weaving spiders apart from the recently discovered ''[[Nephila komaci]]''. The first, second, and fourth pairs of legs of juvenile females have dense hairy brushes, but these brushes disappear as the spider matures.


The ''Nephila pilipes'' golden web is vertical with a fine irregular mesh. It is not symmetrical, with the hub usually nearer the top.<ref name=AU/> Rather than [[egg sac]]s being hung in the web, a pit is dug which is then covered with plant debris or soil.
The ''N. pilipes'' golden web is vertical with a fine irregular mesh. It is not symmetrical, with the hub usually nearer the top.<ref name=AU/> Rather than [[egg sac]]s being hung in the web, a pit is dug which is then covered with plant debris or soil.




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==Description==
==Description==
''Nephila pilipes'' display female gigantism and male dwarfism (see [[#sexual dimorphism|Sexual dimorphism]] section). In terrestrial animals, ''Nephila pilipes'' have the most size differences between males and females.<ref name=":0">{{Cite journal|last=Kuntner|first=Matjaž|last2=Zhang|first2=Shichang|last3=Gregorič|first3=Matjaž|last4=Li|first4=Daiqin|date=November 2012|title=Nephila female gigantism attained through post-maturity molting|url=http://dx.doi.org/10.1636/b12-03.1|journal=Journal of Arachnology|volume=40|issue=3|pages=345–347|doi=10.1636/b12-03.1|issn=0161-8202}}</ref> This can be explained by the evolutionary selection for females with better fecundity.<ref name=":0" /> Female ''Nephila'' ''pilipes'' have huge parental investments to their progenies, including egg production and web construction (see [[#parental care|Parental care]] section).
''N. pilipes'' display female gigantism and male dwarfism (see [[#sexual dimorphism|Sexual dimorphism]] section). In terrestrial animals, ''N. pilipes'' have the most size differences between males and females.<ref name=":0">{{Cite journal|last=Kuntner|first=Matjaž|last2=Zhang|first2=Shichang|last3=Gregorič|first3=Matjaž|last4=Li|first4=Daiqin|date=November 2012|title=Nephila female gigantism attained through post-maturity molting|url=http://dx.doi.org/10.1636/b12-03.1|journal=Journal of Arachnology|volume=40|issue=3|pages=345–347|doi=10.1636/b12-03.1|issn=0161-8202}}</ref> This can be explained by the evolutionary selection for females with better fecundity.<ref name=":0" /> Female ''N''. ''pilipes'' have huge parental investments to their progenies, including egg production and web construction (see [[#parental care|Parental care]] section).


=== Female ===
=== Female ===
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=== Difference from close relatives ===
=== Difference from close relatives ===
Different from ''Nephila clavate'', this species ''Nephila pilipes'' has a horn-like bulge on its tergum.<ref name=":1" /> Also, to distinguish from ''Nephila laurinae'', ''Nephila pilipes'' has a pair of yellow stripes at its back of the abdomen.<ref name=":1" />
Different from ''Nephila clavate'', this species ''N. pilipes'' has a horn-like bulge on its tergum.<ref name=":1" /> Also, to distinguish from ''Nephila laurinae'', ''N. pilipes'' has a pair of yellow stripes at its back of the abdomen.<ref name=":1" />


== Sexual dimorphism ==
== Sexual dimorphism ==
[[File:Giant Wood Spider (Nephila pilipes) W IMG 2806.jpg|thumb|upright=0.8|Size comparison of female with many males]]
[[File:Giant Wood Spider (Nephila pilipes) W IMG 2806.jpg|thumb|upright=0.8|Size comparison of female with many males]]
''Nephila pilipes'' display [[sexual dimorphism]], the presence of distinct difference between the males and females of a species. In ''Nephila pilipes'', female spiders are much larger than their male counterparts,<ref name="ReferenceA"/> and males can be 4–10 times smaller than the females.<ref>{{cite journal|last1=Harvey|first1=Mark S.|last2=Austin|first2=Andrew D.|last3=Adams|first3=Mark|date=2007|title=The systematics and biology of the spider genus ''Nephila'' (Araneae : Nephilidae) in the Australasian region|url=http://www.arachne.org.au/_dbase_upl/NephilidaeHarvey.pdf|journal=Invertebrate Systematics|volume=21|issue=5|pages=407–451|doi=10.1071/IS05016|via=}}</ref>
''N. pilipes'' display [[sexual dimorphism]], the presence of distinct difference between the males and females of a species. In ''N. pilipes'', female spiders are much larger than their male counterparts,<ref name="ReferenceA"/> and males can be 4–10 times smaller than the females.<ref>{{cite journal|last1=Harvey|first1=Mark S.|last2=Austin|first2=Andrew D.|last3=Adams|first3=Mark|date=2007|title=The systematics and biology of the spider genus ''Nephila'' (Araneae : Nephilidae) in the Australasian region|url=http://www.arachne.org.au/_dbase_upl/NephilidaeHarvey.pdf|journal=Invertebrate Systematics|volume=21|issue=5|pages=407–451|doi=10.1071/IS05016|via=}}</ref>


Sexual dimorphism appears to be a shared feature among ''Nephila pilipes'' spiders of different populations, as examinations of populations in Southeast Asia, Australia, and Papua New Guinea show similar patterns of size dimorphism.<ref name="ReferenceC">{{cite journal|last1=Tso|first1=I-Min|last2=Shu-Ya|first2=Chiang|last3=Blackledge|first3=Todd|date=2007|title=Does The Giant Wood Spider ''Nephila pilipes'' Respond To Prey Variation By Altering Web Or Silk Properties?.|url=|journal=Ethology|volume=113|issue=4|pages=324–333|doi=10.1111/j.1439-0310.2007.01318.x}}</ref> Two major hypotheses have been suggested to explain this sexual selection and they are female gigantism and male dwarfism.
Sexual dimorphism appears to be a shared feature among ''N. pilipes'' spiders of different populations, as examinations of populations in Southeast Asia, Australia, and Papua New Guinea show similar patterns of size dimorphism.<ref name="ReferenceC">{{cite journal|last1=Tso|first1=I-Min|last2=Shu-Ya|first2=Chiang|last3=Blackledge|first3=Todd|date=2007|title=Does The Giant Wood Spider ''Nephila pilipes'' Respond To Prey Variation By Altering Web Or Silk Properties?.|url=|journal=Ethology|volume=113|issue=4|pages=324–333|doi=10.1111/j.1439-0310.2007.01318.x}}</ref> Two major hypotheses have been suggested to explain this sexual selection and they are female gigantism and male dwarfism.


=== Female gigantism ===
=== Female gigantism ===
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== Habitat and distribution ==
== Habitat and distribution ==
''Nephila pilipes'' prefers moist habitats with no direct sunlight. In Australia, most ''N. pilipes'' are found in rainforest habitats in northern and eastern Australia, where climate is humid and vegetation offers shade against direct sunlight.<ref name=":32">{{Cite journal|last=Harvey|first=Mark S.|last2=Austin|first2=Andrew D.|last3=Adams|first3=Mark|date=2007|title=The systematics and biology of the spider genus Nephila (Araneae:Nephilidae) in the Australasian region|url=http://dx.doi.org/10.1071/is05016|journal=Invertebrate Systematics|volume=21|issue=5|pages=407|doi=10.1071/is05016|issn=1445-5226}}</ref> In general, ''N. pilipes'' are distributed along coastal lines, where precipitation is ample.<ref name=":32"/> However, reports show that ''N. pilipes'' can be found in dry sclerophyll and low shrublands, hundred miles away from the coast.<ref name=":32"/> ''N. pilipes'' can survive in many climate types, including temperate coastal, Mediterranean, subtropical and tropical savannah climates.<ref name=":32"/> They construct webs in bushes and trees, near surface water, and against buildings and other terrain structures.<ref name=":32"/> To reduce heat from the sun, like other spiders, ''Nephila'' spp. has a thermoregulatory behavior.<ref name=":32"/>. Experiments show that when the temperature reaches 32°C, ''Nephila'' spp. will adjust the angle between its body and the incoming sunlight, orienting its abdomen towards the sun but keeping the cephalothorax parallel to the web.<ref name=":32"/> When temperature further rises, it will align the full body along the sunlight direction, further reducing the area that is receiving heat from the sun.<ref name=":32"/> At temperature above 40°C, ''Nephila'' spp. will abandon the web. Unlike other relatives, the distribution of ''N. pilipes'' doesn’t depend on seasons.<ref name=":32"/> Adult females are active throughout four seasons and continuously lay eggs. Adult males are present in the population for longer times than females.<ref name=":32"/>
''N. pilipes'' prefers moist habitats with no direct sunlight. It can be found in [[Japan]], [[China]], [[Vietnam]], [[Cambodia]], [[Taiwan]], [[Malaysia]], [[Singapore]], [[Myanmar]], [[Indonesia]], [[Thailand]], [[Laos]], [[Philippines]], [[Sri Lanka]], [[India]], [[Nepal]], [[Papua New Guinea]], and [[Australia]]. In Australia, most ''N. pilipes'' are found in rainforest habitats in northern and eastern Australia, where climate is humid and vegetation offers shade against direct sunlight.<ref name=":32">{{Cite journal|last=Harvey|first=Mark S.|last2=Austin|first2=Andrew D.|last3=Adams|first3=Mark|date=2007|title=The systematics and biology of the spider genus Nephila (Araneae:Nephilidae) in the Australasian region|url=http://dx.doi.org/10.1071/is05016|journal=Invertebrate Systematics|volume=21|issue=5|pages=407|doi=10.1071/is05016|issn=1445-5226}}</ref> In general, ''N. pilipes'' are distributed along coastal lines, where precipitation is ample.<ref name=":32"/> However, reports show that ''N. pilipes'' can be found in dry sclerophyll and low shrublands, hundred miles away from the coast.<ref name=":32"/> ''N. pilipes'' can survive in many climate types, including temperate coastal, Mediterranean, subtropical and tropical savannah climates.<ref name=":32"/> They construct webs in bushes and trees, near surface water, and against buildings and other terrain structures.<ref name=":32"/> To reduce heat from the sun, like other spiders, ''Nephila'' spp. has a thermoregulatory behavior.<ref name=":32"/>. Experiments show that when the temperature reaches 32°C, ''Nephila'' spp. will adjust the angle between its body and the incoming sunlight, orienting its abdomen towards the sun but keeping the cephalothorax parallel to the web.<ref name=":32"/> When temperature further rises, it will align the full body along the sunlight direction, further reducing the area that is receiving heat from the sun.<ref name=":32"/> At temperature above 40°C, ''Nephila'' spp. will abandon the web. Unlike other relatives, the distribution of ''N. pilipes'' doesn’t depend on seasons.<ref name=":32"/> Adult females are active throughout four seasons and continuously lay eggs. Adult males are present in the population for longer times than females.<ref name=":32"/>


== Diet ==
== Diet ==
''Nephila'' spp. are somewhat selective on their preys.<ref name=":32"/> Most of their preys are only a few species.<ref name=":32"/> Some studies also reports that ''Nephila'' spp. will remove some specific insects from their webs.<ref name=":32"/> They avoid vespid wasps, alate ants, and other insects that secrete distasteful compounds.<ref name=":32"/> Due to ''Nephila’s'' large body size, it can prey on insects with a broad size range, from 2&nbsp;mm to even larger size than themselves.<ref name=":32"/> They adopt different strategies for different sizes of preys.<ref name=":32"/> Small preys are directly caught and removed from the web.<ref name=":32"/> For larger ones, they inject venom and wait for the decease of their prey.<ref name=":32"/> However, ''Nephila'' spp. have a dramatic change in predation style from the spiderling to adulthood. During the spiderling stage, they feed collectively on a common web.<ref name=":32"/> Upon reaching adulthood, they construct their own webs.<ref name=":32"/> ''Nephila'' spp. also reserve food caches to deal with periods of food shortage.<ref name=":32"/> They wrap surplus prey in silk and store them in the hub of the web.<ref name=":32"/> The silk covering can significantly reduce water loss due to evaporation.<ref name=":32"/> Food cache can reduce weight loss during periods with limited preys.<ref name=":32"/>
''Nephila'' spp. prey upon only a few species.<ref name=":32"/> Some studies also reports that ''Nephila'' spp. will remove some specific insects from their webs.<ref name=":32"/> They avoid vespid wasps, alate ants, and other insects that secrete distasteful compounds.<ref name=":32"/> Due to ''Nephila’s'' large body size, it can prey on insects with a broad size range, from 2&nbsp;mm to even larger size than themselves.<ref name=":32"/> They adopt different strategies for different sizes of preys.<ref name=":32"/> Small preys are directly caught and removed from the web.<ref name=":32"/> For larger ones, they inject venom and wait for the decease of their prey.<ref name=":32"/> However, ''Nephila'' spp. have a dramatic change in predation style from the spiderling to adulthood. During the spiderling stage, they feed collectively on a common web.<ref name=":32"/> Upon reaching adulthood, they construct their own webs.<ref name=":32"/> ''Nephila'' spp. also reserve food caches to deal with periods of food shortage.<ref name=":32"/> They wrap surplus prey in silk and store them in the hub of the web.<ref name=":32"/> The silk covering can significantly reduce water loss due to evaporation.<ref name=":32"/> Food cache can reduce weight loss during periods with limited preys.<ref name=":32"/>


It has been reported that some ''Nephila pilipes'' can adjust the ultraviolet radiation reflected by stripes on their body to attract preys that are UV light oriented.<ref name=":2" /><ref name=":4">{{Cite journal|last=Chiao|first=C.-C.|last2=Wu|first2=W.-Y.|last3=Chen|first3=S.-H.|last4=Yang|first4=E.-C.|date=2009-06-26|title=Visualization of the spatial and spectral signals of orb-weaving spiders, Nephila pilipes, through the eyes of a honeybee|url=http://dx.doi.org/10.1242/jeb.030734|journal=Journal of Experimental Biology|volume=212|issue=14|pages=2269–2278|doi=10.1242/jeb.030734|issn=0022-0949}}</ref> The bands on the legs and body of ''Nephila pilipes'' can reflect UV radiation, so they are more visible to flying insects.<ref name=":2" /> The contrast of colors makes them look like food sources for insects, rather than a predator spider.<ref name=":4" />
It has been reported that some ''N. pilipes'' can adjust the ultraviolet radiation reflected by stripes on their body to attract preys that are UV light oriented.<ref name=":2" /><ref name=":4">{{Cite journal|last=Chiao|first=C.-C.|last2=Wu|first2=W.-Y.|last3=Chen|first3=S.-H.|last4=Yang|first4=E.-C.|date=2009-06-26|title=Visualization of the spatial and spectral signals of orb-weaving spiders, Nephila pilipes, through the eyes of a honeybee|url=http://dx.doi.org/10.1242/jeb.030734|journal=Journal of Experimental Biology|volume=212|issue=14|pages=2269–2278|doi=10.1242/jeb.030734|issn=0022-0949}}</ref> The bands on the legs and body of ''N. pilipes'' can reflect UV radiation, so they are more visible to flying insects.<ref name=":2" /> The contrast of colors makes them look like food sources for insects, rather than a predator spider.<ref name=":4" />


== Web structure ==
== Web structure ==
=== Web type ===
=== Web type ===
Generally, ''Nephila'' spp.'s web is not symmetric.<ref name=":5">{{Cite journal|last=Tso|first=I-Min|last2=Chiang|first2=Shu-Ya|last3=Blackledge|first3=Todd A.|date=2007-03-27|title=Does the Giant Wood Spider Nephila pilipes Respond to Prey Variation by Altering Web or Silk Properties?|url=http://dx.doi.org/10.1111/j.1439-0310.2007.01318.x|journal=Ethology|volume=113|issue=4|pages=324–333|doi=10.1111/j.1439-0310.2007.01318.x|issn=0179-1613}}</ref> Their silk appears to be yellow, hence the name [[golden orb-weavers]].<ref name=":5" /> For adult ''Nephila'', their webs are typically 0.5–1.0 m in diameter.<ref name=":5" /> However, when females aggregate together, their webs can be even larger.<ref name=":5" />. The hub of the web is usually at the top.<ref name=":5" />. Preys will mostly be trapped on the lower web.<ref name=":5" />
Generally, ''Nephila'' spp.'s web is not symmetric.<ref name=":5">{{Cite journal|last=Tso|first=I-Min|last2=Chiang|first2=Shu-Ya|last3=Blackledge|first3=Todd A.|date=2007-03-27|title=Does the Giant Wood Spider Nephila pilipes Respond to Prey Variation by Altering Web or Silk Properties?|url=http://dx.doi.org/10.1111/j.1439-0310.2007.01318.x|journal=Ethology|volume=113|issue=4|pages=324–333|doi=10.1111/j.1439-0310.2007.01318.x|issn=0179-1613}}</ref> Their silk appears to be yellow, hence the name [[golden orb-weavers]].<ref name=":5" /> For adult ''Nephila spp.'', their webs are typically 0.5–1.0 m in diameter.<ref name=":5" /> However, when females aggregate together, their webs can be even larger.<ref name=":5" />. The hub of the web is usually at the top.<ref name=":5" />. Preys will mostly be trapped on the lower web.<ref name=":5" />


=== Construction ===
=== Construction ===
The web constructed by ''Nephila pilipes'' has elastic silk at the center to absorb the kinetic energy of moving insects.<ref name=":5" /> Around the elastic part, some stiff structures fix the web to trees or other terrain structures.<ref name=":5" /> It has been reported that ''Nephila pilipes'' can adjust their silk composition, responding to different diet and environmental conditions.<ref name=":6">{{Cite journal|last=Tso|first=I-M.|date=2005-03-15|title=Giant wood spider Nephila pilipes alters silk protein in response to prey variation|url=http://dx.doi.org/10.1242/jeb.01437|journal=Journal of Experimental Biology|volume=208|issue=6|pages=1053–1061|doi=10.1242/jeb.01437|issn=0022-0949}}</ref> One study investigated the relationship between silk construction and prey types.<ref name=":6" /> The study found that when ''Nephila pilipes'' are fed with flies, small and airborne prey, the silk becomes more elastic, and the mesh size becomes smaller.<ref name=":6" /> When ''Nephila pilipes'' are fed crickets, large and powerful prey, the silk becomes stiff.<ref name=":6" /> Moreover, researchers also found that ''Nephila pilipes'' can change the silk's amino acid composition to fit their needs.<ref name=":6" /> The silk has excellent mechanical properties, more robust than most human-made materials, so the silk of ''Nephila'' spp. draws much attention in the field of [[materials science]].<ref name=":7">{{Cite journal|last=Wang|first=Zuyuan|last2=Cang|first2=Yu|last3=Kremer|first3=Friedrich|last4=Thomas|first4=Edwin L.|last5=Fytas|first5=George|date=2020-01-14|title=Determination of the Complete Elasticity of Nephila pilipes Spider Silk|url=http://dx.doi.org/10.1021/acs.biomac.9b01607|journal=Biomacromolecules|volume=21|issue=3|pages=1179–1185|doi=10.1021/acs.biomac.9b01607|issn=1525-7797}}</ref> However, when silks are artificially spun in the lab, they often have inferior quality.<ref name=":7" />
The web constructed by ''N. pilipes'' has elastic silk at the center to absorb the kinetic energy of moving insects.<ref name=":5" /> Around the elastic part, some stiff structures fix the web to trees or other terrain structures.<ref name=":5" /> It has been reported that ''N. pilipes'' can adjust their silk composition, responding to different diet and environmental conditions.<ref name=":6">{{Cite journal|last=Tso|first=I-M.|date=2005-03-15|title=Giant wood spider Nephila pilipes alters silk protein in response to prey variation|url=http://dx.doi.org/10.1242/jeb.01437|journal=Journal of Experimental Biology|volume=208|issue=6|pages=1053–1061|doi=10.1242/jeb.01437|issn=0022-0949}}</ref> One study investigated the relationship between silk construction and prey types.<ref name=":6" /> The study found that when ''N. pilipes'' are fed with flies, small and airborne prey, the silk becomes more elastic, and the mesh size becomes smaller.<ref name=":6" /> When ''N. pilipes'' are fed crickets, large and powerful prey, the silk becomes stiff.<ref name=":6" /> Moreover, researchers also found that ''N. pilipes'' can change the silk's amino acid composition to fit their needs.<ref name=":6" /> The silk has excellent mechanical properties, more robust than most human-made materials, so the silk of ''Nephila'' spp. draws much attention in the field of [[materials science]].<ref name=":7">{{Cite journal|last=Wang|first=Zuyuan|last2=Cang|first2=Yu|last3=Kremer|first3=Friedrich|last4=Thomas|first4=Edwin L.|last5=Fytas|first5=George|date=2020-01-14|title=Determination of the Complete Elasticity of Nephila pilipes Spider Silk|url=http://dx.doi.org/10.1021/acs.biomac.9b01607|journal=Biomacromolecules|volume=21|issue=3|pages=1179–1185|doi=10.1021/acs.biomac.9b01607|issn=1525-7797}}</ref> However, when silks are artificially spun in the lab, they often have inferior quality.<ref name=":7" />


=== Web damage and repair ===
=== Web damage and repair ===
Their webs, like other spiders, are subject to potential damage by birds and small mammals.<ref name=":32"/> When their webs are partially damaged, it usually takes ''Nephila pilipes'' 10–60 minutes to repair the web.<ref name=":32"/> However, if the damage is critical, the spider will consume the web and construct a new web in another nearby location.<ref name=":32"/>
Their webs, like other spiders, are subject to potential damage by birds and small mammals.<ref name=":32"/> When their webs are partially damaged, it usually takes ''N. pilipes'' 10–60 minutes to repair the web.<ref name=":32"/> However, if the damage is critical, the spider will consume the web and construct a new web in another nearby location.<ref name=":32"/>


== Mating ==
== Mating ==
=== Mate searching behavior ===
=== Mate searching behavior ===
Upon reaching adulthood, males leave their webs and begin the search for females.<ref name=":32"/> They look for the correct web chemical compositions and web characteristics.<ref name=":32"/> It is reported that ''Nephila pilipes'' do not have an airborne pheromone-based signaling system for mating.<ref name=":32"/>
Upon reaching adulthood, males leave their webs and begin the search for females.<ref name=":32"/> They look for the correct web chemical compositions and web characteristics.<ref name=":32"/> It is reported that ''N. pilipes'' do not have an airborne pheromone-based signaling system for mating.<ref name=":32"/>


=== Mating interactions ===
=== Mating interactions ===
In nature, sexual conflicts between males and females are very frequent because males and females have very different reproduction purposes. Males have relatively less parental investment than females, so their goal is to fertilize as many females as possible. Females, on the other hand, tend to be reluctant to mate repeatedly and only select those males with good genes. Female spiders can display aggressiveness in a variety of ways: shake the web and chase the male when approaching; kick the male when the male makes contact with the female’s leg; violently shake body when the male climbs up the female's dorsum; abruptly interrupt mating by kick the male off.<ref name=":8">{{Cite journal|last=Zhang|first=Shichang|last2=Kuntner|first2=Matjaž|last3=Li|first3=Daiqin|date=December 2011|title=Mate binding: male adaptation to sexual conflict in the golden orb-web spider (Nephilidae: Nephila pilipes)|url=http://dx.doi.org/10.1016/j.anbehav.2011.09.010|journal=Animal Behaviour|volume=82|issue=6|pages=1299–1304|doi=10.1016/j.anbehav.2011.09.010|issn=0003-3472}}</ref> Unlike other close relatives, ''Nephila pilipes'' have a less aggressive mating ritual, and sexual cannibalism is very rare.<ref name=":32"/> Males have evolved a mechanism called ''mate binding'' to avoid females' resistance and cannibalism.<ref name=":8" /> It has been reported that males can deposit silks soaked with sexual hormones onto the female’s body to calm the tactile and chemical receptors.<ref name=":8" /> In this way, males can reduce cannibalism and maximize the number of females they fertilize.<ref name=":8" />
In nature, sexual conflicts between males and females are very frequent because males and females have very different reproduction purposes. Males have relatively less parental investment than females, so their goal is to fertilize as many females as possible. Females, on the other hand, tend to be reluctant to mate repeatedly and only select those males with good genes. Female spiders can display aggressiveness in a variety of ways: shake the web and chase the male when approaching; kick the male when the male makes contact with the female’s leg; violently shake body when the male climbs up the female's dorsum; abruptly interrupt mating by kick the male off.<ref name=":8">{{Cite journal|last=Zhang|first=Shichang|last2=Kuntner|first2=Matjaž|last3=Li|first3=Daiqin|date=December 2011|title=Mate binding: male adaptation to sexual conflict in the golden orb-web spider (Nephilidae: Nephila pilipes)|url=http://dx.doi.org/10.1016/j.anbehav.2011.09.010|journal=Animal Behaviour|volume=82|issue=6|pages=1299–1304|doi=10.1016/j.anbehav.2011.09.010|issn=0003-3472}}</ref> Unlike other close relatives, ''N. pilipes'' have a less aggressive mating ritual, and sexual cannibalism is very rare.<ref name=":32"/> Males have evolved a mechanism called ''mate binding'' to avoid females' resistance and cannibalism.<ref name=":8" /> It has been reported that males can deposit silks soaked with sexual hormones onto the female’s body to calm the tactile and chemical receptors.<ref name=":8" /> In this way, males can reduce cannibalism and maximize the number of females they fertilize.<ref name=":8" />


== Parental care ==
== Parental care ==
=== Oviposition ===
=== Oviposition ===
Other ''Nephila'' spp. lay eggs near their webs and cover the eggmass with a thick layer of silk.<ref name=":32"/> Then the eggmass is firmly tied to leaves and tree branches around it.<ref name=":32"/> The eggs are always found beneath leaves or other shades, which prevent direct UV light from the sun.<ref name=":32"/> However, ''Nephila pilipes'' is different from all other ''Nephila'' spp.<ref name=":32"/> Female ''Nephila pilipes'' lay eggs in small pits on the ground to avoid predation and parasitism.<ref name=":32"/>
Other ''Nephila'' spp. lay eggs near their webs and cover the eggmass with a thick layer of silk.<ref name=":32"/> Then the eggmass is firmly tied to leaves and tree branches around it.<ref name=":32"/> The eggs are always found beneath leaves or other shades, which prevent direct UV light from the sun.<ref name=":32"/> However, ''N. pilipes'' is different from all other ''Nephila'' spp.<ref name=":32"/> Female ''N. pilipes'' lay eggs in small pits on the ground to avoid predation and parasitism.<ref name=":32"/>


== Predators ==
== Predators ==
Few studies have been conducted to investigate the enemies of ''Nephila pilipes''.<ref name=":32"/> The primary enemy of ''Nephila pilipes'' in nature is the bird, which evolves to fly by and take them without being entangled by the web.<ref name=":32"/> It is rarely reported that ''Nephila pilipes'' are attacked and killed by parasites like Hymenoptera.<ref name=":32"/> To avoid predation by birds, female ''Nephila'' often construct an aggregated web system with other females or other orb-weavers, so their web system can shield them from birds.<ref name=":32"/>
Few studies have been conducted to investigate the enemies of ''N. pilipes''.<ref name=":32"/> The primary enemy of ''N. pilipes'' in nature is the bird, which evolves to fly by and take them without being entangled by the web.<ref name=":32"/> It is rarely reported that ''N. pilipes'' are attacked and killed by parasites like Hymenoptera.<ref name=":32"/> To avoid predation by birds, female ''Nephila'' often construct an aggregated web system with other females or other orb-weavers, so their web system can shield them from birds.<ref name=":32"/>


== Bites to humans ==
== Bites to humans ==
The bite of ''Nephila pilipes'' to humans is rarely reported. Other orb-weaving spiders are reported to cause acute symptoms, including muscle pain, feeling of tightness, and reflexes exaggeration.<ref name=":9">{{Cite journal|last=Wise|first=David H.|date=March 1981|title=Inter-and intraspecific effects of density manipulations upon females of two orb-weaving spiders (araneae: araneidae)|url=http://dx.doi.org/10.1007/bf00347972|journal=Oecologia|volume=48|issue=2|pages=252–256|doi=10.1007/bf00347972|issn=0029-8549}}</ref> Treatment with [[calcium gluconate]] can relieve victims from acute pains. [[Antiserum]] treatment can speed the victim's recovery.<ref name=":9" />
The bite of ''N. pilipes'' to humans is rarely reported. Other orb-weaving spiders are reported to cause acute symptoms, including muscle pain, feeling of tightness, and reflexes exaggeration.<ref name=":9">{{Cite journal|last=Wise|first=David H.|date=March 1981|title=Inter-and intraspecific effects of density manipulations upon females of two orb-weaving spiders (araneae: araneidae)|url=http://dx.doi.org/10.1007/bf00347972|journal=Oecologia|volume=48|issue=2|pages=252–256|doi=10.1007/bf00347972|issn=0029-8549}}</ref> Treatment with [[calcium gluconate]] can relieve victims from acute pains. [[Antiserum]] treatment can speed the victim's recovery.<ref name=":9" />


== Ballooning ==
== Ballooning ==
[[Ballooning (spider)|Balloning]] is a common behavior among spiders.<ref name=":10">{{Cite journal|last=Lee|first=Vanessa M. J.|last2=Kuntner|first2=Matjaž|last3=Li|first3=Daiqin|date=2015-01-21|title=Ballooning behavior in the golden orbweb spider Nephila pilipes (Araneae: Nephilidae)|url=http://dx.doi.org/10.3389/fevo.2015.00002|journal=Frontiers in Ecology and Evolution|volume=3|doi=10.3389/fevo.2015.00002|issn=2296-701X}}</ref> Spiderlings can disperse via wind to larger areas after birth to avoid overcrowded habitat and competition for resources.<ref name=":10" /> Research conducted in lab has found that ''Nephila pilipes'' will display ballooning behavior when the wind speed reaches a threshold of 3.17&nbsp;m/s.<ref name=":10" /> However, ballooning is subject to many influences in wild habitats, such as wind angles, humidity, temperature, and pressure.<ref name=":10" /> Ballooning is also a social behavior.<ref name=":10" /> When the majority of spiderlings balloons, some spiderlings will sense the decline of the population and stay in the original place.<ref name=":10" /> Ballooning behavior explains the wide spread of ''Nephila pilipes'', and sometimes colonization of islands.<ref name=":10" />
[[Ballooning (spider)|Balloning]] is a common behavior among spiders.<ref name=":10">{{Cite journal|last=Lee|first=Vanessa M. J.|last2=Kuntner|first2=Matjaž|last3=Li|first3=Daiqin|date=2015-01-21|title=Ballooning behavior in the golden orbweb spider Nephila pilipes (Araneae: Nephilidae)|url=http://dx.doi.org/10.3389/fevo.2015.00002|journal=Frontiers in Ecology and Evolution|volume=3|doi=10.3389/fevo.2015.00002|issn=2296-701X}}</ref> Spiderlings can disperse via wind to larger areas after birth to avoid overcrowded habitat and competition for resources.<ref name=":10" /> Research conducted in lab has found that ''N. pilipes'' will display ballooning behavior when the wind speed reaches a threshold of 3.17&nbsp;m/s.<ref name=":10" /> However, ballooning is subject to many influences in wild habitats, such as wind angles, humidity, temperature, and pressure.<ref name=":10" /> Ballooning is also a social behavior.<ref name=":10" /> When the majority of spiderlings balloons, some spiderlings will sense the decline of the population and stay in the original place.<ref name=":10" /> Ballooning behavior explains the wide spread of ''N. pilipes'', and sometimes colonization of islands.<ref name=":10" />


== As food ==
== As food ==

Revision as of 23:18, 3 November 2020

Nephila pilipes
Dorsal side
Ventral side
both Madhya Pradesh, India
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Order: Araneae
Infraorder: Araneomorphae
Family: Nephilidae
Genus: Nephila
Species:
N. pilipes
Binomial name
Nephila pilipes
Synonyms

Aranea longipes
Aranea maculata
Aranea pilipes
Aranea sebae
Epeira chrysogaster
Nephila maculata
Nephila fuscipes
Epeira fuscipes
Epeira doreyana
Epeira caliginosa
Nephila ornata
Epeira penicillum
Epeira harpyia
Nephila chrysogaster
Meta ornata
Nephila pecuniosa
Nephila aurosa
Nephila procera
Nephila sulphurosa
Nephila tenuipes
Nephila submaculata

Nephila pilipes (northern golden orb weaver or giant golden orb weaver[2]) is a species of golden orb-web spider. It resides all over countries in East and Southeast Asia as well as Oceania. It is commonly found in primary and secondary forests and gardens. Females are large and grow to a body size of 30–50 mm (overall size up to 20 cm), with males growing to 5–6 mm. It is the second largest of the orb-weaving spiders apart from the recently discovered Nephila komaci. The first, second, and fourth pairs of legs of juvenile females have dense hairy brushes, but these brushes disappear as the spider matures.

The N. pilipes golden web is vertical with a fine irregular mesh. It is not symmetrical, with the hub usually nearer the top.[2] Rather than egg sacs being hung in the web, a pit is dug which is then covered with plant debris or soil.


Subspecies

Description

N. pilipes display female gigantism and male dwarfism (see Sexual dimorphism section). In terrestrial animals, N. pilipes have the most size differences between males and females.[3] This can be explained by the evolutionary selection for females with better fecundity.[3] Female N. pilipes have huge parental investments to their progenies, including egg production and web construction (see Parental care section).

Female

Females typically have a body size of 30–50 mm.[4] The cephalothorax is about 15 mm long, 10 mm wide.[4] The abdomen is about 30 mm long, 15 mm wide, mostly dark yellow-brown color with yellow stripes.[4] The tergum is generally black or brown, covered with dense hairs.[4] Both rows of eyes bulge towards the rear.[4] The plastron is mostly black and brown.[4] The legs span very long, and they are black and yellow.[4] There are no apparent hairs on all legs.[4] It has been reported that this sharp contrast between yellow and black color can increase foraging success towards visually-oriented prey.[5]

Male

Males’ body size is typically 5 mm to 6.5 mm.[4] The cephalothorax is about 2.5 mm long, 2 mm wide.[4] The abdomen is about 4 mm long, 1.5 mm wide.[4] Front eyes are larger than rear eyes.[4] Males have light brown legs, with some hairs.[4] The carapace is yellow with very few hairs.[4]

Difference from close relatives

Different from Nephila clavate, this species N. pilipes has a horn-like bulge on its tergum.[4] Also, to distinguish from Nephila laurinae, N. pilipes has a pair of yellow stripes at its back of the abdomen.[4]

Sexual dimorphism

Size comparison of female with many males

N. pilipes display sexual dimorphism, the presence of distinct difference between the males and females of a species. In N. pilipes, female spiders are much larger than their male counterparts,[6] and males can be 4–10 times smaller than the females.[7]

Sexual dimorphism appears to be a shared feature among N. pilipes spiders of different populations, as examinations of populations in Southeast Asia, Australia, and Papua New Guinea show similar patterns of size dimorphism.[8] Two major hypotheses have been suggested to explain this sexual selection and they are female gigantism and male dwarfism.

Female gigantism

One explanation for the dimorphism observed in the N. pilipes species is sexual size dimorphism may have evolved due to selection favoring female gigantism. Gigantism may have evolved in response to reduce the effectiveness of male mating plugs.[9] With a mating plug, male spiders ensure that only the male who create the plug post coitus would be the father of the resulting offspring.[9] Although in the species of N. pilipes, plugging does not seem to have an effect as females are still able to have successful matings even with multiple plugs.[9] The evolution of plugs occurred in earlier ancestral species and still persists along. These traits were then passed down to one of its descendants, the living species N. pilipes.[10] In this ancestral species, female gigantism was selected as a positive adaptation. In female spiders with larger bodies, the "embolic plugs" inserted in the N. pilipes females' genitalia become too thin to effectively seal the genitalia.[11] Additionally, female gigantism would have been important to the ancestral species at the time when mating plugs were still effective, as body size has been shown to increase fecundity.[12] By laying more eggs at a time, the ancestral females could have produced more offspring before they were plugged by a male.[8]

Female N. pilipes spiders are able to achieve a large size because they can continue to molt and grow after maturity.[6] This contrasts with most spiders, where growth stops once sexual maturity is reached. Female N. pilipes spiders will stop molting, however, during times of high copulation where it may not be advantageous to continue to grow as sperm are fertilizing eggs.[6]

Male dwarfism

Alternatively, sexual size dimorphism may have evolved due to selection favoring male dwarfism instead of female giganticism. An explanation of how dwarfism was selected for is due to scramble competition in which smaller male size is advantageous for reproduction. In N. pilipes spiders specifically, male spiders that were able to find female spiders first often fertilize a greater percentage of their eggs than other males.[13] Also, because female N. pilipes spiders often do not remain on their webs during sexual maturity, the smaller and more agile male spiders were able to reach them first, compared to the slower, larger males which often waited at the web of the female in advantageous spots.[13] This explanation means that smaller males had a greater fitness due to the behavior of the female spider to move around in maturity as compared to that of larger males who did not move as much and often copulated with females in their webs after smaller males had already done so.[13]

Habitat and distribution

N. pilipes prefers moist habitats with no direct sunlight. It can be found in Japan, China, Vietnam, Cambodia, Taiwan, Malaysia, Singapore, Myanmar, Indonesia, Thailand, Laos, Philippines, Sri Lanka, India, Nepal, Papua New Guinea, and Australia. In Australia, most N. pilipes are found in rainforest habitats in northern and eastern Australia, where climate is humid and vegetation offers shade against direct sunlight.[14] In general, N. pilipes are distributed along coastal lines, where precipitation is ample.[14] However, reports show that N. pilipes can be found in dry sclerophyll and low shrublands, hundred miles away from the coast.[14] N. pilipes can survive in many climate types, including temperate coastal, Mediterranean, subtropical and tropical savannah climates.[14] They construct webs in bushes and trees, near surface water, and against buildings and other terrain structures.[14] To reduce heat from the sun, like other spiders, Nephila spp. has a thermoregulatory behavior.[14]. Experiments show that when the temperature reaches 32°C, Nephila spp. will adjust the angle between its body and the incoming sunlight, orienting its abdomen towards the sun but keeping the cephalothorax parallel to the web.[14] When temperature further rises, it will align the full body along the sunlight direction, further reducing the area that is receiving heat from the sun.[14] At temperature above 40°C, Nephila spp. will abandon the web. Unlike other relatives, the distribution of N. pilipes doesn’t depend on seasons.[14] Adult females are active throughout four seasons and continuously lay eggs. Adult males are present in the population for longer times than females.[14]

Diet

Nephila spp. prey upon only a few species.[14] Some studies also reports that Nephila spp. will remove some specific insects from their webs.[14] They avoid vespid wasps, alate ants, and other insects that secrete distasteful compounds.[14] Due to Nephila’s large body size, it can prey on insects with a broad size range, from 2 mm to even larger size than themselves.[14] They adopt different strategies for different sizes of preys.[14] Small preys are directly caught and removed from the web.[14] For larger ones, they inject venom and wait for the decease of their prey.[14] However, Nephila spp. have a dramatic change in predation style from the spiderling to adulthood. During the spiderling stage, they feed collectively on a common web.[14] Upon reaching adulthood, they construct their own webs.[14] Nephila spp. also reserve food caches to deal with periods of food shortage.[14] They wrap surplus prey in silk and store them in the hub of the web.[14] The silk covering can significantly reduce water loss due to evaporation.[14] Food cache can reduce weight loss during periods with limited preys.[14]

It has been reported that some N. pilipes can adjust the ultraviolet radiation reflected by stripes on their body to attract preys that are UV light oriented.[5][15] The bands on the legs and body of N. pilipes can reflect UV radiation, so they are more visible to flying insects.[5] The contrast of colors makes them look like food sources for insects, rather than a predator spider.[15]

Web structure

Web type

Generally, Nephila spp.'s web is not symmetric.[16] Their silk appears to be yellow, hence the name golden orb-weavers.[16] For adult Nephila spp., their webs are typically 0.5–1.0 m in diameter.[16] However, when females aggregate together, their webs can be even larger.[16]. The hub of the web is usually at the top.[16]. Preys will mostly be trapped on the lower web.[16]

Construction

The web constructed by N. pilipes has elastic silk at the center to absorb the kinetic energy of moving insects.[16] Around the elastic part, some stiff structures fix the web to trees or other terrain structures.[16] It has been reported that N. pilipes can adjust their silk composition, responding to different diet and environmental conditions.[17] One study investigated the relationship between silk construction and prey types.[17] The study found that when N. pilipes are fed with flies, small and airborne prey, the silk becomes more elastic, and the mesh size becomes smaller.[17] When N. pilipes are fed crickets, large and powerful prey, the silk becomes stiff.[17] Moreover, researchers also found that N. pilipes can change the silk's amino acid composition to fit their needs.[17] The silk has excellent mechanical properties, more robust than most human-made materials, so the silk of Nephila spp. draws much attention in the field of materials science.[18] However, when silks are artificially spun in the lab, they often have inferior quality.[18]

Web damage and repair

Their webs, like other spiders, are subject to potential damage by birds and small mammals.[14] When their webs are partially damaged, it usually takes N. pilipes 10–60 minutes to repair the web.[14] However, if the damage is critical, the spider will consume the web and construct a new web in another nearby location.[14]

Mating

Mate searching behavior

Upon reaching adulthood, males leave their webs and begin the search for females.[14] They look for the correct web chemical compositions and web characteristics.[14] It is reported that N. pilipes do not have an airborne pheromone-based signaling system for mating.[14]

Mating interactions

In nature, sexual conflicts between males and females are very frequent because males and females have very different reproduction purposes. Males have relatively less parental investment than females, so their goal is to fertilize as many females as possible. Females, on the other hand, tend to be reluctant to mate repeatedly and only select those males with good genes. Female spiders can display aggressiveness in a variety of ways: shake the web and chase the male when approaching; kick the male when the male makes contact with the female’s leg; violently shake body when the male climbs up the female's dorsum; abruptly interrupt mating by kick the male off.[19] Unlike other close relatives, N. pilipes have a less aggressive mating ritual, and sexual cannibalism is very rare.[14] Males have evolved a mechanism called mate binding to avoid females' resistance and cannibalism.[19] It has been reported that males can deposit silks soaked with sexual hormones onto the female’s body to calm the tactile and chemical receptors.[19] In this way, males can reduce cannibalism and maximize the number of females they fertilize.[19]

Parental care

Oviposition

Other Nephila spp. lay eggs near their webs and cover the eggmass with a thick layer of silk.[14] Then the eggmass is firmly tied to leaves and tree branches around it.[14] The eggs are always found beneath leaves or other shades, which prevent direct UV light from the sun.[14] However, N. pilipes is different from all other Nephila spp.[14] Female N. pilipes lay eggs in small pits on the ground to avoid predation and parasitism.[14]

Predators

Few studies have been conducted to investigate the enemies of N. pilipes.[14] The primary enemy of N. pilipes in nature is the bird, which evolves to fly by and take them without being entangled by the web.[14] It is rarely reported that N. pilipes are attacked and killed by parasites like Hymenoptera.[14] To avoid predation by birds, female Nephila often construct an aggregated web system with other females or other orb-weavers, so their web system can shield them from birds.[14]

Bites to humans

The bite of N. pilipes to humans is rarely reported. Other orb-weaving spiders are reported to cause acute symptoms, including muscle pain, feeling of tightness, and reflexes exaggeration.[20] Treatment with calcium gluconate can relieve victims from acute pains. Antiserum treatment can speed the victim's recovery.[20]

Ballooning

Balloning is a common behavior among spiders.[21] Spiderlings can disperse via wind to larger areas after birth to avoid overcrowded habitat and competition for resources.[21] Research conducted in lab has found that N. pilipes will display ballooning behavior when the wind speed reaches a threshold of 3.17 m/s.[21] However, ballooning is subject to many influences in wild habitats, such as wind angles, humidity, temperature, and pressure.[21] Ballooning is also a social behavior.[21] When the majority of spiderlings balloons, some spiderlings will sense the decline of the population and stay in the original place.[21] Ballooning behavior explains the wide spread of N. pilipes, and sometimes colonization of islands.[21]

As food

Raglai people in Vietnamese Bình Thuận Province consume the spiders after roasting, and consider them a healthy food source. These spiders can save local people who live in the forest from hunger if they run out of food.[22]

Different views and aspects

References

  1. ^ "Taxon details Nephila pilipes (Fabricius, 1793)". World Spider Catalog. Natural History Museum Bern. Retrieved 2017-05-15.
  2. ^ a b Nephila pilipes, Arachne.org.au
  3. ^ a b Kuntner, Matjaž; Zhang, Shichang; Gregorič, Matjaž; Li, Daiqin (November 2012). "Nephila female gigantism attained through post-maturity molting". Journal of Arachnology. 40 (3): 345–347. doi:10.1636/b12-03.1. ISSN 0161-8202.
  4. ^ a b c d e f g h i j k l m n o p Zhu, M. Chinese Zoology. 35: 402. {{cite journal}}: Missing or empty |title= (help)
  5. ^ a b c Tso, I-M. (2004-07-01). "Colourful orb-weaving spiders, Nephila pilipes, through a bee's eyes". Journal of Experimental Biology. 207 (15): 2631–2637. doi:10.1242/jeb.01068. ISSN 0022-0949.
  6. ^ a b c Kuntner, Matjaž; Zhang, Shichang; Gregorič, Matjaž; Li, Daiqin (2012). "Nephila Female Gigantism Attained through Post-maturity Molting". Journal of Arachnology. 40 (3): 345–347. doi:10.1636/b12-03.1. S2CID 30430184.
  7. ^ Harvey, Mark S.; Austin, Andrew D.; Adams, Mark (2007). "The systematics and biology of the spider genus Nephila (Araneae : Nephilidae) in the Australasian region" (PDF). Invertebrate Systematics. 21 (5): 407–451. doi:10.1071/IS05016.
  8. ^ a b Tso, I-Min; Shu-Ya, Chiang; Blackledge, Todd (2007). "Does The Giant Wood Spider Nephila pilipes Respond To Prey Variation By Altering Web Or Silk Properties?". Ethology. 113 (4): 324–333. doi:10.1111/j.1439-0310.2007.01318.x.
  9. ^ a b c Kuntner, Matjaž; Coddington, Jonathan; Schneider, Jutta (2009). "Intersexual Arms Race? Genital Coevolution In Nephilid Spiders (Araneae, Nephilidae)". Evolution. 63 (6): 1451–1463. doi:10.1111/j.1558-5646.2009.00634.x. PMID 19492993. S2CID 6321371.
  10. ^ Coddington, Jonathan A.; Hormiga, Gustavo; Scharff, Nikolaj (1997). "Giant female or dwarf male spiders?". Nature. 385 (6618): 687–688. doi:10.1038/385687a0. S2CID 4308746.
  11. ^ Kuntner, Matjaž.; Kralj-Fišer, S.; Schneider, Jutta; Li, Daiqin (2009). "Mate Plugging Via Genital Mutilation In Nephilid Spiders: An Evolutionary Hypothesis". Journal of Zoology. 277 (4): 257–266. doi:10.1111/j.1469-7998.2008.00533.x.
  12. ^ Legrand, Rebecca S.; Morse, Douglass H. (2000). "Factors driving extreme sexual size dimorphism of a sit-and-wait predator under low density". Biological Journal of the Linnean Society. 71 (4): 643–664. doi:10.1006/bijl.2000.0466.
  13. ^ a b c Danielson-François, Anne; Hou, Chueh; Cole, Nina; Tso, I-Min (2012). "Scramble Competition For Moulting Females As A Driving Force For Extreme Male Dwarfism In Spiders". Animal Behaviour. 84 (4): 937–945. doi:10.1016/j.anbehav.2012.07.018. S2CID 53186187.
  14. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am Harvey, Mark S.; Austin, Andrew D.; Adams, Mark (2007). "The systematics and biology of the spider genus Nephila (Araneae:Nephilidae) in the Australasian region". Invertebrate Systematics. 21 (5): 407. doi:10.1071/is05016. ISSN 1445-5226.
  15. ^ a b Chiao, C.-C.; Wu, W.-Y.; Chen, S.-H.; Yang, E.-C. (2009-06-26). "Visualization of the spatial and spectral signals of orb-weaving spiders, Nephila pilipes, through the eyes of a honeybee". Journal of Experimental Biology. 212 (14): 2269–2278. doi:10.1242/jeb.030734. ISSN 0022-0949.
  16. ^ a b c d e f g h Tso, I-Min; Chiang, Shu-Ya; Blackledge, Todd A. (2007-03-27). "Does the Giant Wood Spider Nephila pilipes Respond to Prey Variation by Altering Web or Silk Properties?". Ethology. 113 (4): 324–333. doi:10.1111/j.1439-0310.2007.01318.x. ISSN 0179-1613.
  17. ^ a b c d e Tso, I-M. (2005-03-15). "Giant wood spider Nephila pilipes alters silk protein in response to prey variation". Journal of Experimental Biology. 208 (6): 1053–1061. doi:10.1242/jeb.01437. ISSN 0022-0949.
  18. ^ a b Wang, Zuyuan; Cang, Yu; Kremer, Friedrich; Thomas, Edwin L.; Fytas, George (2020-01-14). "Determination of the Complete Elasticity of Nephila pilipes Spider Silk". Biomacromolecules. 21 (3): 1179–1185. doi:10.1021/acs.biomac.9b01607. ISSN 1525-7797.
  19. ^ a b c d Zhang, Shichang; Kuntner, Matjaž; Li, Daiqin (December 2011). "Mate binding: male adaptation to sexual conflict in the golden orb-web spider (Nephilidae: Nephila pilipes)". Animal Behaviour. 82 (6): 1299–1304. doi:10.1016/j.anbehav.2011.09.010. ISSN 0003-3472.
  20. ^ a b Wise, David H. (March 1981). "Inter-and intraspecific effects of density manipulations upon females of two orb-weaving spiders (araneae: araneidae)". Oecologia. 48 (2): 252–256. doi:10.1007/bf00347972. ISSN 0029-8549.
  21. ^ a b c d e f g Lee, Vanessa M. J.; Kuntner, Matjaž; Li, Daiqin (2015-01-21). "Ballooning behavior in the golden orbweb spider Nephila pilipes (Araneae: Nephilidae)". Frontiers in Ecology and Evolution. 3. doi:10.3389/fevo.2015.00002. ISSN 2296-701X.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  22. ^ VnExpress. "Giant wood spider, a specialty food in forests of south central Vietnam - VnExpress International".
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