Nephila

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Golden silk orb-weaver
Temporal range: Ypresian–present
Giant Golden Orb Weaver (Ventral Side).jpg
Ventral side of N. pilipes , Agumbe Rainforest, India
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Order: Araneae
Infraorder: Araneomorphae
Family: Nephilidae
Genus: Nephila
Leach, 1815 [1]
Type species
Aranea pilipes
(Fabricius, 1793) [1]
Species

See text

Diversity [1]
11 species

Nephila is a genus of araneomorph spiders noted for the impressive webs they weave. Nephila consists of numerous species found in warmer regions around the world, although some species formerly included in the genus have been moved to Trichonephila . They are commonly called golden silk orb-weavers, golden orb-weavers, giant wood spiders, or banana spiders.

Etymology

The genus name Nephila is derived from Ancient Greek, meaning "fond of spinning", [2] from the words νεῖν (nein) = to spin (related to nema νήμα "thread") + φίλος (philos) = "love".

Description

Nephila spiders vary from reddish to greenish yellow in color with distinctive whiteness on the cephalothorax and the beginning of the abdomen. Like many species of the superfamily Araneoidea, most of them have striped legs specialized for weaving (where their tips point inward, rather than outward as is the case with many wandering spiders). Their contrast of dark brown/black and green/yellow allows warning and repelling of potential predators to which their venom might be of little danger.

Golden orb-weavers reach sizes of 4.8–5.1 cm (1.5–2 in) in females, not including legspan, with males being usually two-thirds smaller (less than 2.5 cm, 1 in). In 2012, a large individual was photographed killing and consuming a 0.5-m-long brown tree snake in Freshwater, Queensland. [3] Species from Taiwan have been known to reach over 130 mm (5.1 in), legspan included, in mountainous country. [4] In 2014, a study discovered that golden orb-weavers living in urban areas, particularly areas of a high socioeconomic status, grew larger and carried more eggs than those in their native habitats. A number of possible explanations were suggested, such as increased food supplies due to artificial light or lack of predators and parasites. [5]

Species

In 2018, twelve Nephila species were reclassified as Trichonephila , [6] with another two (N. kuhlii and N. robusta) considered in 2020 to be junior synonyms of N. pilipes. [7] Species whose placement has been changed by some sources include:

As of April 2024, the World Spider Catalog does not accept all of these changes, listing the following species in the genus Nephila: [1]

Additional fossil species are known from the Cenozoic. In 2012 Geratonephila burmanica was described from the Cenomanian aged Burmese amber, Wunderlich 2015 synonymised Geratonephilia with Nephilia tenuis, a species from the Dominican Amber, as he considered it unlikely that the amber was actually Burmese in origin. [9] Though largely ambiguous, the origins of Nephila are undoubtedly Gondowanan. With the prime candidates being Africa, Indomalaya, and Australasia. [10]

Distribution and habitat

Golden silk orb-weavers are widespread in warmer regions throughout the world, with species in Australia, Asia, Africa (including Madagascar), and the Americas. Spiderlings can be carried by the wind over long distances, and each year, a small number of female golden orb web spiders are found in New Zealand (where they are not endemic) after having been blown across the Tasman Sea; the spiders usually end up in the North Island. [11]

Whilst the geographic distribution of Nephila is large, many habitat similarities are seen between these locations. A warm and reasonably wet climate is generally preferred, as these are some of the environmental cues that induce spiderling hatching. [12] Locally, spiders look for relatively dense vegetation where webs can be set up in areas that insects will regularly fly through. Urban environments are also attractive due to the large prey concentrations and lower levels of predation. [13]

Behavior

Web spinning and structure

Nephila spiders produce large asymmetric orb webs up to 1.5 m (5 ft) in diameter.[ citation needed ]Nephila species remain in their webs permanently, so have a higher predation risk. The golden silk orb-weaver is named for the yellow color of the spider silk used to construct these webs.

Yellow threads of their web shine like gold in sunlight. Carotenoids are the main contributors to this yellow color, but xanthurenic acid, two quinones, and an unknown compound may also aid in the color. [14] Experimental evidence suggests that the silk's color may serve a dual purpose: sunlit webs ensnare bees that are attracted to the bright yellow strands, whereas in shady spots, the yellow blends in with background foliage to act as a camouflage. The spider is able to adjust pigment intensity relative to background light levels and color; the range of spectral reflectance is specifically adapted to insect vision.

The webs of most Nephila spiders are complex, with a fine-meshed orb suspended in a maze of non-sticky barrier webs. As with many weavers of sticky spirals, the orb is renewed regularly if not daily, apparently because the stickiness of the orb declines with age. When weather is good (and no rain has damaged the orb web), subadults and adults often rebuild only a portion of the web. The spider removes and consumes the portion to be replaced, builds new radial elements, then spins the new spirals. This partial orb renewal is distinct from other orb-weaving spiders that usually replace the entire orb web. The web of Nephila antipodiana contains ant-repellent chemicals to protect the web. [15]

Typically, the golden orb-weaver first weaves a nonsticky spiral with space for two to 20 more spirals in between (the density of sticky spiral strands decreases with increasing spider size).[ citation needed ] When she has completed the coarse weaving, she returns and fills in the gaps. Whereas most orb-weaving spiders remove the nonsticky spiral when spinning the sticky spiral, Nephila spiders leave it. This produces a "manuscript paper" effect when the orb is seen in the sun: groups of sticky spirals reflecting light with "gaps" where the nonsticky spiral does not reflect the light.

In relation to the ground, the webs of adults may be woven from eye-level upwards high into the tree canopy. The orb web is usually truncated by a top horizontal support strand, giving it an incomplete look.

Adjacent to one face of the main orb, a rather extensive and haphazard-looking network of guard-strands may be suspended a few centimeters distant across a free space. This network is often decorated with a lumpy string or two of plant detritus and insect carcasses clumped with silk. This "barrier web" may function as a kind of early-warning system for incoming prey or against spider-hunting predators, or as a shield against windblown leaves; it may also be remnants of the owner's previous web. At least one reference explains the suspended debris-chain as a cue for birds to avoid blundering into and destroying the web.

Prey capture and feeding

The golden silk orb-weaver targets many different organisms as prey, ranging from small flies and beetles to larger cicadas and locusts. As a result of their strong web structure, small birds and bats can also become trapped and fed upon. Whilst most of the captured prey is relatively small compared to Nephila, the majority of biomass consumed comes from larger, rarer prey. Prey larger than 66% of the captor's size accounts for just 16.5% of prey captured, but 85% of prey consumed, indicating the spider is selective in its feeding habits. [16]

Spiders are notified that potential prey has been caught in the web through vibrations along strands, and these can be followed to the prey location on the web and be used to estimate prey size. [13]

Nephila species also create caches of food for storage, which can be found above the hub of the web and contain up to 15 prey items. These items are arranged in a line vertically and are wrapped in silk to reduce dehydration. Caches are created and grow when prey is readily available and more biomass is available for consumption than is required by the spider. The purpose of caches is to have a backup food source when prey is scarce and occasionally to provide bait to attract more prey to the web. [17] Nephila species may also respond to food shortages by moving their webs, but this is a response to longer periods of prey scarcity than cache creation. [18] Web moving is seen as a result of environmental change, whereas caches occur from environmental fluctuation.

Mating and reproduction

N. pilipes female with many males Giant Wood Spider (Nephila pilipes) W IMG 2806.jpg
N. pilipes female with many males

Nephila spiders display large sexual dimorphism in size, with females being greatly larger than males. Debate exists as to whether this is a result of male dwarfism or female gigantism.[ citation needed ] Smaller males may be selected for due to the presence of competition for mating. Smaller males are quicker and more nimble, allowing them to be able to catch the females more easily, as well as to escape when threatened. Larger males may have to wait for the female to come close due to their slower speed. [19] Larger females may have been selected for as a result of males using mating plugs upon copulation. Larger individuals reduce the success of these plugs, allowing for multiple mating and reducing the risk of genital mutilation. [20] Gigantism in females is also associated with fecundity, as larger individuals can produce more eggs and therefore increase reproductive success. [21]

When males are fully mature, they leave their webs to search for a suitable female, often using web characteristics to identify potential mates. Often, multiple males attempt to court the same female, thus competition for territory on the web occurs, but is rarely physical, as smaller males surrender area to larger ones. When males approach females, they are often feeding, allowing the males to get closer without an aggressive response and also meaning the female is not moving. On approach, the male makes himself known by tapping on a web strand to ensure the female is amenable before proceeding to mate. When met with aggression, males stop approaching and remain in the same location until the female relaxes or they retreat. Females engage in multiple mating, but no benefit to the offspring occurs as a result of this; however, the energy cost of repelling a male is higher than that of allowing him to copulate. [22] As a result of this, sperm competition occurs through males altering the duration and frequency of mating, with longer mating being proportional to a greater likelihood of success. Sexual cannibalism is uncommon in Nephila as a result of male mating behaviours. By copulating when females are immobile after molting or inactive due to feeding, the males increase their chances of survival. Males also approach from the side of the web opposite the female, increasing the odds of a successful approach. [23] Male Nephila pilipes is reported to have a mate binding behavior to avoid sexual cannibalism. [13] Sexual cannibalism does still occur, but generally is more common with larger males, and from older females.

Females produce an egg sac in the surrounding environs of the web to protect their eggs. The eggs are deposited on a silk platform, then are covered in loose silk to form a sac, which is firmly attached to surrounding vegetation so that it is hidden from the view of predators. It is reported that egg sacs are mostly under leaves and other coverings. However, only Nephila pilipes is different than other Nephila species. They lay eggs in small pits on the ground to avoid parasitism. [13] These sacs can contain from 300 to 3000 eggs, depending on mating success and particular species. [18] Once hatched, the spiderlings inhabit a communal web to begin their lives.

Thermoregulation

Nephila spiders change their body positioning relative to the sun to maintain internal temperatures at an optimal level. As ambient temperatures increase, the spiders position themselves so the abdomen shades the cephalothorax from the sun. Spiders may also hang from their hind legs as a result of the heat due to a loss of hydrostatic pressure. Conversely, as temperatures cool down, the spiders position themselves perpendicular to the sun to retain as much heat energy as possible. When ambient temperatures reach extreme highs (above 40 °C), they may leave their webs and seek shade in the surrounding environment. [24]

Predation and parasitism

Predation of Nephila species is relatively uncommon; when it does occur, the main group affected are the juvenile individuals. The major predators are birds, but wasps and damselflies also prey upon smaller juveniles.

Nephila species are frequently parasitized by Argyrodes , a genus of very small black-and-silver spiders that are kleptoparasitic. As many as a few dozen may infest a single Nephila web to feed from the host spider's captured prey. The frequent rebuilding or abandoning of webs by Nephila may be a tactic for controlling Argyrodes. Spiny orb-weaver spiders of the genus Gasteracantha also inhabit the webs of Nephila as kleptoparasites.

Egg sacs generally remain free from both predation and parasitism, often due to the close proximity of the mother and how well it is hidden.

Life cycle

Spiderlings

Nephila spiderlings leave the egg sac as a result of environmental cues, often warmer and wetter conditions in spring. They then live on a communal web, eating dead siblings and web debris for around a week before dispersing to make individual webs. [12]

Young spiders do not generally build yellow-colored silk, and the young themselves can be easily mistaken for young orchard spiders ( Leucauge ) in general color and shape (both genera sport silver stripes or patches on their abdomens, described in some references as a form of heat control). The best distinction between Leucauge and Nephila juveniles is web structure: Leucauge species tend to build horizontal orbs that form a perfect circle, whereas Nephila species build vertical, elliptical orbs that are incomplete (missing the portion of the orb over the hub, the center where the spider sits). The latter seem to prefer more open habitat such as second-growth scrub or forest edges. Fences or building overhangs often do just as nicely.

Molting and growth

Once they are juvenile, Nephila spiders inhabit their individual webs, then begin their growth by the molting process. The time between molts is called an instar and seven to 12 of these can occur depending on food availability. Ecdysis, the shedding of the exoskeleton, occurs through the formation of a soft exoskeleton inside the current one. Once the old exoskeleton is shed, the new, larger one begins to harden. Ecdysis occurs when the spider's mass becomes too great for the current exoskeleton to support. [25] Male spiders seek out females for copulation and live on their webs. When mating season arrives, both males and females stop molting and remain the same size for the remainder of their lives.

Venom

The venom of the golden silk orb-weaver is effective in action on prey, but has not been reported to be of any notable consequence for humans if accidentally bitten. In the literature, Nephila is one of several genera where the venom "must be considered as more or less ineffectual in human beings". [26] That said, the potentially large size of several members of the genus means that they possess relatively strong chelicerae, so any bite can cause some mechanical damage, but only of short-term localised effect for humans. However, further studies of the venom components are needed to better understand pathways associated with any toxicity.

Interaction with humans

Nephila do not seem to form either beneficial or harmful relationships with humans. Females often construct their webs using human structures as a base for support strands due to their stability. Individuals are often found in urban and suburban environments due to the protection from predation and greater prey availability. [27] As they weave their webs in bushes and near flowers, they might present a nuisance for gardeners or flower pickers. Some nests near fruits may repel or destroy known pests, such as Tephritid fruit flies, without the need to use insecticides. Spiders may bite humans if provoked but more often flee if confronted.

Golden silk

Cape made from Madagascar golden orb-weaver spider silk, Victoria and Albert Museum, London Spider silk cape.jpg
Cape made from Madagascar golden orb-weaver spider silk, Victoria and Albert Museum, London

There have been several efforts in the past to produce garments from Nephila silk although none commercially viable. [29] These include two bed hangings that were shown at the 1900 Paris Exhibition. [30] In 2004 a textile designer, Simon Peers, and an entrepreneur, Nicholas Godley managed in three years' work and using 1.2 million Golden silk orb-weavers (collected in the wild and released some 30 minutes later after they produced the silk) to produce a shawl that was exhibited at the American Museum of Natural History in 2009. [31] [32] By 2012 they managed to produce a second, bigger garment, a cape, that, together with the shawl, were exhibited at the Victoria and Albert Museum in London. [33] Two shawls and a traditional Madagascan lamba made of this spider silk were included in an exhibition of curios from the natural world in 2021. [34]

Another possible use of Nephila silk lies in tissue engineering. A study from the Medizinische Hochschule Hannover reports that processed Nephila silk is an excellent scaffold material thanks to its biocompatibility, mechanical strengths, and its property to promote cell adhesion and proliferation.[ clarification needed ] [35] In particular, the silk acts as a suitable guiding material for peripheral nerve regrowth. [36]

Fishermen on the coasts of the Indo-Pacific ocean remove Nephila webs and form them into a ball, which is thrown into the water. There it unfolds and is used to catch bait fish. [29]

Related Research Articles

<span class="mw-page-title-main">Orb-weaver spider</span> Family of spiders

Orb-weaver spiders are members of the spider family Araneidae. They are the most common group of builders of spiral wheel-shaped webs often found in gardens, fields, and forests. The English word "orb" can mean "circular", hence the English name of the group. Araneids have eight similar eyes, hairy or spiny legs, and no stridulating organs.

<i>Trichonephila inaurata</i> Species of spider

Trichonephila inaurata, synonym Nephila inaurata, commonly known as the red-legged golden orb-weaver spider or red-legged nephila, is a species of spider of the genus Trichonephila. It is native to southern and East Africa, as well as several islands of the western Indian Ocean.

<i>Trichonephila clavata</i> Species of spider

Trichonephila clavata, also known as the Joro spider (ジョロウグモ , is a member of the Trichonephila genus. The spider can be found throughout Japan, Korea, Taiwan, China. Due to its large size and the bright, unique colors of the female Trichonephila, the spider is well-favored in Japan.

<i>Trichonephila clavipes</i> Species of spider native to the Americas

Trichonephila clavipes, commonly known as the golden silk orb-weaver, golden silk spider, or colloquially banana spider, is an orb-weaving spider species which inhabits forests and wooded areas ranging from the southern US to Argentina. It is indigenous to both continental North and South America. Known for the golden color of their silk, the large size of their females, and their distinctive red-brown and yellow coloring, T. clavipes construct large, asymmetrical circular webs attached to trees and low shrubs in woods to catch small- and medium-size flying prey, mostly insects. They are excellent web-builders, producing and utilizing seven different types of silk, and they subdue their prey by injecting them with venom, as opposed to related species which immobilize their prey by wrapping them in silk first. They are not known to be aggressive towards humans, only biting out of self-defense if touched, and their relatively harmless venom has a low toxicity, posing little health concern to healthy human adults. Due to their prevalence in forests, T. clavipes may be encountered by hikers.

<i>Zygiella x-notata</i> Species of spider

Zygiella x-notata, sometimes known as the missing sector orb weaver or the silver-sided sector spider, is a spider species in the family Araneidae. They are solitary spiders, residing in daily spun orb webs. Z. x-notata is a member of the genus Zygiella, the orb-weaving spiders. The adult female is easily recognized by the characteristic leaf-like mark on her posterior opisthosoma, caudal to the yellow-brown cephalothorax.

<i>Trichonephila edulis</i> Species of spider

Trichonephila edulis is a species of large spider of the family Nephilidae, formerly placed in the genus Nephila. It is referred to by the common name Australian golden orb weaver. It is found in Indonesia from Java eastwards, Papua New Guinea, Australia, northern New Zealand, and New Caledonia.

<i>Nephila pilipes</i> Species of spider

Nephila pilipes 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, 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.

Nephilengys is a genus of tropical spiders of the family Nephilidae, consisting of two currently described species. The genus Nephilingis has been split off from this genus. Both genera have been called hermit spiders from the habit staying in their retreats during the day; the name eunuch spiders has been used for Nephilengys alone. Males may sever parts of their palpal bulbs after copulation.

<span class="mw-page-title-main">Sexual cannibalism</span> Practice of animals eating their own mating partners

Sexual cannibalism is when an animal, usually the female, cannibalizes its mate prior to, during, or after copulation. It is a trait observed in many arachnid orders, several insect and crustacean clades, gastropods, and some snake species. Several hypotheses to explain this seemingly paradoxical behavior have been proposed. The adaptive foraging hypothesis, aggressive spillover hypothesis and mistaken identity hypothesis are among the proposed hypotheses to explain how sexual cannibalism evolved. This behavior is believed to have evolved as a manifestation of sexual conflict, occurring when the reproductive interests of males and females differ. In many species that exhibit sexual cannibalism, the female consumes the male upon detection. Females of cannibalistic species are generally hostile and unwilling to mate; thus many males of these species have developed adaptive behaviors to counteract female aggression.

<i>Trichonephila plumipes</i> Species of spider

Trichonephila plumipes, the Pacific golden orb weaver, is a species of spider found in Australia, Indonesia and some Pacific Islands, which exhibits extreme sexual dimorphism through its sexual cannibalism behavior. It is sometimes called the tiger spider due to its markings which look similar to a tiger. This species was formerly called Nephila plumipes. As with other spiders from the genus Nephila, these spiders have a distinct golden web.

<span class="mw-page-title-main">Spider web</span> Structure created by a spider from silk

A spider web, spiderweb, spider's web, or cobweb is a structure created by a spider out of proteinaceous spider silk extruded from its spinnerets, generally meant to catch its prey.

<i>Cyrtophora citricola</i> Species of spider

Cyrtophora citricola, also known as the tropical tent-web spider, is an orb-weaver spider in the family Araneidae. It is found in Asia, Africa, Australia, Costa Rica, Hispaniola, Colombia, and Southern Europe and in 2000, it was discovered in Florida. C. citricola differs from many of its close relatives due its ability to live in a wide variety of environments. In North America and South America, the spider has caused extensive damage to agricultural operations.

<span class="mw-page-title-main">Darwin's bark spider</span> Species of spider

Darwin's bark spider is an orb-weaver spider that produces the largest known orb webs, ranging from 900 to 28,000 square centimetres, with bridge lines spanning up to 25 metres (82 ft). The spider was discovered in Madagascar in the Andasibe-Mantadia National Park in 2009. Its silk is the toughest biological material ever studied. Its tensile strength is 1.6 GPa. The species was named in honour of the naturalist Charles Darwin on November 24, 2009—precisely 150 years after the publication of The Origin of Species.

<i>Socca pustulosa</i> Species of arachnid

Socca pustulosa is a Orb-weaver spider species in the family Araneidae, and it was first described by a French scientist Charles Athanase Walckenaer in 1841 from Tasmania, Australia, but later on when Walckenaer examined the specimen collected from New Zealand and renamed it as a different species- Epeira verrucosa. Until 1917, Dalmas reviewed the Australian pustulosa and New Zealand verrucosa and realized they were the same species. Although S. pustulosa has been accepted for some time in the genus of Eriophora, the evidence supporting its placement within this genus were not fully convincing. Therefore, an alternative view was proposed in 2022- a new genus established to accommodate pustulosa along with 11 other spider species from Australia; the diagnostic test based on haplotype analysis and systematic morphology study by arachnologists and found the anatomical features of male pedipalp terminal apophysis differs from other orb-web species.

<i>Cyclosa argenteoalba</i> Species of spider

Cyclosa argenteoalba, in the trashline orbweavers genus, is a species of orb weaver in the spider family Araneidae. It is found in East Asia in the countries of China, Japan, and Korea. C. argenteoalba are diurnal, which means they are active during the day. Each individual has a unique appearance due to their differences in the ratio of black to silver coloring on their abdomen. Spiders with less silver coloring are better at catching prey, since the silver is bright and warns their prey. They catch their prey by waiting in the hub of their web until their prey is close enough to catch. Parasitic larvae are often found attached to C. argenteoalba, and the larvae are able to manipulate the spider's behavior. Females are on average 2 mm longer in size than males. During mating, female genital mutilation is common in order to increase the fitness of the male. On their webs, they often attach silk "decorations" that are thought to deter predators. Relocating to a different place to build a new web occurs frequently until they find a location with a significant amount of prey.

<i>Nephilingis</i> Genus of spiders

Nephilingis is a genus of spiders in the family Nephilidae. It was split off from the genus Nephilengys in 2006. Both genera have been called hermit spiders from the habit of staying in their retreats during the day; alternatively the name "hermit spider" may be reserved for Nephilingis, with Nephilengys species called "eunuch spiders".

<span class="mw-page-title-main">Nephilidae</span> Spider family

Nephilidae is a spider family commonly referred to as golden orb-weavers. The various genera in Nephilidae were formerly placed in Tetragnathidae and Araneidae. All nephilid genera partially renew their webs.

<i>Tetragnatha versicolor</i> Species of spider

Tetragnatha versicolor is a species of long-jawed orb weaver in the spider family Tetragnathidae. It is found throughout North America, Canada, Central America, and Cuba, but are most common in the United States. T. versicolor is heavily concentrated in New England and the west coast in states like California and Washington. T. versicolor is considered a habitat generalist, and can thrive in many different environments. While they can be found in places like Grasslands, Wetlands, Forests, etc., they prefer dryer areas like normal trees and shrubs. Unlike other spiders in the genus Tetragnatha, T. versicolor will rarely reside near aquatic environments. T. versicolor will typically be colored dark yellow or pale orange and average around 5 mm for males and 6.5 mm for females in length, which is very small for a spider. They are much longer than they are wide, making them very distinct. In addition, T. versicolor can be distinguished from other spiders in Tetragnatha by the distinct separation of the anterior/posterior eyes and the appearance of their reproductive organs. As an orb weaver spider, T. versicolor creates a web to hunt for prey. It will wait at night for prey to stumble into its web and use vibrational signals throughout the web to sense trapped prey. In terms of mating behavior, T. versicolor lacks a distinct courting ritual and will mate with any others in the proximity. Mating behavior is heavily affected by female mating history. In terms of interactions with humans, the bite of T. versicolor is venomous, but not known to cause significant harm.

<i>Trichonephila</i> Genus of spiders

Trichonephila is a genus of golden orb-weaver spiders that was first described by Friedrich Dahl in 1911, as a subgenus of Nephila. Trichonephila was elevated to the level of genus by Kuntner et al. in 2019. The genus Trichonephila belongs to the Nephilidae family.

<i>Leucauge mariana</i> Species of spider

Leucauge mariana is a long-jawed orb weaver spider, native to Central America and South America. Its web building and sexual behavior have been studied extensively. Males perform several kinds of courtship behavior to induce females to copulate and to use their sperm.

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