Bolas spider

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Bolas spiders
Bolas Spider - Mastophora phrynosoma hunting, Julie Metz Wetlands, Woodbridge, Virginia (24843011518).jpg
Mastophora phrynosoma with bolas, Virginia, US
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Order: Araneae
Infraorder: Araneomorphae
Family: Araneidae
Subfamily: Cyrtarachninae s.l.
Informal group: Bolas spiders
Genera [1] [2] [3]

A bolas spider is a member of the orb-weaver spider (family Araneidae) that, instead of spinning a typical orb web, hunts by using one or more sticky "capture blobs" on the end of a silk line, known as a "bolas". By swinging the bolas at flying male moths or moth flies nearby, the spider may snag its prey rather like a fisherman snagging a fish on a hook. Because of this, they are also called angling or fishing spiders [4] (although the remotely related genus Dolomedes is also called a fishing spider). The prey is lured to the spider by the production of up to three sex pheromone-analogues.

Contents

Bolas spiders have been treated as either the whole or part of either the tribe "Mastophoreae" or Mastophorini, the subfamily Mastophorinae, or the informal group mastophorines. Recent studies show that the genus Celaenia , which does not use a bolas, belongs in the same taxonomic group.

Description

Mastophora bisaccata resembles a snail when at rest Bolas Spider - Mastophora bisaccata, Okaloacoochee Slough WMA, Florida crop.jpg
Mastophora bisaccata resembles a snail when at rest

Bolas spiders are small nocturnal animals with conspicuous outgrowths on the upper (dorsal) surface of the cephalothorax. Mature females have a body length of 10–20 mm (1⁄2–3⁄4 in), while the males, which are much smaller, have a body length usually less than 2 mm (1⁄16 in). The abdomen of adult females is large relative to the cephalothorax, partially covering it. In some species, the female's abdomen is comparatively smooth, while in others it has humps and tubercules. The females of some bolas spiders (e.g. Mastophora cornigera ) look remarkably like a bird dropping, thanks to their large, globular abdomen and brownish cephalothorax. This is a form of defensive mimicry as the animals that prey on spiders pay little attention to bird droppings, which enables the spiders to rest unnoticed during the day in fairly exposed places. M. bisaccata resembles the shell of a snail that is abundant in the spider's habitat. While the bird dropping mimics rest on the upper surface of leaves, M. bisaccata rests on the undersides. If removed from the surface, Mastophora females produce a pungent odor, which is highly unusual for spiders (other than bolas spiders, only one species of Cyrtarachne is known to do this). They do not attempt to flee when handled. Some species of Cladomelea also rest exposed on leaves during the day. However, the Australian Ordgarius magnificus , which displays eye spots on its back that make it resemble the moth it hunts, ties leaves together with silk to form a retreat. [5] [6]

Taxonomy

Genera

Bolas-using spiders are species of four related genera. [1] [2] [3] (Distributions are from the World Spider Catalog. [7] )

The use of a bolas by the Madagascan genus Exechocentrus was not reported until 2012, [3] so the genus is not included in earlier accounts of bolas spiders.

Classification

Bolas spiders are members of the family Araneidae, the orb-weaver spiders, although they do not weave webs. They may be treated as a group within the subfamily Cyrtarachninae sensu lato, [8] or as the whole or part of a separate taxon. In 1931, Mello-Leitão created a tribe under the name Mastophoreae, which included genera now known to use a bolas as well the genus Acantharachne . [9] Gertsch in 1955 also placed bolas spiders in the tribe Mastophoreae, again including Acantharachne (under the synonym Acantharanea). [10] Article 29.2 of the International Code of Zoological Nomenclature states that the ending "-ini" should be used for tribes, so the correct name for the tribe is now "Mastophorini". [11] The tribe has been elevated to a separate subfamily Mastophorinae. [12] The informal name "mastophorines" has also been used, including both bolas-using spiders and a related genus. [13]

A 2014 molecular phylogenetic study of a representative set of genera placed in Cyrtarachninae s.l. showed that the species of the two bolas spider genera included in the study formed a strongly supported clade nested within Cyrtarachninae s.l. [8] A wider study of the family Araneidae published in 2020 reached a similar conclusion, separating Cyrtarachninae s.l. into two informal sister groups "mastophorines" and "cyrtarachines". Mastophorines included the genus Celaenia that does not use a bolas. [13] Bolas-using spiders thus form a subgroup within the narrowly defined mastophorines or the broadly defined Cyrtarachninae.

Phylogeny and web evolution

The Cyrtarachninae, defined broadly, construct several kinds of web, which differ from the orb webs of other Araneidae. Some construct "spanning-thread webs", which have a small number of radii and widely spaced sticky threads that do not form spirals. Others construct triangular webs. These are formed in the same way as spanning-thread webs, but with only three radii, so that they appear triangular. Bolas spiders do not spin webs at all, adult females catching their prey on single sticky threads. Finally, some species capture their prey without a web, using their outstretched legs, as do juvenile and male bolas spiders. [8]

One hypothesis was that the evolution of the web types involved successive reduction: spanning-thread webs → triangular webs → bolas → no web. However, the 2014 molecular phylogenetic study by Tanikawa et al. showed that this hypothesis did not fit with the evolutionary relationships they derived. Their cladogram is shown below. [8] (The dashed line shows where the genus Exechocentrus would be expected to fall based on a 2020 study. [13] ) Triangular webs could have been derived from spanning-thread webs, but spiders with no webs or that used bolas formed a completely separate monophyletic group. [8]

Cyrtarachninae s.l.

Scharff et al. in a molecular phylogenetic study of the family Araneidae published in 2020 reached a similar conclusion concerning the relationships within the broadly defined Cyrtarachninae. They included the bolas-using genus Exechocentrus in their analysis (which has been added to the cladogram above), dividing Cyrtarachninae s.l. into "mastophorines" and "cyrtarachnines". [13]

Distribution

Bolas spiders are found in America, Africa including Madagascar, and Australasia northwards into Asia. They do not occur in temperate Eurasia. About half of the known Mastophora species occur in South America. The genus is distributed from southern Chile to the northern US (to 45° north latitude in Minnesota). M. archeri, M. bisaccata, M. hutchinsoni and M. phrynosoma occur widely in the US east of the Great Plains. M. cornigera occurs from Alabama to California, as well as in northern Mexico.

Life-cycle

Mastophora phrynosoma resting in full view, resembling a bird dropping; egg sac under the leaf Arachtober 23 -2 - Bolas Spider - Mastophora phrynosoma and egg sac, Julie Metz Wetlands, Woodbridge, Virginia (37840647492).jpg
Mastophora phrynosoma resting in full view, resembling a bird dropping; egg sac under the leaf

The life-cycle of the North American Mastophora hutchinsoni is best known. As is probably the case for other Mastophora species in temperate regions, mating takes place in late summer or early fall, after which egg cases are produced. Adults do not survive the winter, males typically dying before females. A similar pattern is observed for bolas spiders in the Southern Hemisphere, depending on the climate. In the subtropical climate of Edmonton in north Queensland, Australia, a female Ordgarius monstrosus matured in June and produced egg cases between late June and early September. In Pietermaritzburg, South Africa, adult females of Cladomelea akermani were observed with egg cases during July. The number of egg cases varies widely among species, with temperate species producing fewer than tropical ones (e.g. one to three for Mastophora species in temperate regions of the United States, and up to 11 for M. dizzydeani near the equator). The number of eggs per case also varies depending on the climate, so that the total number of spiderlings produced by one female ranges from several hundred in the temperate US to several thousand in Colombia. [6]

Some male bolas spiders, such as those of Mastophora cornigera and Ordgarius magnificus , appear to be mature on emergence from their egg cases and so capable of inseminating females. Others, such as those of M. dizzydeani, M. hutchinsoni and M. phrynosoma require one or two moults before becoming mature. By contrast, Mastophora females may require up to eight moults to reach maturity. [6]

Prey capture

Cladomelea sp. from South Africa with multiple drops on its bolas line Bola Spider (Cladomelea sp.) (Photo Sylvain REMY) (9286584444).jpg
Cladomelea sp. from South Africa with multiple drops on its bolas line

Bolas spiders are defined by the way in which adults catch their prey. Like other members of the subfamily Cyrtarachninae sensu lato, bolas spiders specialize in catching moths. The scales covering moth wings are detachable, so that most ordinary orb webs are not effective at capturing moths. Adult female bolas spiders spend the day resting, disguised in various ways, including as bird droppings. At night, they feed on male moths which they attract by producing a scent that mimics the sex pheromones emitted by the female of one or more species of moth. They capture approaching male moths by using a "bolas", a silken thread with one or more sticky drops at the end which they swing, rather than throw, at the moth. [2] [6] Other spiders in the subfamily Cyrtarachninae, including Celaenia species from Australia and Taczanowskia species from South America, also use chemical lures to attract moths, but they catch them with their front legs. [6]

Bolas spiders will try and often succeed in catching any insect that is flying nearby. They seem able to detect prey by the sound of their approaching flight. In experiments, M. cornigera attacked a hand-held moth whose wings were fluttering, but did not attack if the moth's wings were immobilized. [6]

Left: Mastophora cornigera attracting a moth while holding its bolas; right: with captured moth Mastophora et sa proie 2.jpg
Left: Mastophora cornigera attracting a moth while holding its bolas; right: with captured moth

The manner in which the bolas is used differs among the genera. Mastophora species hold the bolas stationary with a front leg until a moth approaches, and then cock the leg and swing the bolas towards the prey with a rapid pendulum-like stroke. Other genera use their second leg. Ordgarius species begin to whirl the bolas rapidly when detecting an incoming moth. Cladomelea akermani whirls the bolas immediately after it has been prepared for about 15 minutes, even when there is no moth present. [6] Instead of preparing a single bolas, at least four Mastophora species are known to sometimes hang up to nine droplets on one horizontal line. This has not been observed to result in prey capture, but is remarkably similar to the midline of a Pasilobus web in which the spanning threads have been broken. [6]

Spiderlings of both sexes and adult males hunt without a bolas. This might be because the small globule these spiders could produce would dry out rapidly. Rather, they position themselves at edges of leaves and grab prey with their frontal pair of legs. Juvenile females and males of all ages of M. hutchinsoni have a row of strong bristles on the prolateral surface of the tibiae and metatarsi of each of their first two pairs of legs which may aid in prey capture. Females lose these bristles after a few moults. Moth flies of the genus Psychoda are prominent prey of juvenile and adult male Mastophora. [14] [15]

Drop structure

The sticky drops of the bolas have a diameter of about 2.5 mm (1⁄10 in). Their internal structure is complex, consisting of a mass of curled or folded fibre embedded in a viscid matrix which is in turn surrounded by a less viscous layer. This results in the low viscosity liquid flowing past the moth's scales to reach the cuticle below, while the more viscous liquid forms a bond to the thread to sustain the moth's weight. The folded thread inside the ball permits elastic elongations which extend the spider's striking range. Ordgarius sometimes has smaller droplets above the terminal one. [6] Exechocentrus lancearius uses a bolas with two globules. [3] The other genera usually produce only one terminal globule. The drop shrinks over time, thus presumably reducing its effectiveness in catching prey. Bolas spiders consume unused bolas after at most half an hour if the hunt was fruitless. [6]

Prey variety

Female Mastophora catch an average of two moths per night, but as many as six or seven catches during a single night have been observed. Bolas spiders vary in the range of moth species they capture, with 20 moth species recorded for Mastophora cornigera females, mostly noctuids, and four for Mastophora hutchinsoni females, three of which were noctuids. [6] A study of M. hutchinsoni showed that the blend of chemicals produced by the spider was very similar to that produced by the female of the bristly cutworm, Lacinipolia renigera , whose males make up about two thirds of the biomass consumed by the spider. [16] A study by K.F. Haynes showed that M. hutchinsoni females adapt to changes in prey availability, with one major prey species flying from early evening until 10:30 pm or so and the other only after 11:00 pm. The first moth ignores the pheromone of the second, so the spider produces a mix of both pheromones early in the evening. However, since the second moth is repulsed by the other moth's pheromone, the spider ceases to produce the first pheromone later at night. [17] [16]

Natural enemies

Yeargan noted that records of natural enemies of bolas spiders all concern predators or parasitoids of the egg stage. A gryllacridid was observed feeding on the eggs of Ordgarius magnificus , and a flesh fly parasitoid has been reared from egg cases of the same species. Five species of hymenopteran parasitoids are known to attack Mastophora eggs. Tromatobia notator parasitizes M. cornigera, M. bisaccata and M. phrynosoma; a Gelis species parasitizes M. cornigera. Eupelmids in the genus Arachnophaga also parasitize Mastophora species. [6]

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>Ordgarius</i> Genus of spiders

Ordgarius is a genus of orb-weaver spiders first described by Eugen von Keyserling in 1886. Adult females of the genus are bolas spiders, capturing their prey with one or more sticky drops at the end of a single line of silk rather than in a web. Males and juvenile females capture their prey directly with their legs.

<i>Kaira</i> (spider) Genus of spiders

Kaira, sometimes called frilled orbweavers, is a mostly neotropical genus of orb-weaver spiders first described by O. Pickard-Cambridge in 1889. It includes sixteen described species that occur from South America up to the southern and eastern USA. It is presumably related to Aculepeira, Amazonepeira and Metepeira.

<i>Mastophora</i> (spider) Genus of spiders

Mastophora, also known as bolas spiders, is a genus of orb-weaver spiders first described by E. L. Holmberg in 1876. They can be identified by a pair of lumps on the dorsal surface of the opisthosoma, though not all males will have these lumps.

Exechocentrus is a genus of Madagascan orb-weaver spiders first described by Eugène Simon in 1889. It is a bolas-using spider, capturing its prey with one or more sticky drops at the end of a single line of silk rather than in a web.

<i>Pasilobus</i> Genus of spiders

Pasilobus is a genus of orb-weaver spiders first described by Eugène Simon in 1895.

Taczanowskia is a genus of orb-weaver spiders first described by Eugen von Keyserling in 1879. Contrary to the common name of the group, spiders of the genus Taczanowskia do not build webs and are furtive hunters, deceiving their prey by producing sex pheromones that attract male moths, and catching their prey by using a pair of enlarged claws at the tip of their anterior legs.

<i>Mastophora cornigera</i> Species of spider

Mastophora cornigera is a species of orb weaver in the spider family Araneidae. It is found in a range from the United States to Nicaragua. Like all known species of the genus Mastophora, adult females are bolas spiders, capturing their prey with one or more sticky drops at the end of a single line of silk rather than in a web. Males and juvenile females capture their prey directly with their legs.

Mastophora leucabulba is a species of orb weaver in the spider family Araneidae. It is found in a range from the United States to Honduras. Like all known species of the genus Mastophora, adult females are bolas spiders, capturing their prey with one or more sticky drops at the end of a single line of silk rather than in a web. Males and juvenile females capture their prey directly with their legs.

Mastophora yeargani is a species of orb weaver in the spider family Araneidae. It is found in the United States. Like all known species of the genus Mastophora, adult females are bolas spiders, capturing their prey with one or more sticky drops at the end of a single line of silk rather than in a web. Males and juvenile females capture their prey directly with their legs.

<i>Mastophora phrynosoma</i> Species of spider

Mastophora phrynosoma is a species of orb weaver in the spider family Araneidae. It is found in the United States. Like all known species of the genus Mastophora, adult females are bolas spiders, capturing their prey with one or more sticky drops at the end of a single line of silk rather than in a web. Males and juvenile females capture their prey directly with their legs.

<i>Mastophora hutchinsoni</i> Species of spider

Mastophora hutchinsoni, also known as the American bolas spider, is a species of orb weaver in the genus Mastophora. The genus is distributed extensively throughout various subtropical geographical areas including Australia, South Africa, Oriental Asia, and the Americas and is not found in Europe. The hunting behavior of adult female M. hutchinsoni is unusual because they are bolas spiders. They mimic moth pheromones to attract male moths, and female M. hutchinsoni have evolved to alter their chemical release to target different moths. They then capture their prey with a sticky drop on the end of a silk line, resembling a bolas.

<span class="mw-page-title-main">Cyrtarachninae</span> Subfamily of spiders

Cyrtarachninae is a subfamily of spiders in the family Araneidae. The group has been circumscribed in several different ways. It originated as the group Cyrtarachneae, described by Eugène Simon in 1892. The group was later treated at different ranks: as a tribe, both under Simon's name and as Cyrtarachnini, and as the subfamily Cyrtarachninae. Circumscriptions have varied. The broadest circumscription, Cyrtarachninae sensu lato (s.l.), includes three of Simon's original groups, including the bolas spiders. Unlike most araneids, members of the subfamily do not construct orb webs, some not using webs at all to capture prey, some using one or more sticky drops on a single line, while others construct webs with few widely spaced non-spiral threads, some triangular. Many have been shown to attract prey by producing analogues of insect sex pheromones, particularly to attract male moths. Adult females may mimic snails, bird droppings and other objects, and so are able to remain exposed during the day time, capturing prey at night.

Mastophora extraordinaria is a species of spider in the orb-weaver spider family Araneidae. It is found in South America. Like some other species of the genus Mastophora, adult females resemble bird droppings. Mastophora species, including M. extraordinaria, are "bolas spiders" – adult females capture their prey by using a sticky drop on the end of a single line which they swing at the target, usually a male moth attracted by the release of an analogue of the attractant sex pheromone produced by the female moth. Juveniles and adult males do not use a bolas, catching prey with their legs alone.

Cladomelea akermani is a species of spider in the orb-weaver spider family Araneidae, found in South Africa. Cladomelea species, including C. akermani, are "bolas spiders" – adult females capture their prey by using a sticky drop on the end of a single line which they swing, usually catching male moths attracted by the release of an analogue of the attractant sex pheromone produced by the female moth. Juvenile and adult male bolas spiders do not use a bolas, catching prey with their legs alone.

<i>Cladomelea debeeri</i> Species of spider

Cladomelea debeeri is a species of spider in the orb-weaver spider family Araneidae, found in South Africa. It was first described in 2004. Cladomelea species, including C. debeeri, are "bolas spiders" – adult females capture their prey by using one or more sticky drops on the end of a line which they swing, usually catching male moths attracted by the release of an analogue of the attractant sex pheromone produced by the female moth. Juvenile and adult male bolas spiders do not use a bolas, catching prey with their legs alone.

Exechocentrus lancearius is a species of spider in the orb-weaver spider family Araneidae, found only in Madagascar. It was initially described from a partial specimen of an adult female. The first description of a complete specimen and its prey-catching behaviour was published in 2012. E. lancearius is a bolas spider. Rather than using a web, adult females catch their prey by using a line with one or two sticky drops which they swing.

Ordgarius monstrosus is a species of spider in the orb-weaver spider family Araneidae, found in Queensland, Australia. O. monstrosus is a bolas spider. Rather than using a web, adult females catch their prey by using a line with one or two sticky drops which they swing.

<i>Ordgarius sexspinosus</i> Species of spider

Ordgarius sexspinosus is a species of spider in the orb-weaver spider family Araneidae, found from India to Japan and Indonesia. O. sexspinosus is a bolas spider. Rather than using a web, adult females catch their prey by using a line with one or two sticky drops which they swing.

<i>Pasilobus hupingensis</i> Species of spider

Pasilobus hupingensis is a species of spider in the orb-weaver spider family Araneidae, found in China and Japan. Females of the genus Pasilobus construct "spanning-thread webs" with only two sectors, making them appear triangular. Widely spaced threads with sticky drops span the three radii of these webs. One end is attached in such a way that it readily breaks free. When a prey item is caught on one of these threads, the line parts at this end and the prey hangs from the web until it is hauled up by the spider.

References

  1. 1 2 Foelix (2011), p. 185.
  2. 1 2 3 Levi, H.W. (2003). "The bolas spiders of the genus Mastophora (Araneae: Araneidae)". Bulletin of the Museum of Comparative Zoology. 157: 309–382.
  3. 1 2 3 4 Scharff, Nikolaj & Hormiga, Gustavo (2012). "First evidence of aggressive chemical mimicry in the Malagasy orb weaving spider Exechocentrus lancearius Simon, 1889 (Arachnida: Araneae: Araneidae) and description of a second species in the genus" (PDF). Arthropod Systematics & Phylogeny. 70 (2): 107–118. doi: 10.3897/asp.70.e31757 . Retrieved 2021-01-17.
  4. ↑ Anon. (1998). "Fishing spider's dinner makes a magnificent tale" (PDF). Savanna Links. Vol. 5. p. 10. Archived from the original (PDF) on 2006-08-19.
  5. ↑ Piper, Ross (2007). Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals. Westport, Conn.: Greenwood Press. ISBN   978-0-313-33922-6.
  6. 1 2 3 4 5 6 7 8 9 10 11 12 Yeargan, K.V. (1994). "Biology of Bolas Spiders". Annual Review of Entomology. 39: 81–99. doi:10.1146/annurev.en.39.010194.000501.
  7. ↑ "Search". World Spider Catalog. Natural History Museum Bern. Retrieved 2021-01-16.
  8. 1 2 3 4 5 Tanikawa, Akio; Shinkai, Akira & Miyashita, Tadashi (2014). "Molecular Phylogeny of Moth-Specialized Spider Sub-Family Cyrtarachninae, which Includes Bolas Spiders". Zoological Science. 31 (11): 716–720. doi:10.2108/zs140034. PMID   25366153. S2CID   20031154.
  9. ↑ Mello-Leitão, C.F. de (1931). "Contribuicao ao estudo da tribu Mastophoreas". Anais da Academia Brasileira de Ciências. 3: 65–74 – via World Spider Catalog.. Mello-Leitão only uses the Portuguese form Mastophoreas, which has been latinized to Mastophoreae by later authors.
  10. ↑ Gertsch, W.J. (1955). "The North American bolas spiders of the genera Mastophora and Agatostichus". Bulletin of the American Museum of Natural History. 106: 221–254. hdl:2246/1022 . Retrieved 2021-01-16.
  11. ↑ ICZN (1999), "Article 29", International Code of Zoological Nomenclature (fourth edition), London, UK: The International Trust for Zoological Nomenclature, retrieved 2021-01-18
  12. ↑ Dippenaar-Schoeman, Anna Sophia & Foord, Stefan Hendrik (2019). "New records of Cladomelea from South Africa, including the first records of C. longipes (O. Pickard-Cambridge, 1877) (Araneae, Araneidae) outside its type locality". Check List. 15 (6): 1071–1075. doi: 10.15560/15.6.1071 .
  13. 1 2 3 4 5 Scharff, N.; Coddington, J.A.; Blackledge, Todd A.; Agnarsson, Ingi; Framenau, Volker W.; Szűts, Tamás; Hayashii, Cheryl Y. & Dimitrov, Dimitar (2020). "Phylogeny of the orb-weaving spider family Araneidae (Araneae: Araneoidea)". Cladistics. 36 (1): 1–21. doi:10.1111/cla.12382. hdl: 1956/22200 . PMID   34618955. S2CID   149824795.
  14. ↑ Yeargan, K. V.; Quate, L. W. (1997). "Adult male bolas spiders retain juvenile hunting tactics". Oecologia. 112 (4): 572–576. Bibcode:1997Oecol.112..572Y. doi:10.1007/s004420050347. PMID   28307636. S2CID   424262.
  15. ↑ Yeargan, K.V. & Quate, L.W. (1996). "Juvenile bolas spiders attract psychodid flies". Oecologia. 106 (2): 266–271. Bibcode:1996Oecol.106..266Y. doi:10.1007/BF00328607. PMID   28307652. S2CID   23146947.
  16. 1 2 Gemeno, C.; Yeargan, K.V. & Haynes, K.F. (2000). "Aggressive Chemical Mimicry by the Bolas Spider Mastophora hutchinsoni: Identification and Quantification of a Major Prey's Sex Pheromone Components in the Spider's Volatile Emissions". Journal of Chemical Ecology. 26 (3): 1235–1243. doi:10.1023/A:1005488128468. S2CID   40716552.
  17. ↑ Haynes, K.F.; Yeargan, K.V.; Millar, J.G. & Johnson, K.M. (2002). "How does this specialist predator attract more than one species of prey?". Chemoecology. 12 (2): 99–105. doi:10.1007/s00049-002-8332-2. S2CID   41663936.

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