{{Short description|Group of spiders that capture prey with a bolas}} {{Automatic taxobox |name=Bolas spiders |image=Bolas Spider - Mastophora phrynosoma hunting, Julie Metz Wetlands, Woodbridge, Virginia (24843011518).jpg |image_caption=''Mastophora phrynosoma'' with bolas, Virginia, US |taxon=Bolas spiders |subdivision_ranks=Genera |subdivision = {{Linked genus list |Cladomelea|Simon, 1895 |Exechocentrus|Simon, 1889 |''Mastophora''|Holmberg, 1876 |Ordgarius|Keyserling, 1886 }} |subdivision_ref={{sfnp|Foelix|2011|p=185}}<ref name=Levi03/><ref name=SchaHorm12/> }}
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'''<ref name=Anon98/> (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.
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== [[File:Bolas Spider - Mastophora bisaccata, Okaloacoochee Slough WMA, Florida crop.jpg|thumb|''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 {{cvt|10|–|20|mm|in|frac=4}}, while the males, which are much smaller, have a body length usually less than {{cvt|2|mm|in|frac=16}}. 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.<ref name=Pipe07/><ref name=Year94/>
==Taxonomy== ===Genera=== Bolas-using spiders are species of four related genera.{{sfnp|Foelix|2011|p=185}}<ref name=Levi03/><ref name=SchaHorm12/> (Distributions are from the World Spider Catalog.<ref name=WSC_search/>) * ''Cladomelea'' <small>Simon, 1895</small> – Central and South Africa * ''Exechocentrus'' <small>Simon, 1889</small> – Madagascar * ''Mastophora'' <small>Holmberg, 1876</small> (syn. ''Agathostichus'') – North and South America * ''Ordgarius'' <small>Keyserling, 1886</small> (syn. ''Dicrostichus'') – Australia north to India, China and Japan The use of a bolas by the Madagascan genus ''Exechocentrus'' was not reported until 2012,<ref name=SchaHorm12/> 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'',<ref name=TaniShinMiya14/> 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''.<ref name=Mell31/> Gertsch in 1955 also placed bolas spiders in the tribe Mastophoreae, again including ''Acantharachne'' (under the synonym ''Acantharanea'').<ref name=Gert55/> 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".<ref name=ICZN99/> The tribe has been elevated to a separate subfamily Mastophorinae.<ref name=DippFoor19/> The informal name "mastophorines" has also been used, including both bolas-using spiders and a related genus.<ref name=SchaCoddBlacAgna20/>
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.''<ref name=TaniShinMiya14/> 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.<ref name=SchaCoddBlacAgna20/> 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.<ref name=TaniShinMiya14/>
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.<ref name=TaniShinMiya14/> (The dashed line shows where the genus ''Exechocentrus'' would be expected to fall based on a 2020 study.<ref name=SchaCoddBlacAgna20/>) 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.<ref name=TaniShinMiya14/> {{clade |label1=Cyrtarachninae ''s.l.'' |1={{clade |label1=mastophorines |1={{clade |grouplabel1=webless |1=''Celaenia excavata'' |barbegin1=#666 |barend1=#666 |grouplabel2=bolas |2={{clade |1=''Exechocentrus lancearius''<ref name=SchaCoddBlacAgna20/> |barbegin1=#F66|state1=dashed |2={{clade |1={{clade |1=''Mastophora phrynosoma'' |bar1=#F66 |2=''Ordgarius magnificus'' |bar2=#F66 }} |2={{clade |1=''Ordgarius hobsoni'' |bar1=#F66 |2=''Ordgarius sexspinosus'' |barend2=#F66 }} }} }} }} |label2=cyrtarachnines |2={{clade |grouplabel1=spanning-thread web |1={{clade |1=''Cyrtarachne'' (5 spp.) |barbegin1=#66F |2=''Paraplectana'' (2 spp.) |bar2=#66F |3={{clade |1=''Poecilopachys australasia'' |bar1=#66F |2={{clade |1=''Cyrtarachne nagasakiensis'' |bar1=#66F |2={{clade |1=''Cyrtarachne yunoharuensis'' |barend1=#66F |2={{clade |grouplabel1=triangular web |1=''Pasilobus hupingensis'' |barbegin1=#0F0 |barend1=#0F0 }} }} }} }} }} }} }} }}
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".<ref name=SchaCoddBlacAgna20/>
==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== [[File:Arachtober 23 -2 - Bolas Spider - Mastophora phrynosoma and egg sac, Julie Metz Wetlands, Woodbridge, Virginia (37840647492).jpg|thumb|''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.<ref name=Year94/>
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.<ref name=Year94/>
==Prey capture== thumb|upright|''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.<ref name=Levi03/><ref name=Year94/> 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.<ref name=Year94/>
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.<ref name=Year94/> [[File:Mastophora et sa proie 2.jpg|thumb|centre|upright=2.5|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.<ref name=Year94/> 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.<ref name=Year94/>
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''.<ref name=Year97/><ref name=YearQuat96/>
===Drop structure=== The sticky drops of the bolas have a diameter of about {{cvt|2.5|mm|in|frac=10}}. 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.<ref name=Year94/> ''Exechocentrus lancearius'' uses a bolas with two globules.<ref name=SchaHorm12/> 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.<ref name=Year94/>
===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.<ref name=Year94/> 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.<ref name=GemeYearHayn00/> 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.<ref name=HaynYearMillJohn02/><ref name=GemeYearHayn00/>
==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.<ref name=Year94/>
==References== {{Reflist|refs= <ref name=Anon98>{{Citation |mode=cs1 |last1=Anon. |date=1998 |title=Fishing spider's dinner makes a magnificent tale |work=Savanna Links |volume=5 |page=10 |url=http://www.savanna.ntu.edu.au/downloads/savlinks5.pdf |archive-url=https://web.archive.org/web/20060819183222/http://www.savanna.ntu.edu.au/downloads/savlinks5.pdf |archive-date=2006-08-19 }}</ref>
<ref name=DippFoor19>{{Citation |mode=cs1 |last1=Dippenaar-Schoeman |first1=Anna Sophia |last2=Foord |first2=Stefan Hendrik |date=2019 |title=New records of ''Cladomelea'' from South Africa, including the first records of ''C. longipes'' (O. Pickard-Cambridge, 1877) (Araneae, Araneidae) outside its type locality |journal=Check List |volume=15 |issue=6 |pages=1071–1075 |doi=10.15560/15.6.1071 |name-list-style=amp |doi-access=free |hdl=10019.1/121599 |hdl-access=free }}</ref>
<ref name=GemeYearHayn00>{{Citation |mode=cs1 |last1=Gemeno |first1=C. |last2=Yeargan |first2=K.V. |last3=Haynes |first3=K.F. |date=2000 |title=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=Journal of Chemical Ecology |volume=26 |issue=3 |pages=1235–1243 |doi=10.1023/A:1005488128468 |s2cid=40716552 |name-list-style=amp }}</ref>
<ref name=Gert55>{{Citation |mode=cs1 |last1=Gertsch |first1=W.J. |date=1955 |title=The North American bolas spiders of the genera ''Mastophora'' and ''Agatostichus'' |journal=Bulletin of the American Museum of Natural History |volume=106 |pages=221–254 |hdl=2246/1022 |url=http://digitallibrary.amnh.org/handle/2246/1022 |access-date=2021-01-16 }}</ref>
<ref name=HaynYearMillJohn02>{{Citation |mode=cs1 |last1=Haynes |first1=K.F. |last2=Yeargan |first2=K.V. |last3=Millar |first3=J.G. |last4=Johnson |first4=K.M. |date=2002 |title=How does this specialist predator attract more than one species of prey? |journal=Chemoecology |volume=12 |issue=2 |pages=99–105 |doi=10.1007/s00049-002-8332-2 |s2cid=41663936 |name-list-style=amp }}</ref>
<ref name=ICZN99>{{Citation |last1=ICZN |date=1999 |contribution=Article 29 |title=International Code of Zoological Nomenclature (fourth edition) |publication-place=London, UK |publisher=The International Trust for Zoological Nomenclature |contribution-url=https://code.iczn.org/formation-and-treatment-of-names/article-29-family-group-names |access-date=2021-01-18 }}</ref>
<ref name=Levi03>{{Citation |mode=cs1 |last1=Levi |first1=H.W. |date=2003 |title=The bolas spiders of the genus ''Mastophora'' (Araneae: Araneidae) |journal=Bulletin of the Museum of Comparative Zoology |volume=157 |pages=309–382 }}</ref>
<ref name=Mell31>{{Citation |mode=cs1 |last1=Mello-Leitão |first1=C.F. de |date=1931 |title=Contribuicao ao estudo da tribu Mastophoreas |journal=Anais da Academia Brasileira de Ciências |volume=3 |pages=65–74 |via=World Spider Catalog }}. Mello-Leitão only uses the Portuguese form ''Mastophoreas'', which has been latinized to Mastophoreae by later authors.</ref>
<ref name=Pipe07>{{Citation |mode=cs1 |last1=Piper |first1=Ross |date=2007 |title=Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals |publication-place=Westport, Conn. |publisher=Greenwood Press |isbn=978-0-313-33922-6 }}</ref>
<ref name=SchaCoddBlacAgna20>{{Citation |mode=cs1 |last1=Scharff |first1=N. |last2=Coddington |first2=J.A. |last3=Blackledge |first3=Todd A. |last4=Agnarsson |first4=Ingi |last5=Framenau |first5=Volker W. |last6=Szűts |first6=Tamás |last7=Hayashii |first7=Cheryl Y. |last8=Dimitrov |first8=Dimitar |date=2020 |title=Phylogeny of the orb-weaving spider family Araneidae (Araneae: Araneoidea) |journal=Cladistics |volume=36 |issue=1 |pages=1–21 |doi=10.1111/cla.12382 |pmid=34618955 |name-list-style=amp |hdl=1956/22200 |s2cid=149824795 |hdl-access=free }}</ref>
<ref name=SchaHorm12>{{Citation |mode=cs1 |last1=Scharff |first1=Nikolaj |last2=Hormiga |first2=Gustavo |date=2012 |title=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 |journal=Arthropod Systematics & Phylogeny |volume=70 |issue=2 |pages=107–118 |doi=10.3897/asp.70.e31757 |url=https://core.ac.uk/download/pdf/269220898.pdf |access-date=2021-01-17 |name-list-style=amp |doi-access=free }}</ref>
<ref name=TaniShinMiya14>{{Citation |mode=cs1 |last1=Tanikawa |first1=Akio |last2=Shinkai |first2=Akira |last3=Miyashita |first3=Tadashi |date=2014 |title=Molecular Phylogeny of Moth-Specialized Spider Sub-Family Cyrtarachninae, which Includes Bolas Spiders |journal=Zoological Science |volume=31 |issue=11 |pages=716–720 |doi=10.2108/zs140034 |pmid=25366153 |s2cid=20031154 |name-list-style=amp }}</ref>
<ref name=Year94>{{Citation |mode=cs1 |last1=Yeargan |first1=K.V. |date=1994 |title=Biology of Bolas Spiders |journal=Annual Review of Entomology |volume=39 |pages=81–99 |doi=10.1146/annurev.en.39.010194.000501 }}</ref>
<ref name=Year97>{{Cite journal |last1=Yeargan |first1=K. V. |last2=Quate |first2=L. W. |date=1997 |title=Adult male bolas spiders retain juvenile hunting tactics |journal=Oecologia |volume=112 |issue=4 |pages=572–576 |doi=10.1007/s004420050347 |pmid=28307636|bibcode=1997Oecol.112..572Y |s2cid=424262 }}</ref>
<ref name=YearQuat96>{{Citation |mode=cs1 |last1=Yeargan |first1=K.V. |last2=Quate |first2=L.W. |date=1996 |title=Juvenile bolas spiders attract psychodid flies |journal=Oecologia |volume=106 |issue=2 |pages=266–271 |doi=10.1007/BF00328607 |pmid=28307652 |bibcode=1996Oecol.106..266Y |s2cid=23146947 |name-list-style=amp }}</ref>
<ref name=WSC_search>{{Citation |mode=cs1 |title=Search |work=World Spider Catalog |publisher=Natural History Museum Bern |url=https://www.wsc.nmbe.ch/search |access-date=2021-01-16 }}</ref> }}
==Bibliography== *{{Citation |mode=cs1 |last=Foelix |first=Rainer F. |year=2011 |title=Biology of Spiders |edition=3rd p/b |publisher=Oxford University Press |isbn=978-0-19-973482-5 }} *{{Cite journal |last1=Scharff |first1=N. |last2=Coddington |first2 =J.A. |date=1997 |title=A phylogenetic analysis of the orb-weaving spider family Araneidae (Arachnida, Araneae) |journal=Zoological Journal of the Linnean Society |volume=120 |issue=4 |pages=355–434 |doi=10.1111/j.1096-3642.1997.tb01281.x |doi-access=free }}
==External links== * [http://www.deh.gov.au/cgi-bin/species-bank/sbank-treatment.pl?id=75613 Description, Australian distribution and pictures of ''Ordgarius magnificus''] * [https://web.archive.org/web/20060823084521/http://www.amonline.net.au/factsheets/bolas_spider.htm Australian Museum Online - Bolas spiders] * [https://davids-pics.blogspot.com/2007/05/id-please.html Magnificent Bolas Spider - Pictures and Egg Sac] * [http://www.dailykos.com/story/2006/6/2/191228/0852 Science Spider Friday: Bolas spiders] * https://www.archentoflor.com/Mastophora__sauvegard%E9.html {{Taxonbar|from1=Q104864420|from2=Q891260}}
{{DEFAULTSORT:Bolas Spiders}} Category:Araneidae Category:Arthropod common names