Orb-weaver spider

Orb weaver spider
Argiope catenulata
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Arachnida
Order:	Araneae
Suborder:	Araneomorphae
Superfamily:	Araneoidea
Family:	Araneidae Simon, 1895
Selected genera
Aculepeira Arachnura Araneus Araniella Argiope Augusta Austracantha Bertrana Celaenia Chorizopes Cyclosa Cyrtophora Eriophora Gasteracantha Kaira Larinioides Mangora Mastophora Melychiopharis Micrathena Neoscona Nuctenea Ordgarius Perilla Zygiella
Diversity
168 genera, 3,006 species[1]

The "typical" orb-weaver spiders (family Araneidae) are the most common group of builders of spiral wheel-shaped webs often found in gardens, fields and forests. Their common name is taken from the round shape of this typical web, and the taxon was formerly also referred to as the Orbiculariae.

Orb-weavers have eight similar eyes, legs hairy or spiny and no stridulating organs. The Araneidae family is cosmopolitan, including many well-known large or brightly colored garden spiders. There are 3,006 species in 168 genera worldwide, making Araneidae the third largest family of spiders known (behind Salticidae and Linyphiidae).^[1] The orb-weavers include over 10,000 species and make up about 25% of spider diversity.^[2]

However, orb-webs are also produced by members of other families. The large "golden" orb-weavers (Nephilidae) and the long-jawed orb weavers (Tetragnathidae) were formerly included in the Araneidae; they are indeed closely related to them, being part of superfamily Araneoidea. Their webs are similar to those of the typical orb-weavers, but tend to be less sophisticated and often have an irregular instead of a neat spiral arrangement of the prey-capturing threads. The cribellate or hackled orb-weavers (Uloboridae) belong to a distinct superfamily of the suborder Araneomorphae; their webs are often very sophisticated but Uloboridae use neither poison to kill their prey, nor sticky threads in their web, and probably evolved the orb structure independently. Uloboridae are cribellate, and their threads can be recognized by the fuzzy and dull appearance, which captures prey by a velcro-like mechanism. Even among the Araneoidea, the orb-webs may have evolved at least twice from the three-dimensional webs most common in this superfamily, as typically produced e.g. by tangle-web spiders (Theridiidae).

The orb-web

Argiope sp. sitting on the stabilimentum at the center of the web

Spiderlings in the web near where they hatched

Close up of the cephalothorax on Eriophora sp. (possibly E. heroine or E. pustuosa

Generally, orb-weaving spiders are three-clawed builders of flat webs with sticky spiral capture silk. The building of a web is an engineering feat, begun when the spider floats a line on the wind to another surface. The spider secures the line and then drops another line from the center, making a "Y". The rest of the scaffolding follows with many radii of non-sticky silk being constructed before a final spiral of sticky capture silk. The third claw is used to walk on the non-sticky part of the web. Characteristically, the prey insect that blunders into the sticky lines is stunned by a quick bite and then wrapped in silk. If the prey is a venomous insect, such as a wasp, wrapping may precede biting.

Many orb-weavers build a new web each day. Generally, towards evening, the spider will consume the old web, rest for approximately an hour, then spin a new web in the same general location. Thus, the webs of orb-weavers are generally free of the accumulation of detritus common to other species such as black widow spiders.

Some orb-weavers do not build webs at all. Members of the genera Mastophora in the Americas, Cladomelea in Africa and Ordgarius in Australia produce sticky globules, which contain a pheromone analog. The globule is hung from a silken thread dangled by the spider from its front legs. The pheromone analog attracts male moths of only a few species. These get stuck on the globule and are reeled in to be eaten. Interestingly, both types of bolas spiders are highly camouflaged and difficult to locate.

The spiny orb-weaving spiders in the genera Gasteracantha and Micrathena look like plant seeds or thorns hanging in their orb-webs. Some species of Gasteracantha have very long horn-like spines protruding from their abdomens.

One feature of the webs of some orb-weavers is the stabilimentum, a crisscross band of silk through the center of the web. It is found in a number of genera, but Argiope, which includes the common garden spider of Europe as well as the yellow and banded garden spiders of North America, is a prime example. The band has been hypothesized to be a lure for prey, a marker to warn birds away from the web and a camouflage for the spider when it sits in the center of the web. However, recent research suggests that the stabilimentum actually decreases the visibility of the silk to insects, thus making it harder for prey to avoid the web.^[3] The orb-web consists of a frame and supporting radii overlaid with a sticky capture spiral, and the silks used by orb-weaver spiders have exceptional mechanical properties to withstand the impact of flying prey.^[4]

The orb web derived from substrate-bound web, likely an irregular ground web or brushed sheet web.^[5] The change from a ground web to an aerial web allowed for both horizontal and vertical orb webs that could capture flying prey. During the Cretaceous, a radiation of angiosperm plants and their insect pollinators occurred. Fossil evidence shows that the orb web was in existence at this time, which permitted a concurrent radiation of the spider predators along with their insect prey.^[5] The capacity of orb webs to absorb the impact of flying prey led orbicularian spiders to become the dominant predators of aerial insects in many ecosystems.^[2] Insects and spiders have comparable rates of diversification, suggesting that they co-radiated, and the peak of this radiation occurred 100 Mya before the origin of angiosperms.^[6] Vollrath and Selden (2007) make the bold proposition that insect evolution was driven less by flowering plants than by spider predation – particularly through orb webs – as a major selective force.

Most arachnid webs are vertical and the spiders usually hang with their head downward. A few webs, such as those of orb-weaver in the genus Metepiera have the orb hidden within a tangled space of web. Some Metepiera are semi-social and live in communal webs. In Mexico such communal webs have been cut out of trees or bushes and used for living fly paper.^{[citation needed]}

Natural history

The oldest known true orb-weaver is Mesozygiella dunlopi, from the Lower Cretaceous. Several fossils provide direct evidence that the three major orb weaving families, namely Araneidae, Tetragnathidae and Uloboridae, had evolved by this time, about 140 million years ago.^[7] They probably originated during the Jurassic (200 to 140 million years ago). Based on new molecular evidence in silk genes, all three families are very likely to have a common origin.^[5][4][2]

The two families, Deinopoidea and Araneoidea, have similar behavioral sequences and spinning apparatuses to produce architecturally similar webs. The Araneidae weave true viscid silk with an aqueous glue property and the Deinopoidea use dry fibrils and sticky silk.^[4][8] The Deinopoidea (including the Uloboridae), have a cribellum – a flat, complex spinning plate from which the cribellate silk is released. They also have a calamistrum – an apparatus of bristles used to comb the cribellate silk from the cribellum. The Araneoidea, or the "ecribellate" spiders do not have these two structures. The two families of orb-weaving spiders are morphologically very distinct, yet there is much similarity between their web form and web construction behavior. The cribellates retained the ancestral character, yet the cribellum was lost in the escribellates. The lack of a functional cribellum in Araneoids is most likely synapomorphic. If the orb-weaver spiders are a monophyletic group, the fact that only some species in the group lost a feature adds to the controversy. The cribellates are split off as a separate taxon that retained the primitive feature, which makes the lineage paraphyletic and non-synonymous with any real evolutionary lineage. The morphological and behavioral evidence surrounding orb webs led to the disagreement over a single origin or a dual origin.^[9] However, molecular analysis provides more support for a monophyletic origin.^[5][4][2]

Systematics

Main article: List of Araneidae genera

Eriophora transmarina from Australia

The categorization into subfamilies and tribes follows Joel Hallan's Biology Catalog.^[10]

The Nephilidae have been elevated to family status in 2006. Some researchers also consider the genera Leviellus, Parazygiella, Stroemiellus and Zygiella to reside in the family Zygiellidae.

• Araneinae Simon, 1895

• Anepsiini
• Arachnurini
• Araneini
• Arkycini
• Bertranini
• Celaenini
• Cyclosini
• Dolophonini
• Exechocentrini
• Heterognathini
• Hypognathini
• Mangorini
• Poltyini
• Pseudartonini
• Testudinarini
• Ursini
• incertae sedis

Glyptogona Simon, 1884

• Argiopinae

• Cyrtarachninae Simon

• Cyrtarachnini Simon
• Mastophorini

• Cyrtophorinae

• Gasteracanthinae

• Caerostrini
• Gasteracanthini

• Micratheninae

• incertae sedis

• Artonis Simon, 1895
• Colphepeira Archer, 1941
• Enacrosoma Mello-Leitão, 1932
• Encyosaccus Simon, 1895
• Melychiopharis Simon, 1895
• Micrepeira Schenkel, 1953
• Parmatergus Emerit, 1994
• Pronous Keyserling, 1881
• Sedasta Simon, 1894
• Singafrotypa Benoit, 1962
• Tethneus Scudder, 1890 † (fossil, Oligocene)

• Tethneus guyoti (Scudder, 1890) †
• Tethneus henzii (Scudder) †
• Tethneus robustus (Petrunkevitch) †

See also

Arthropods portal

• List of Araneidae species
• Spider families

References

1. ^ ^{a b} Platnick, Norman I. (29 December, 2010). "Currently valid spider genera and species". The World Spider Catalog, Version 11.5. American Museum of Natural History. http://research.amnh.org/iz/spiders/catalog/COUNTS.html. Retrieved 24 May 2011. 
2. ^ ^{a b c d} Todd A. Blackledge, Nikolaj Scharff, Jonathan A. Coddington, Tamas Szüts, John W. Wenzel, Cheryl Y. Hayashi & Ingi Agnarsson (2009). "Reconstructing web evolution and spider diversification in the molecular era". Proceedings of the National Academy of Sciences 106: 5229–5234. doi:10.1073/pnas.0901377106. PMC 2656561. PMID 19289848. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2656561. 
3. ^ Todd A. Blackledge & John W. Wenzel (2000). "The evolution of cryptic spider silk: a behavioral test" (PDF). Behavioral Ecology 11 (2): 142–145. doi:10.1093/beheco/11.2.142. http://www3.uakron.edu/biology/blackledge/PDFs/Behav_Ecol_2000.pdf. 
4. ^ ^{a b c d} Jessica E. Garb, Teresa DiMauro, Victoria Vo & Cheryl Y. Hayashi (2006). "Silk genes support the single origin of orb webs". Science 312: 1762–1762. doi:10.1126/science.1127946. PMID 16794073. 
5. ^ ^{a b c d} David Penney & Vicente M. Ortuño (2006). "Oldest true orb-weaving spider (Araneae: Araneidae)" (PDF). Biology Letters 2: 447–450. doi:10.1098/rsbl.2006.0506. PMC 1686203. PMID 17148427. http://www.pubs.royalsoc.ac.uk/media/biology_letters/rsbl20060506.pdf. 
6. ^ Fritz Vollrath & Paul Selden (2007). "The role of behavior in the evolution of spiders, silks, and webs" (PDF). Annual Review of Ecology, Evolution, and Systematics 38: 819–846. doi:10.1146/annurev.ecolsys.37.091305.110221. http://homepage.mac.com/paulselden/Sites/Website/ARES.pdf. 
7. ^ Enrique Peñalver, David A. Grimaldi & Xavier Delclòs (2006). "Early Cretaceous spider web with its prey" (PDF). Science 312 (5781): 1761–1761. doi:10.1126/science.1126628. PMID 16794072. http://www.bio-nica.info/biblioteca/Penalver2006SpiderWebCretaceous.pdf. 
8. ^ William A. Shear (1986). "The evolution of web-building behavior in spiders: a third generation of hypotheses". In William A. Shear. Spiders: webs, behavior, and evolution. Stanford, California: Stanford University Press. pp. 364–400. ISBN 9780804712033. http://books.google.co.uk/books?id=rc6Vz-cWickC&pg=PA364. 
9. ^ Jonathan A. Coddington (1986). "The monophyletic origin of the orb web". In William A. Shear. Spiders: webs, behavior, and evolution. Stanford, California: Stanford University Press. pp. 319–363. ISBN 9780804712033. http://books.google.co.uk/books?id=rc6Vz-cWickC&pg=PA319. 
10. ^ Joel Hallan. "Araneidae". Biology Catalog. Texas A&M University. http://insects.tamu.edu/research/collection/hallan/Acari/Family/Araneidae.txt. Retrieved February 11, 2011. 

Further reading

• The Life of the Spider by John Crompton. Mentor, 1950.
• "The Orb-Weaving Spiders of Canada and Alaska. Araneae: Uloboridae, Tetragnathidae, Araneidae, Theridiosomatidae. Insects and Arachnids of Canada Series, Part 23." By C. D. Dondale, J. H. Redner, P. Paquin, and H. W. Levi. NRC Research Press, Ottawa, 2003. ISBN 978-0-660-18898-0
• How to Know the Spiders by B. J. Kaston. Dubuque, 1953.
• Spiders by Barbara York Main. Sidney, 1976.
• Biology of Spiders, by Rainer F. Foelix, second edition, 1996
• Levi, H. W. (1993): The new orb-weaver genus Lewisepeira (Araneae: Araneidae). Psyche 100: 127–136. PDF
• Platnick, Norman I. (2010): The world spider catalog: Araneidae, version 11.5. American Museum of Natural History.

External links

• Spiders of Australia
• Spiders of northwestern Europe
• Araneae, Arachnology Home Pages
• World Spider Catalog
• Orb weavers of Kentucky, University of Kentucky
• Pictures of Mangora species
• Gasteracantha cancriformis, spinybacked orbweaver on the University of Florida / Institute of Food and Agricultural Sciences Featured Creatures website
• Neoscona crucifera and N. domiciliorum on the University of Florida / Institute of Food and Agricultural Sciences Featured Creatures website HOE