Zoraptera
| Zoropterans Temporal range: 145–0 Ma Cretaceous to Recent | |
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| Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Subphylum: | Hexapoda |
| Class: | Insecta |
| Subclass: | Pterygota |
| Order: | Zoraptera Silvestri, 1913 |
| Family: | Zorotypidae Silvestri, 1913 |
| Genus: | Zorotypus Silvestri, 1913 |
| Species | |
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See Text | |
The insect order Zoraptera (angel insect) contains a single family, the Zorotypidae, which in turn contains one extant genus with 39 species, Zorotypus as well as 9 extinct species.[1]
Phylogeny
The phylogenetic relationship of the order remains controversial and elusive. At present the best supported position based on morphological traits recognizes the Zoraptera as polyneopterous insects related to the webspinners of the Order Embioptera. However, molecular analysis of 18s ribosomal DNA supports a close relationship with the Superorder Dictyoptera.[2][3][4][5][6]
Description
The name Zoraptera, given by Filippo Silvestri in 1913,[7] is misnamed and potentially misleading: "zor" is Greek for pure and "aptera" means wingless. "Pure wingless" clearly does not fit the winged alate forms, which were discovered several years after the wingless forms had been described.
The members of this order are small insects, 3 millimetres (0.12 in) or less in length, that resemble termites in appearance and in their gregarious behavior. They are short and swollen in appearance. They belong to the hemimetabolous insects. They possess mandibulated biting mouthparts, short cerci (usually 1 segment only), and short antennae with 9 segments. The abdomen is segmented in 11 sections.[8] The maxillary palps have five segments, labial palps three, in both the most distal segment is enlarged. Immature nymphs resemble small adults.
Each species shows polymorphism. Most individuals are the apterous form or "morph", with no wings, no eyes, and no or little pigmentation. A few females and even fewer males are in the alate form with relatively large membranous wings that can be shed at a basal fracture line. Alates also have compound eyes and ocelli, and more pigmentation. This polymorphism can be observed already as two forms of nymphs. Wingspan can be up to 7 millimetres (0.28 in), and the wings can be shed spontaneously. When observed, wings have simple venation.[8] Under good conditions the blind and wingless form predominates, but if their surroundings become too tough, they produce offspring which develop into winged adults with eyes. The wings are paddle shaped, and have reduced venation.
Behavior and ecology
They live in small colonies beneath rotting wood, lacking in mouthparts able to tunnel into sound wood, eating fungal spores and detritus but also can hunt smaller arthropods like mites and collembola.[9]
Zorotypus gurneyi lives in colonies consisting of up to several hundred of individuals. Most commonly the colonies have a size of around 30 individuals, of which about 30% are nymphs, the remainder adults. Zoraptera spend most of their time grooming one another. The grooming process is thought to be a way of removing fungal pathogens.[10]
When two colonies of Z. hubbardi are brought together experimentally, there is no difference in behavior towards members of the own or new colony. Therefore, colonies in the wild might merge easily. Winged forms are rare. The males in such average colonies establish a linear dominance hierarchy in which age or duration of colony membership is the prime factor determining dominance. Males appearing later in colonies are at the bottom of the hierarchical ladder, regardless of their body size. By continually attacking other males, the dominant male monopolizes a harem of females. The members of this harem stay clumped together. There is a high correlation between rank and reproductive success of the males.[11] <ref name Choe87-93">Choe JC 1994; "Sexual selection and mating system in Zorotypus gurneyi Choe (Insecta, Zoraptera ). I Dominance hierarchy and mating success." Behavioral Ecology and Sociobiology 34: 87-93</ref>
Z. barberi lack such a dominance structure but display complex courtship behavior including nuptial feeding. The males possess a cephalic gland that opens in the middle of their head. During courtship they secrete a fluid from this gland and offer it to the female. Acceptance of this droplet by the female acts as behavioral releaser and immediately leads to copulation.[9]
In Z. impolitus, copulation does not occur, but fertilisation is accomplished instead by transfer of a spermatophore from the male to the female. This 0.1-millimetre (0.0039 in) spermatophore contains a single giant sperm cell, which unravels to about the same length as the female herself, 3 millimetres (0.12 in). It is thought that this large sperm cell prevents fertilisation by other males, by physically blocking the female's genital tract.[12][13]
Effects on ecosystem functioning
Zoraptera are thought to provide some important supporting and provisioning services in ecosystems. By consuming detritus, such as dead arthropods, they assist in decomposition and nutrient cycling.[14]
Species
48 living and fossil species are found worldwide, mainly in tropical and subtropical regions around the world. Four species occur north of the Tropic of Cancer, two in the USA and two in Tibet.
Currently there are 39 extant and 9 extinct species.[1]
Zorotypus absonus Engel 2008 - Dominican Republic (Miocene) (fossil)
Zorotypus acanthothorax Engel & Grimaldi 2002 - Myanmar (Cretaceous) (fossil)
Zorotypus amazonensis Rafael & Engel 2006 - Brazil (Amazonas)
Zorotypus barberi Gurney 1938 - Costa Rica, Dominican Republic, French Guiana, Panama, Trinidad and Tobago, Venezuela
Zorotypus brasiliensis Silvestri 1947 - Brazil (Minas Gerais, Parana, Rio de Janeiro, Santa Catarina, São Paulo)
Zorotypus buxtoni Karny 1932 - Samoa
Zorotypus caudelli Karny 1922 - Indonesia (Sumatra), Malaysia (peninsular)
Zorotypus caxiuana Rafael, Godoi & Engel 2008 - Brazil (Para)
Zorotypus cervicornis Mashimo, Yoshizawa & Engel 2013 - Malaysia (peninsular)
Zorotypus ceylonicus Silvestri 1913 - Sri Lanka
Zorotypus congensis van Ryn-Tournel 1971 - Congo (Dem.Rep.)
Zorotypus cramptoni Gurney 1938 - Guatemala
Zorotypus cretatus Engel & Grimaldi 2002 - Myanmar (Cretaceous) (fossil)
Zorotypus delamarei Paulian 1949 - Madagascar
Zorotypus goeleti Engel & Grimaldi 2000 - Dominican Republic (Miocene) (fossil)
Zorotypus guineensis Silvestri 1913 - Ghana, Guinea, Ivory Coast
Zorotypus gurneyi Choe 1989 - Costa Rica, Panama
Zorotypus hamiltoni New 1978 - Colombia
Zorotypus hubbardi Caudell 1918 - USA (Alabama, Arkansas, Delaware, District of Columbia, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maryland, Mississippi, Missouri, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, West Virginia)
Zorotypus hudae Kaddumi 2005 - Jordan (Cretaceous) (fossil)
Zorotypus huxleyi Bolivar y Pieltain & Coronado 1963 - Brazil (Amazonas), Peru
Zorotypus impolitus Mashimo, Engel, Dallai, Beutel & Machida 2013 - Malaysia (peninsular)
Zorotypus javanicus Silvestri 1913 - Indonesia (Java)
Zorotypus juninensis Engel 2000 - Peru
Zorotypus lawrencei New 1995 - Christmas Island
Zorotypus leleupi Weidner 1976 - Ecuador (Galapagos Islands)
Zorotypus longicercatus Caudell 1927 - Jamaica
Zorotypus magnicaudelli Mashimo, Engel, Dallai, Beutel & Machida 2013 - Malaysia (peninsular)
Zorotypus manni Caudell 1923 - Bolivia, Peru
Zorotypus medoensis Huang 1978 - China (Xizang)
Zorotypus mexicanus Bolivar y Pieltain 1940 - Mexico
Zorotypus mnemosyne Engel 2008 - Dominican Republic (Miocene) (fossil)
Zorotypus nascimbenei Engel & Grimaldi 2002 - Myanmar (Cretaceous) (fossil)
Zorotypus neotropicus Silvestri 1916 - Costa Rica
Zorotypus newi Chao & Chen 2000 - Taiwan
Zorotypus novobritannicus Terry & Whiting 2012 - Papua New Guinea (New Britain)
Zorotypus palaeus Poinar 1988 - Dominican Republic (Eocene) (fossil)
Zorotypus philippinensis Gurney 1938 - Philippines
Zorotypus sechellensis Zompro 2005 - Seychelles
Zorotypus shannoni Gurney 1938 - Brazil (Amazonas, Mato Grosso)
Zorotypus silvestrii Karny 1927 - Indonesia (Mentawai Islands)
Zorotypus sinensis Huang 1974 - China (Xizang)
Zorotypus snyderi Caudell 1920 - Jamaica, USA (Florida)
Zorotypus swezeyi Caudell 1922 - USA (Hawaii)
Zorotypus vinsoni Paulian 1951 - Mauritius
Zorotypus weidneri New 1978 - Brazil (Amazonas)
Zorotypus zimmermani Gurney 1939 - Fiji
Xenozorotypus burmiticus Engel & Grimaldi 2002 - Myanmar (Cretaceous) (fossil)
References
- 1 2 Rafael JA, Godoi FDP, Engel MS, 2008; "A new species of Zorotypus from eastern Amazonia, Brazil (Zoraptera: Zorotypidae)" Transactions of the Kansas Academy of Science 111:193-202
- ↑ Yoshizawa K 2007; "The Zoraptera problem: evidence for Zoraptera + Embiodea from the wing base." Systematic Entomology 32: 197-204
- ↑ Yoshizawa K & Johnson KP; 2005; "Aligned 18S for Zoraptera (Insecta): Phylogenetic position and molecular evolution." Molecular Phylogenetics and Evolution 37: 572-580
- ↑ Engel MS & Grimaldi DA; 2002; "The first mesozoic Zoraptera (Insecta)." American Museum Novitates 3362:1-20
- ↑ Ishiwata K, Sasaki G,Ogawa J, Miyata T & Su Z-H; 2011; "Phylogenetic relationships among insect orders based on three nuclear protein-coding gene sequences." Molecular Phylogenetics and Evolution 58:169-180
- ↑ Wang, X.; Engel, M.S.; Rafael, J.A.; Dang, K.; Wu, H.; Wang, Y.; Xie, Q.; Bu, W. (2013). "A unique box in 28S rRNA is shared by the enigmatic insect order Zoraptera and Dictyoptera". PLoS ONE 8 (1): e53679. doi:10.1371/journal.pone.0053679.
- ↑ Silvestri, F. (1913); "Descrizione di un nuovo ordine di insetti." Bol. Lab. Zool. Gen. Agric. Portici 7:193–209.
- 1 2 Gullan; Granston (2005). The Insects: An Outline of Entomology.
- 1 2 Choe JC; 1997; "The evolution of mating systems in the Zoraptera: Mating variations and sexual conflicts." Pages 130-145 in Choe JC & Crespi BJ. (Editors), "The Evolution of Mating Systems in Insects and Arachnids." Cambridge Univ. Press, Cambridge, 575 pp.
- ↑ "Zoraptera (Insects)". what-when-how.com. Retrieved 2016-02-22.
- ↑ Choe JC 1994; "Sexual selection and mating system in Zorotypus gurneyi Choe (Insecta, Zoraptera). II Determinants and dynamics of dominance." Behavioral Ecology and Sociobiology 34: 233-237
- ↑ R Dallai et al, "Divergent mating patterns and a unique mode of external sperm transfer in Zoraptera, an enigmatic group of pterygote insects",Naturwissenschaften, June 2013, 100: 6 , pp 581-594, doi:10.1007/s00114-013-1055-0
- ↑ "The tiny insect with the massive sperm", New Scientist 29 May 2013, 2919 p 17
- ↑ Engel, Michael (2007). "The Zorotypidae of Fiji (Zoraptera)" (PDF). Bishop Museum Occasional Papers.
General references
- Costa JT 2006 Psocopera and Zoraptera. In: Costa JT The other Insect Societies. The Belknap Press of Harvard University Press, Cambridge, MA and London, UK pp 193–211
- Grimaldi, D. and Engel, M.S. (2005). Evolution of the Insects. Cambridge University Press. ISBN 0-521-82149-5.
- Hubbard, M D (1990) A Catalog of the Order Zoraptera (Insecta). Insecta Mundi 4(1-4)
- Rafael, J.A.; Engel, M.S. (2006). "A new species of Zorotypus from Central Amazonia, Brazil (Zoraptera: Zorotypidae)". American Museum Novitates 3528: 1–11. doi:10.1206/0003-0082(2006)3528[1:ANSOZF]2.0.CO;2.
- Kaddumi, H.F. (2005). Amber of Jordan, the Oldest Prehistoric Insects in Fossilized Resin. Publications of the Eternal River Museum of Natural History, Amman. 168 pp.
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