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Revision as of 15:19, 23 October 2012

Sphaerotheriida
Scientific classification
Kingdom:
Animalia
Phylum:
Arthropoda
Subphylum:
Myriapoda
Class:
Diplopoda
Superorder:
Oniscomorpha
Order:
Sphaerotheriida

Brandt, 1833
Families
Synonyms

Sphaerotheriida (from the Latin sphaerium = ball) is an order of millipedes[1] in the subclass Pentazonia,[2] sometimes known as giant pill millipedes. They inhabit the Southern Hemisphere.[3] Like the Northern Hemisphere pill millipedes of the order Glomerida, these millipedes can roll into a ball when disturbed. When they are rolled-up, most sphaerotheriids reach a maximum size of a cherry [4] or golf ball,[5] but some species from Madagascar can even reach the size of a baseball[6] (an example of island gigantism). When rolled-up, predators are unable to unravel giant pill millipedes since the margins of their second and last dorsal plates fit perfectly into one another, creating a sealed ball. A few giant pill millipede species are able to produce sound, the only millipede taxa known to do this.[7] This order of millipedes is also unique in that some African species are used for medicinal purposes.[8]

Morphology

Sphaerotheriida are characterized by a relatively conservative body morphology; superficially all species and genera look the same.[3] Dorsally, their head is followed by twelve body tergites (collum, thoracic shield, and ten normal tergites) and the anal shield. Ventrally, females possess twenty-one leg pairs (forty-two legs in total), while males carry two additional modified leg pairs, the anterior and posterior telopods under their anal shield. The telopods resemble chelae and/or clamp-like structures, which are probably used in holding the female during mating.[3] In the position of the male telopods, the females instead have a sclerotized subanal plate, which in some species such as those belonging to the family Arthrosphaeridae, is enlarged and is used to produce vibrations (stridulation).[3] Furthermore unlike other large-bodied millipede orders, Sphaerotheriida do not have glands that excrete poisonous or ill-smelling substances. Therefore, they depend entirely on their rolling-up behavior for protection.

Sphaerotheriida somewhat resemble the North American and Eurasian pill millipedes of the order Glomerida, but are generally larger in size (20–80 millimetres or 0.8–3.1 inches body length): only the smallest Sphaerotheriida species are as small as the largest members of the order Glomerida. The orders differ in the number of tergites (10 or 11 in Glomerida, 12 in Sphaerotheriida) and legs (17 or 19 in Glomerida, 21 or 23 in Sphaerotheriida), and show great differences in their head morphology and genital openings, among other characters.[9] Both orders have the ability to roll into a perfect ball, protecting the head, antennae, and the vulnerable underside. However, this rolled-up position (volvation) is achieved differently. In Glomerida, the enlarged second body ring (thoracic shield) has a more or less visible gap within which fit the tips of tergites 3–11, whereas in Sphaerotheriida the tips of tergites 3–12 fit perfectly into a groove on the thoracic shield. Juvenile sphaerotheriids show the same gap as the Glomerida.[10] Many giant pill millipede species have special ledges ('locking carinae') on the underside of the tergite tips and the anal shield which can be moved above a brim on the thoracic shield. These millipedes remain passively locked-up since they need not continued muscle contraction to remain in the rolled-up position.

On Madagascar, some giant pill millipede species exhibit island gigantism, reaching a size comparable to a tennis ball, baseball or even a small orange when rolled up.[11] Recently, a very small form was also described from Madagascar: full-grown individuals of the species Microsphaerotherium ivohibiensis being just the size of a pea.[12]

Distribution

In general, sphaerotheriids are found only in the Southern Hemisphere in a Gondwanan distribution.[13] Gondwana was the large southern continent that formed after the breakup of the supercontinent Pangaea.[14] It included the modern-day landmasses of South America, New Zealand, Australia, India, and Madagascar. It is believed that the Sphaerotheriida originated in Gondwana and then diverged into its various forms. Currently, 322 species[15] in 23 genera[3] are known, with numerous new species having been discovered recently.[16] The sphaerotheriids are divided into four families whose distributions do not overlap: Procyliosomatidae, Zephronidae, Sphaerotheriidae and Arthrosphaeridae. The most basal family, Procyliosomatidae, lives in Australia and New Zealand.[3][17] The Zephronidae (synonym Sphaeropoeidae) occurs in southeast Asia from the Himalayas and China south and east to Sulawesi and inhabits some Philippines islands.[3] The family Sphaerotheriidae only occurs in southern Africa with isolated populations in Zimbabwe and Malawi (probably introduced).[5] The Arthrosphaeridae are distributed in southern India and Madagascar.[3]

A few giant pill millipede species have been dispersed by humans. Examples include the Sri Lankan Arthrosphaeridae species A. brandtii which has established a population in the Usambara hills, Tanzania, Africa,[18] as well as some South African Sphaerotherium species which have isolated populations in Malawi.[19] Another likely candidate is Sechelliosoma forcipatum, a small species of the southeast Asian family Zephronidae, currently only known from a single island in the Seychelles.[20]

Ecology

File:Treeclimbing Zarborealis TWesener2004.jpg
Zoosphaerium arborealis

Little is known about the ecology, development and life history of Sphaerotheriids, but all species are most likely detritivores, feeding on dead organic matter such as leaves and wood on the forest floor. Giant pill millipedes, like earthworms, play an important role in decomposition, releasing nutrients that are locked up in decaying organic matter back into the soil.[21][22][23][24] This process is essential for plant nutrition. It is possible that giant pill millipedes use special bacteria in their gut, similar to the way termites do, to get energy out of non-easily digestible material such as lignin.

Like most millipedes, sphaerotheriids mainly inhabit the leaf litter of humid forests. Some species, however, show an arboreal (tree-living) lifestyle,[25] and for these organisms the rolling-up reflex has been suppressed.[11]

Defense against predation

File:Sphaerotheriida.JPG
Giant pill millipede in rolled-up position

Giant pill millipedes are preyed upon by a wide variety of organisms, and their rolling-up ability and tough skeletal armor offer protection against some predators.[26] A few animals specialize on feeding on giant pill millipedes and have evolved special structures or behaviors to overcome their defensive mechanism. Examples include the South African snail family Chlamydephoridae which almost exclusively feeds on giant pill millipedes[27] along with meerkats (Suricata suricata) which have been reported (at least in captivity) to throw rolled-up sphaerotheriids against rocks in order to break them.[26] Besides their rolling-up behavior, it has been suggested that camouflage may be another defense mechanism that giant pill millipedes use specifically to protect them from animals that hunt using vision, such as birds. Sphaerotheriids also need to cope with internal parasites, with several species of nematodes living in them exclusively.[28][29]

Pets

Sphaerotheriids are commonly found in the pet trade and are kept by terrarium enthusiasts in Japan, Europe, and the United States. The species usually either come from Madagascar or Malaysia. The survival rate for giant pill millipedes in captivity is quite low; they rarely live longer than one month. The reasons for their low survival rate in captivity is little understood. Some explanations include their reliance on low calorie foods, their susceptibility to parasites, their dependence on special climatic conditions (<24°C/75°F at day, <18°C/65°F at night), and weakness caused during transport.[30] Since no captive breeding programs have been initiated, all animals found in stores are caught in the wild.

Taxonomy

Arthrosphaera Pocock, 1895 – 40 species, India, Sri Lanka
Zoosphaerium Pocock, 1895 – 55 species, Madagascar
Sphaeromimus de Saussure & Zehntner, 1902 – 3 species, southeast Madagascar
Microsphaerotherium Wesener & van den Spiegel, 2007 – 2 species, Madagascar
Procyliosoma Silvestri, 1917 – 11 species, Eastern Australia, Tasmania, New Zealand
Bothrobelum Verhoeff, 1924 – monotypic, Borneo
Cryxus Leach, 1814 – monotypic, Asia
Indosphaera Attems, 1935 – 2 species N. India, Myanmar
Kophosphaera Attems, 1935 – 5 species, N. India, Nepal
Leptotelopus Silvestri, 1897 – monotypic, Myanmar
Prionobelum Verhoeff, 1924 – 8 species, Vietnam, SW China
Sphaerobelum Verhoeff, 1924 – 4 species, Vietnam
Zephronia Gray, 1832 – 37 species N. India, Myanmar, Malayan Peninsula, Java, Sumatra, Borneo
Sphaeropoeus Brandt, 1833 – 22 species, N. India, Myanmar, Malayan Peninsula, Java, Sumatra, Borneo
Tigridosphaera Jeekel, 2000 – 4 species, Malayan Peninsula
Castanotherium Pocock, 1895 – 50 species, Indonesian Islands, Philippines
Castanotheroides Chamberlin, 1921 – 3 species, Philippines
Sechelliosoma Mauriès, 1980 – monotypic, Seychelles
Rajasphaera Attems, 1935 – monotypic, Borneo
Sphaerotherium Brandt, 1833 – 54 species, South Africa, Zimbabwe
Kylindotherium Attems, 1926 – monotypic, South Africa
Epicyliosoma Silvestri, 1917 – 15 species, Australia
Cynotelopus Jeekel, 1986 – monotypic, southwest Australia

References

  1. ^ R. L. Hoffman (1980). "Contributions à l'étude de la faune terrestre des îles granitiques de l'archipel des Séchelles". Revue Zoologique africaine. 94 (1): 138–168.
  2. ^ William A. Shear & Gregory D. Edgecombe (2010). "The geological record and phylogeny of the Myriapoda". Arthropod Structure & Development. 39 (2–3): 174–190. doi:10.1016/j.asd.2009.11.002. PMID 19944188.
  3. ^ a b c d e f g h Thomas Wesener & Didier van den Spiegel (2009). "A first phylogenetic analysis of giant pill millipedes (Diplopoda: Sphaerotheriida), a new model Gondwanan taxon, with special emphasis on island gigantism". Cladistics. 25 (6): 545–573. doi:10.1111/j.1096-0031.2009.00267.x. {{cite journal}}: Text "Science" ignored (help)
  4. ^ Beverley A. Holloway (1956). "Revision of the New Zealand pill millipedes (Oniscomorpha, Sphaerotheridae)" (PDF). Transactions of the Royal Society of New Zealand. 84 (2): 431–446.
  5. ^ a b D. van den Spiegel, S. I. Golovatch & M. Hamer (2003). "Revision of some of the oldest species in the millipede genus Sphaerotherium, Brandt, 1833 (Diplopoda, Sphaerotheriida, Sphaerotheriidae), with new synonymies". African Invertebrates. 43: 143–181.
  6. ^ Thomas Wesener & Johann-Wolfgang Wägele (2008). "The giant pill millipedes of Madagascar: revision of the genus Zoosphaerium (Myriapoda, Diplopoda, Sphaerotheriida)" (PDF). Zoosystema. 30 (1): 5–85.
  7. ^ U. Haacker (1968). "Das Sexualverhalten von Sphaerotherium dorsale (Myriapoda, Diplopoda)". Verhandlungen der Deutschen Zoologischen Gesellschaft. 3: 454–463.
  8. ^ D. G. Herbert, M. L. Hamer, M. Mander, N. Mkhize & F. Prins (2003). "Invertebrate animals as a component of the traditional medicine trade in KwaZulu-Natal, South Africa". African Invertebrates. 44 (2): 327–344.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  9. ^ H. Enghoff (1984). "Phylogeny of millipedes – a cladistic analysis". Journal of Zoological Systematics and Evolutionary Research. 22: 8–26. doi:10.1111/j.1439-0469.1984.tb00559.x.
  10. ^ H. P. Wesner. "Die Einkugelung" (in German).
  11. ^ a b T. Wesener & P. Sierwald (2005). "New giant pill millipede species from the littoral forest of Madagascar (Diplopoda, Sphaerotheriida, Zoosphaerium)" (PDF). Zootaxa. 1097: 1–60.
  12. ^ T. Wesener & D. van den Spiegel (2007). "Microsphaerotherium ivohibiensis, a new genus and species of giant-pill millipedes from Madagascar (Diplopoda, Sphaerotheriidaa, Arthrosphaerinae)". Journal of Afrotropical Zoology. 3: 153–160.
  13. ^ C. A. W. Jeekel (1974). "The group taxonomy and geography of the Sphaerotheriida (Diplopoda)". Symposia of the Zoological Society of London. 32: 41–52.
  14. ^ Jason R. Ali & Jonathan C. Aitchison (2008). "Gondwana to Asia: plate tectonics, paleogeography and the biological connectivity of the Indian subcontinent from the Middle Jurassic through latest Eocene (166–35 Ma". Earth-Science Reviews. 88 (3): 145–166. doi:10.1016/j.earscirev.2008.01.007.
  15. ^ T. Wesener, I. Bespalova & P. Sierwald (2010). "Madagascar's living giants: discovery of five new species of endemic giant pill millipedes from Madagascar (Diplopoda: Sphaerotheriida: Arthrosphaeridae: Zoosphaerium)". African Invertebrates. 51 (1): 133–161.
  16. ^ Thomas Wesener (2009). "Unexplored richness: discovery of 31 new species of giant pill millipedes endemic to Madagascar, with a special emphasis on microendemism (Diplopoda, Sphaerotheriida)" (PDF). Zootaxa. 2097: 1–134.
  17. ^ C. A. W. Jeekel (1986). "Millipedes from Australia". Beaufortia. 36 (3): 35–50.
  18. ^ H. Enghoff (1978). "Arthrosphaera cf. brandti (Humbert), a giant pill millipede found in Tanzania, probably introduced from Sri Lanka". Revue de zoologie africaine. 91 (4): 997–999.
  19. ^ D. van den Spiegel (2002). "On the occurrence of Sphaerotherium punctulatum in Malawi (Diplopoda: Sphaerotheriidae)". Musee Royal de L'Afrique Centrale, Tervuren, Zoologique. 290: 171–174.
  20. ^ J.-P. Mauriés (1980). "Contributions à l'étude de la faune terrestre des îles granitiques de l'archipel des Séchelles". Revue Zoologique africaine. 94 (1): 138–168.
  21. ^ C. S. Crawford (1992). "Millipedes as model detritivores". Berichte des Naturwissenschaftlich-Medizinischen Verein Innsbruck. 10: 277–288.
  22. ^ J. P. Curry (1994). Grassland Invertebrates. Chapman and Hall, London. p. 437. ISBN 0-412-16520-1.
  23. ^ James M. Lawrence & Michael J. Samways (2003). "Litter breakdown by the Seychelles giant millipede and the conservation of soil process on Cousine Island, Seychelles". Biological Conservation. 113: 125–132. doi:10.1016/S0006-3207(02)00355-5.
  24. ^ K. M. Ashwini & K. R. Sridhar (2002). "Towards organic farming with the millipede Arthrosphaera magna" (PDF). Current Science (Bangalore). 82 (1): 20–22.
  25. ^ U. Haacker & S. Fuchs (1972). "Tree climbing in pill-millipedes". Oecologia. 10 (2): 191–192. doi:10.1007/BF00347991.
  26. ^ a b Thomas Eisner & Joseph A. David (1967). "Mongoose throwing and smashing millipedes". Science. 155 (3762): 577–579. doi:10.1126/science.155.3762.577. PMID 17737407.
  27. ^ D. G. Herbert. (2000). "Dining on diplopods: remarkable feeding behaviour in chlamydephorid slugs (Mollusca: Gastropoda)". Journal of Zoology. 251 (1): 1–5. doi:10.1111/j.1469-7998.2000.tb00586.x.
  28. ^ Martin L. Adamson & Daniel van Waerebeke (1984). "Xustrostoma margarettae n. gen., n. sp. (Rhigonematidae; Nematoda) from a sphaeroteroid (order Glomerida) diplopod in Madagascar". Canadian Journal of Zoology. 62 (10): 2092–2096. doi:10.1139/z84-303. ISSN 0008-4301.
  29. ^ Daniel van Waerebeke (1985). "Glomerinema ratsimamangi n.gen., n.sp. (Nematoda, Rhigonematidae) parasite de Glomeris (Diplopode) à Madagascar: description et spermiogenese". Annales de Parasitologie Humaine et Comparée. 60 (1): 23–32. ISSN 0003-4150.
  30. ^ "Pill Millipede Care Sheet". Golden Phoenix Exotica.
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