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Giant Pill-Millipedes (Order Sphaerotheriida)

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Giant Pill-Millipede
Pill Millipede, Spherotheriida
Scientific classification
Kingdom:
Phylum:
Class:
Order:
Sphaerotheriida
Distribution
  • 15
File:Distribution-Sphaerotheriida.jpeg
Range of the Giant Pill-Millipedes(green)

Order Sphaerotheriida Brandt, 1833 (Class Diplopoda, Millipedes)

Synonym: Chorizocerata Verhoeff

Families:
Arthrosphaeridae
Procyliosomatidae
Sphaerotheriidae
Zephronidae

Giant Pill-Millipedes (order Sphaerotheriida, Latin from sphaerium = ball) is an order of millipedes[1] in the subclass Pentazonia.[2] inhabiting the Southern Hemisphere.[3] Like the Northern Hemisphere Pill-Millipedes (order Glomerida), these millipedes can roll into a perfect 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] or 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 plate fit perfectly into one another creating a sealed ball. A few Giant Pill-Millipede species are able to produce sound, being the only millipede taxa known to do this[7] or an example of Island Gigantism). This order of millipedes is also unique since some African species are used for medicinal purposes.[8]

Morphology

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The Sphaerotheriida are characterized by a relatively conservative body morphology; superficially all species and genera look the same.[3] Like the Northern Hemisphere Pill-Millipedes. 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] Like the Northern Hemisphere Pill-Millipedes. 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] Like the Northern Hemisphere Pill-Millipedes Furthermore unlike other large-bodied millipede orders, Sphaerotheriida do not have repugnatorial glands which excrete poisonous or ill-smelling substances. Therefore, they entirely depend on their rolling-up behavior for protection.

Sphaerotheriida somewhat resemble the North American and Euro-Asian Pill-Millipedes of the order Glomerida, but are generally larger in size (20-80 mm body length) with only the smallest Sphaerotheriida species being as large 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 (17/19 in Glomerida, 21/23 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, antenna, 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 well-visible gap where the tips of tergites 3–11 fit inside, whereas in Sphaerotheriida the tips of tergites 3-12 fit perfectly into a groove on the thoracic shield. Interestingly, 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. This permits the millipedes to remain passively locked-up and not stretch their muscles permanently.

On Madagascar, some Giant Pill-Millipede species exhibit Island Gigantism, reaching a size comparable to a tennis ball, baseball or even small orange when rolled-up.[11] Recently, an example of Island Dwarfism was also described from Madagascar: full-grown individuals of the species Microsphaerotherium ivohibiense being just the size of a pea.[12]

Distribution

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In general, Sphaerotherrids are found only in the South Hemisphere and their distribution mirrors that of a Gondwanan distribution.[13] Gondwana was the large continent that formed to the south 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. These include Procyliosomatidae, Zephronidae, Sphaerotheriidae and Arthrosphaeridae. The basal most family, Procyliosomatidae, lives in Australia and New Zealand.[3][17] The Zephronidae (synonym Sphaeropoeidae) occurs in southeast Asia spanning from the Himalayas and China north to Sulawesi and inhabits some Philippines Islands to the south.[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 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 Seychellois island.[20]

Ecology

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Little is known about the ecology, development and life history of Sphaerotheriids, but all species are most likely detritovores, 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

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Giant-Pill Millipedes are predated on by a wide variety of organisms and their rolling-up ability seems to be a good defense against most predators. Only a few animals specialize on feeding on Giant Pill-Millipedes and as a result 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[26] 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.[27] 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]

Giant Pill-Millipedes as Pets

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Sphaerotheriids are commonly found in the pet trade and are kept by terraristic 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 a 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 (<75° F at day, <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

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  • Family: Arthrosphaeridae
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
  • Family: Procyliosomatidae
Procyliosoma, Silvestri, 1917: 11 species, Eastern Australia, Tasmania, New Zealand
  • Family: Zephronidae
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
  • Family: Sphaerotheriidae


Sphaerotherium Brandt, 1833, 54 species, South Africa, Zimbabwe
Kylindotherium Attems, 1926, monotypic, South Africa
  • Incertae Sedis
Epicyliosoma Silvestri, 1917, 15 species, Australia
Cynotelopus Jeekel, 1986, monotypic, southwest Australia

References

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  1. ^ Hoffman, R. L (1980). "Contributions à l'étude de la faune terrestre des îles granitiques de l'archipel des Séchelles". Revue Zoologique africaine. 94 (1): 138–168. {{cite journal}}: Text "Science" ignored (help)
  2. ^ Shear, W. A. & G. D. Edgecombe (2010). "The geological record and phylogeny of the Myriapoda". Arthropod Structure & Development. 39: 174–190. {{cite journal}}: Text "Science" ignored (help)
  3. ^ a b c d e f g h Wesener, T. & van den Spiegel, D. (2009). "A first phylogenetic analysis of Giant Pill-Millipedes (Diplopoda: Sphaerotheriida), a new model Gondwanan taxon, with special emphasis on island gigantism". Cladistics. 25: 545–573. {{cite journal}}: Text "Science" ignored (help)CS1 maint: multiple names: authors list (link)
  4. ^ Holloway, B. A. (1956). "Revision of the New Zealand Pill Millipedes (Oniscomorpha, Sphaerotheridae)". Transactions of the Royal Society of New Zealand. 84 (2): 431–446. {{cite journal}}: Text "Science" ignored (help)
  5. ^ a b Van den Spiegel, D., Golovatch, S.I., Hamer, M. (2003). "Revision of some of the oldest species in the millipede genus Sphaerotherium, Brandt, 1833 (Diplopoda, Sphaerotheriida, Sphaerotheriidae), with new synonymies". Afr. Invertebr. 43: 143–181. {{cite journal}}: Text "Science" ignored (help)CS1 maint: multiple names: authors list (link)
  6. ^ Wesener, T. & J.-W. Wägele (2008). "The giant pill-millipedes of Madagascar: revision of the genus Zoosphaerium (Myriapoda, Diplopoda, Sphaerotheriida)" (PDF). Zoosystema. 30 (1): 5–85. {{cite journal}}: Text "Science" ignored (help)
  7. ^ Haacker, U (1968). "Das Sexualverhalten von Sphaerotherium dorsale (Myriapoda, Diplopoda)". Verhandlungen der Deutschen Zoologischen Gesellschaft. 3: 454–463. {{cite journal}}: Text "Science" ignored (help)
  8. ^ 8. Herbert, D. G., M. L. Hamer, M. Mander, N. Mkhize, and F. Prins (2003). "Invertebrate animals as a component of the traditional medicine trade in KwaZulu-Natal, South Africa". African Invertebrates. 44 (2). {{cite journal}}: Text "Science" ignored (help); horizontal tab character in |author= at position 3 (help)CS1 maint: multiple names: authors list (link) CS1 maint: numeric names: authors list (link)
  9. ^ Enghoff, H (1984). "Phylogeny of millipedes – a cladistic analysis". Zeitschrift für zoologische Systematik und Evolutionsforschung. 22: 8–26. {{cite journal}}: Text "Science" ignored (help)
  10. ^ "Die Einkugelung". {{cite web}}: Unknown parameter |name= ignored (help)
  11. ^ a b Wesener T. & P. Sierwald (2005a). "New giant pill-millipede species from the littoral forest of Madagascar (Diplopoda, Sphaerotheriida, Zoosphaerium)" (PDF). Zootaxa. 1097: 1–60. {{cite journal}}: Text "Science" ignored (help)
  12. ^ Wesener, T. & D. VandenSpiegel (2007). "Microsphaerotherium ivohibiense, a new genus and species of Giant-Pill Millipedes from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaerinae)". Journal of Afrotropical Zoology. 3: 153–160. {{cite journal}}: Text "Science" ignored (help)
  13. ^ Jeekel, C. A. W (1974). "The group taxonomy and geography of the Sphaerotheriida (Diplopoda)". Symposia of the Zoological Society of London. 32: 41–52. {{cite journal}}: Text "Science" ignored (help)
  14. ^ Ali, J. R. & Aitchison, J. C (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 Review. 88 (3): 145–166. {{cite journal}}: Text "Science" ignored (help)CS1 maint: multiple names: authors list (link)
  15. ^ Wesener, T. Bespalova, I. & P. Sierwald (in press). "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. {{cite journal}}: Text "Science" ignored (help)CS1 maint: multiple names: authors list (link)
  16. ^ Wesener, T (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. {{cite journal}}: Text "Science" ignored (help)
  17. ^ Jeekel, C. A. W (1986). "Millipedes from Australia". Beaufortia. 36 (3): 35–50. {{cite journal}}: Text "Science" ignored (help)
  18. ^ Enghoff, H (1978). "Arthrosphaera cf. brandti (Humbert), a giant pill-millipede found in Tansania, probably introduced from Sri Lanka". Revue de zoologie africaine. 91 (4): 997–999. {{cite journal}}: Text "Science" ignored (help)
  19. ^ Van den Spiegel, D (2002). "On the occurence of Sphaerotherium punctulatum in Malawi (Diplopoda: Sphaerotheriidae)". Musee Royal de L'Afrique Centrale, Tervuren, Zoologique. 290: 171–174. {{cite journal}}: Text "Science" ignored (help)
  20. ^ Mauriés, J.-P (1980a). "Contributions à l'étude de la faune terrestre des îles granitiques de l'archipel des Séchelles". Revue Zoologique africaine. 94 (1): 138–168. {{cite journal}}: Text "Science" ignored (help)
  21. ^ Crawford, C. S (1992). "Millipedes as model detritivores". Berichte des Naturwissenschaftlich-Medizinischen Verein Innsbruck. 10: 277–288. {{cite journal}}: Text "Science" ignored (help)
  22. ^ Curry, J. P (1994). Grassland Invertebrates. Chapman and Hall, London. p. 437.
  23. ^ Lawrence, J. M. & Samways, M. J. (2003). "Litter breakdown by the Seychelles giant millipede and the conservation of soil process on Cousine Island, Seychelles". Biological Conservation. 113: 125–132. {{cite journal}}: Text "Science" ignored (help)CS1 maint: multiple names: authors list (link)
  24. ^ Ashwini, K. M. & Sridhar, K. R (2002). "Towards organic farming with the millipede Arthrosphaera magna". Current Science (Bangalore). 82 (1): 20–22. {{cite journal}}: Text "Science" ignored (help)CS1 maint: multiple names: authors list (link)
  25. ^ Haacker, U. & Fuchs, S (1972). "Tree climbing in Pill-Millipedes". Oecologia. 10: 191–192. {{cite journal}}: Text "Science" ignored (help)CS1 maint: multiple names: authors list (link)
  26. ^ Herbert, D. G. (2000). "Dining on diplopods: remarkable feeding behaviour in chlamydephorid slugs (Mollusca: Gastropoda)". J. Zool., Lond. 251: 1–5. {{cite journal}}: Text "Science" ignored (help)
  27. ^ Eisner, J .C. & David, J. A (1967). "Mongoose throwing and smashing millipedes". Science. 155 (3762): 577–579. {{cite journal}}: Text "Science" ignored (help)CS1 maint: multiple names: authors list (link)
  28. ^ Adamson, Martin L., Waerebeke, Daniel van (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. ISSN 0008-4301. {{cite journal}}: Text "Science" ignored (help)CS1 maint: multiple names: authors list (link)
  29. ^ Waerebeke, Daniel van (1985). "Glomerinema ratsimamangi n.gen., n.sp. (Nematoda,Rhigonematidae) parasite de Glomeris (Diplopode) a Madagascar: description et spermiogenese". Annales de Parasitologie Humaine et Comparee. 60 (1): 23–32. ISSN 0003-4150. {{cite journal}}: Text "Science" ignored (help)
  30. ^ "Pill Millipede Care Sheet". {{cite web}}: Unknown parameter |name= ignored (help)