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== <big>'''Giant Pill-Millipedes (Order Sphaerotheriida)'''</big> ==
{{Draft article check| Sphaerotheriida}}
{{About|the type of animal}}
{{Taxobox
| name = Giant Pill-Millipede
| image = Pill_millipede.jpg
| image_caption = [[Pill Millipede]], ''Spherotheriida''
| regnum = [[Animal]]ia
| phylum = [[Arthropods|Arthropoda]]
| classis = [[Diplopoda]]
| ordo = '''Sphaerotheriida'''
| subdivision_ranks = Distribution
| subdivision = * 15
| range_map = Distribution-Sphaerotheriida.jpeg
| range_map_caption = Range of the Giant Pill-Millipedes(green)
}}

Order Sphaerotheriida Brandt, 1833 (Class Diplopoda, Millipedes)

Synonym: Chorizocerata Verhoeff

Families:<br />
Arthrosphaeridae<br />
Procyliosomatidae<br />
Sphaerotheriidae<br />
Zephronidae<br />
<br />
Giant Pill-Millipedes (order Sphaerotheriida, [[Latin]] from sphaerium = ball) is an [[order]] of [[millipedes]]<ref>{{cite journal |author= Hoffman, R. L |year=1980 |title=Contributions à l'étude de la faune terrestre des îles granitiques de l'archipel des Séchelles |journal= Revue Zoologique africaine |Science |volume=94 |issue=1 |pages=138-168}}</ref> in the subclass [[Pentazonia]].<ref>{{cite journal |author= Shear, W. A. & G. D. Edgecombe |year=2010 |title= The geological record and phylogeny of the Myriapoda|journal= Arthropod Structure & Development |Science |volume=39 |pages=174-190}}</ref> inhabiting the Southern Hemisphere.<ref name=TW>{{cite journal |author= Wesener, T. & van den Spiegel, D. |year=2009 |title=A first phylogenetic analysis of Giant Pill-Millipedes (Diplopoda: Sphaerotheriida), a new model Gondwanan taxon, with special emphasis on island gigantism |journal= Cladistics |Science |volume=25 |pages=545-573 |url= http://www3.interscience.wiley.com/journal/122536047/abstract}}</ref>  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 <ref>{{cite journal |author= Holloway, B. A. |year=1956 |title= Revision of the New Zealand Pill Millipedes (Oniscomorpha, Sphaerotheridae)|journal= Transactions of the Royal Society of New Zealand |Science |volume=84 |issue=2 |pages=431-446}}</ref> or golf ball,<ref name=VS>{{cite journal |author= Van den Spiegel, D., Golovatch, S.I., Hamer, M. |year=2003 |title=Revision of some of the oldest species in the millipede genus Sphaerotherium, Brandt, 1833 (Diplopoda, Sphaerotheriida, Sphaerotheriidae), with new synonymies |journal= Afr. Invertebr. |Science |volume=43 |pages=143-181}}</ref> but some species from Madagascar can even reach the size of a baseball<ref>{{cite journal |author= Wesener, T. & J.-W. Wägele |year=2008 |title= The giant pill-millipedes of Madagascar: revision of the genus Zoosphaerium (Myriapoda, Diplopoda, Sphaerotheriida) |journal= Zoosystema |Science |volume=30 |issue=1 |pages=5-85 |url= http://www.mnhn.fr/museum/front/medias/publication/13338_z08n1a1.pdf}}</ref> 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<ref>{{cite journal |author= Haacker, U |year=1968 |title= Das Sexualverhalten von Sphaerotherium dorsale (Myriapoda, Diplopoda) |journal= Verhandlungen der Deutschen Zoologischen Gesellschaft |Science |volume=3 |pages=454-463}}</ref> or an example of Island Gigantism).  This order of millipedes is also unique since some African species are used for medicinal purposes.<ref>{{cite journal |author= 8.	Herbert, D. G., M. L. Hamer, M. Mander, N. Mkhize, and F. Prins |year=2003 |title= Invertebrate animals as a component of the traditional medicine trade in KwaZulu-Natal, South Africa |journal= African Invertebrates |Science |volume=44 |issue=2}}</ref><br />
<br />

== '''Morphology''' ==
 

The Sphaerotheriida are characterized by a relatively conservative body [[morphology]]; superficially all species and [[genera]] look the same.<ref name=TW/>  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.<ref name=TW/>  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]).<ref name=TW/>  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.<br /><br />

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]].<ref>{{cite journal |author= Enghoff, H |year=1984 |title=Phylogeny of millipedes – a cladistic analysis |journal= Zeitschrift für zoologische Systematik und Evolutionsforschung |Science |volume=22 |pages=8-26}}</ref>  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.<ref>{{cite web |name=Wesner HP [in German] |title=Die Einkugelung |url=http://homepage.ruhr-uni-bochum.de/Thomas.Wesener/einkugelung.htm}}</ref>  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.<br /><br />

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.<ref name=WP>{{cite journal |author= Wesener T. & P. Sierwald |year=2005a |title= New giant pill-millipede species from the littoral forest of Madagascar (Diplopoda, Sphaerotheriida, Zoosphaerium) |journal= Zootaxa |Science |volume=1097 |pages=1-60 |url=http://www.mapress.com/zootaxa/2005f/zt01097p060.pdf}}</ref>  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.<ref>{{cite journal |author= Wesener, T. & D. VandenSpiegel |year=2007 |title= Microsphaerotherium ivohibiense, a new genus and species of Giant-Pill Millipedes from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaerinae) |journal= Journal of Afrotropical Zoology |Science |volume=3 |pages=153-160}}</ref><br />

== '''Distribution''' ==

In general, Sphaerotherrids are found only in the South Hemisphere and their distribution mirrors that of a Gondwanan distribution.<ref>{{cite journal |author= Jeekel, C. A. W |year=1974 |title= The group taxonomy and geography of the Sphaerotheriida (Diplopoda) |journal= Symposia of the Zoological Society of London |Science |volume=32 |pages=41-52}}</ref>   [[Gondwana]] was the large continent that formed to the south after the breakup of the supercontinent [[Pangaea]].<ref>{{cite journal |author= Ali, J. R. & Aitchison, J. C |year=2008 |title= Gondwana to Asia: Plate tectonics, paleogeography and the biological connectivity of the Indian subcontinent from the Middle Jurassic through latest Eocene (166–35 Ma) |journal= Earth-Science Review |Science |volume=88 |issue=3 |pages=145-166}}</ref>  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<ref>{{cite journal |author= Wesener, T. Bespalova, I. & P. Sierwald (in press) |title= Madagascar's living giants: discovery of five new species of endemic giant pill-millipedes from Madagascar (Diplopoda: Sphaerotheriida: Arthrosphaeridae: Zoosphaerium) |journal= African Invertebrates |Science |volume=51 |issue=1 |pages=133-161}}</ref> in 23 genera<ref name=TW/> are known with numerous new species having been discovered recently.<ref>{{cite journal |author= Wesener, T |year=2009 |title= Unexplored richness: discovery of 31 new species of Giant Pill-Millipedes endemic to Madagascar, with a special emphasis on microendemism (Diplopoda, Sphaerotheriida) |journal= Zootaxa |Science |volume=2097 |pages=1-134 |url=http://www.mapress.com/zootaxa/2009/f/z02097p134f.pdf}}</ref>  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.<ref name=TW/><ref>{{cite journal |author= Jeekel, C. A. W |year=1986 |title= Millipedes from Australia |journal= Beaufortia |Science |volume=36 |issue=3 |pages=35-50}}</ref>  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.<ref name=TW/>  The family Sphaerotheriidae only occurs in southern Africa with isolated populations in Zimbabwe and Malawi (probably introduced).<ref name=VS/>  The Arthrosphaeridae are distributed in southern India and Madagascar.<ref name=TW/><br />
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<ref>{{cite journal |author= Enghoff, H |year=1978 |title= Arthrosphaera cf. brandti (Humbert), a giant pill-millipede found in Tansania, probably introduced from Sri Lanka |journal= Revue de zoologie africaine |Science |volume=91 |issue=4 |pages=997-999}}</ref>, as well as some South African Sphaerotherium species which have isolated populations in Malawi.<ref>{{cite journal |author= Van den Spiegel, D |year=2002 |title=On the occurence of Sphaerotherium punctulatum in Malawi (Diplopoda: Sphaerotheriidae) |journal= Musee Royal de L'Afrique Centrale, Tervuren, Zoologique |Science |volume=290 |pages=171-174}}</ref>  Another likely candidate is ''Sechelliosoma forcipatum'', a small species of the southeast Asian family Zephronidae, currently only known from a single [[Seychellois]] island.<ref>{{cite journal |author= Mauriés, J.-P |year=1980a |title=Contributions à l'étude de la faune terrestre des îles granitiques de l'archipel des Séchelles |journal= Revue Zoologique africaine |Science |volume=94 |issue=1 |pages=138-168}}</ref>

== '''Ecology''' ==

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.<ref>{{cite journal |author= Crawford, C. S |year=1992 |title= Millipedes as model detritivores |journal= Berichte des Naturwissenschaftlich-Medizinischen Verein Innsbruck |Science |volume=10 |pages=277-288}}</ref><ref>{{cite book |title= Grassland Invertebrates |publisher=Chapman and Hall, London|year=1994 |author=Curry, J. P |pages=437}}</ref><ref>{{cite journal |author= Lawrence, J. M. & Samways, M. J. |year=2003 |title= Litter breakdown by the Seychelles giant millipede and the conservation of soil process on Cousine Island, Seychelles |journal= Biological Conservation |Science |volume=113 |pages=125-132}}</ref><ref>{{cite journal |author= Ashwini, K. M. & Sridhar, K. R |year=2002 |title= Towards organic farming with the millipede Arthrosphaera magna |journal= Current Science (Bangalore) |Science |volume=82 |issue=1 |pages=20-22}}</ref>  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]].<br /><br />

Like most millipedes, Sphaerotheriids mainly inhabit the leaf litter of humid forests.  Some species, however, show an [[arboreal]] (tree-living) lifestyle <ref>{{cite journal |author= Haacker, U. & Fuchs, S |year=1972 |title= Tree climbing in Pill-Millipedes |journal= Oecologia |Science |volume=10 |pages=191-192}}</ref> and for these organisms the rolling-up reflex has been suppressed..<ref name=WP/><br />

== '''Defense against Predation''' ==

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<ref>{{cite journal |author= Herbert, D. G. |year=2000 |title= Dining on diplopods: remarkable feeding behaviour in chlamydephorid slugs (Mollusca: Gastropoda)|journal= J. Zool., Lond |Science |volume=251 |pages=1-5}}</ref> 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.<ref>{{cite journal |author= Eisner, J .C. & David, J. A |year=1967 |title=Mongoose throwing and smashing millipedes |journal= Science |Science |volume=155 |issue=3762 |pages=577-579}}</ref>  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.<ref>{{cite journal |author= Adamson, Martin L., Waerebeke, Daniel van  |year=1984 |title= Xustrostoma margarettae n. gen., n. sp. (Rhigonematidae; Nematoda) from a sphaeroteroid (order Glomerida) diplopod in Madagascar|journal= Canadian Journal of Zoology |Science |volume=62 |issue=10 |pages=2092-2096 |issn=0008-4301}}</ref><ref>{{cite journal |author= Waerebeke, Daniel van |year=1985 |title= Glomerinema ratsimamangi n.gen., n.sp. (Nematoda,Rhigonematidae) parasite de Glomeris (Diplopode) a Madagascar: description et spermiogenese|journal= Annales de Parasitologie Humaine et Comparee |Science |volume=60 |issue=1 |pages=23-32 |issn=0003-4150}}</ref><br />
<br />

== '''Giant Pill-Millipedes as Pets''' ==

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.<ref>{{cite web |name=Golden Phoenix Exotic |title=Pill Millipede Care Sheet  |url=http://www.goldenphoenixexotica.com/pmilcare.html}}</ref>  Since no captive breeding programs have been initiated, all animals found in stores are caught in the wild.<br />

== '''Taxonomy''' ==


* Family: Arthrosphaeridae<br />
:''[[Arthrosphaera]]'' Pocock, 1895, 40 species, India, Sri Lanka<br />
:''[[Zoosphaerium]]'' Pocock, 1895, 55 species, Madagascar<br />
:''[[Sphaeromimus]]'' de Saussure & Zehntner, 1902, 3 species, southeast Madagascar<br />
:''[[Microsphaerotherium]]'' Wesener & van den Spiegel, 2007, 2 species, Madagascar

* Family: Procyliosomatidae<br />
:''[[Procyliosoma]]'', Silvestri, 1917: 11 species, Eastern Australia, Tasmania, New Zealand

* Family: Zephronidae<br />
:''[[Bothrobelum]]'' Verhoeff, 1924, monotypic, Borneo<br />
:''[[Cryxus]]'' Leach, 1814, monotypic, 'Asia'<br />
:''[[Indosphaera]]'' Attems, 1935, 2 species N. India, Myanmar<br />
:''[[Kophosphaera]]'' Attems, 1935, 5 species, N. India, Nepal<br />
:''[[Leptotelopus]]'' Silvestri, 1897, monotypic, Myanmar<br />
:''[[Prionobelum]]'' Verhoeff, 1924, 8 species, Vietnam, SW China<br />
:''[[Sphaerobelum]]'' Verhoeff, 1924, 4 species, Vietnam<br />
:''[[Zephronia]]'' Gray, 1832, 37 species N. India, Myanmar, Malayan Peninsula, Java, Sumatra, Borneo<br />
:''[[Sphaeropoeus]]'' Brandt, 1833, 22 species, N. India, Myanmar, Malayan Peninsula, Java, Sumatra, Borneo<br />
:''[[Tigridosphaera]]'' Jeekel, 2000, 4 species, Malayan Peninsula<br />
:''[[Castanotherium]]'' Pocock, 1895, 50 species, Indonesian Islands, Philippines<br />
:''[[Castanotheroides]]'' Chamberlin, 1921, 3 species, Philippines<br />
:''[[Sechelliosoma]]'' Mauriès, 1980, monotypic, Seychelles<br />
:''[[Rajasphaera]]'' Attems, 1935, monotypic, Borneo

* Family: Sphaerotheriidae
<br />
:''[[Sphaerotherium]]'' Brandt, 1833, 54 species, South Africa, Zimbabwe<br />
:''[[Kylindotherium]]'' Attems, 1926, monotypic, South Africa

* Incertae Sedis<br />
:''[[Epicyliosoma]]'' Silvestri, 1917, 15 species, Australia<br />
:''[[Cynotelopus]]'' Jeekel, 1986, monotypic, southwest Australia<br />
== '''References''' ==
{{Reflist}}