Sphaerotheriida

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Sphaerotheriida
Pill millipede.jpg
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Myriapoda
Class: Diplopoda
Subclass: Chilognatha
Infraclass: Pentazonia
Superorder: Oniscomorpha
Order: Sphaerotheriida
Brandt, 1833
Families
Distribution Sphaerotheriida.jpg
Synonyms

Sphaerotheriida is an order of millipedes [1] in the infraclass Pentazonia, [2] sometimes known as giant pill millipedes. They inhabit Southern Africa, Madagascar, South and Southeast Asia, Australia and New Zealand. [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 sphaerotheriidans reach a maximum size of a cherry [4] or golf ball, [5] but some species from Madagascar can even reach the size of an orange [6] (an example of island gigantism; illustration - ). 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 millipedes known to do this. [7] This order of millipedes is also unique in that some African species are used for medicinal purposes. [8]

Contents

Morphology

Sphaerotheriidans are characterized by a relatively conservative body morphology; superficially all species and genera look the same. [3] Dorsally, the head of an adult is followed by twelve body tergites (collum, thoracic shield, and ten normal tergites) and the anal shield. Ventrally, adult females possess twenty-one leg pairs (forty-two legs in total), while adult 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. Instead they depend entirely on their rolling-up behavior for protection.

A pill millipede from the Western Ghats, India Pillmillipede talakaveri.jpg
A pill millipede from the Western Ghats, India

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). Until the end of the 20th century only the largest known species in the order Glomerida rivalled the size of even the smallest known Sphaerotheriida, but in the early 21st century a much smaller Sphaerotheriid was described from Madagascar: full-grown specimens of Microsphaerotherium ivohibiensis are just the size of a pea. [9] Also on Madagascar, some giant pill millipede species exhibit island gigantism, reaching more than 9.5 cm (3.7 in) in outstretched length and a size comparable to an orange when rolled up. [6]

The orders differ in the number of tergites in adults (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 traits. [10] Millipedes in both orders develop by hemianamorphosis. [11]

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 sphaerotheriidans show the same gap as the Glomerida. [12] 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 do not need continuous muscle contractions to remain in the rolled-up position.

Distribution

Arthrosphaera bicolor (Arthrosphaeridae), named and painted by Reginald Innes Pocock Arthrosphaera bicolor.png
Arthrosphaera bicolor (Arthrosphaeridae), named and painted by Reginald Innes Pocock

In general, Sphaerotheriida have a Gondwanan distribution (the exception is Zephroniidae from southeast Asia and adjacent regions). [3] [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 (which however lacks sphaerotheriidans), New Zealand, Australia, India, and Madagascar. It is believed that the Sphaerotheriida originated in Gondwana and then diverged into its various forms.

Sphaerotheriida is divided into four families whose distributions do not overlap: Procyliosomatidae, Zephroniidae, Sphaerotheriidae and Arthrosphaeridae. The most basal family, Procyliosomatidae, lives in Australia and New Zealand. [3] [15] The Zephroniidae (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 South 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, probably inadvertently. Examples include the Sri Lankan Arthrosphaeridae species Arthrosphaera brandtii which has established a population in the Usambara Mountains, Tanzania, [16] as well as some South African Sphaerotherium species which have isolated populations in Malawi. [17] Another likely candidate is Sechelliosoma forcipatum, a small species of the southeast Asian family Zephroniidae, currently only known from a single island in the Seychelles. [18]

Ecology

Little is known about the ecology, development and life history of Sphaerotheriida, but apparently all species are detritivores, feeding on dead organic matter such as leaves and wood on the forest floor. Like earthworms, they play an important role in decomposition; by breaking down decaying organic matter they release locked up nutrients back into the soil. [19] [20] [21] [22] Such recycling is essential for plant nutrition and accordingly for the entire ecology. It is possible that giant pill millipedes rely on special bacteria in their gut, much as termites do, to exploit the nutritional value of otherwise indigestible material such as lignin.

Like most millipedes, Sphaerotheriida inhabit mainly the leaf litter of humid forests. Some species, however, show an arboreal (tree-living) lifestyle, [23] and in these the rolling-up reflex has been suppressed. [24]

Defense against predation

Giant pill millipede in rolled-up position Sphaerotheriid pill millipede (Order Procyliosomatidae) (4364601900).jpg
Giant pill millipede in rolled-up position

The rolling-up ability and tough skeletal armor of the Sphaerotheriida offer protection against some predators, but a wide variety of predators feed on them, or even specialise in them as a source of food. [25] Species that specialize in feeding on giant pill millipedes necessarily have evolved special structures or behaviors to overcome their defences. Examples include the South African snail family Chlamydephoridae which almost exclusively feeds on giant pill millipedes [26] Another example is the meerkat (Suricata suricata) which has been reported (at least in captivity) to throw rolled-up sphaerotheriids against rocks in order to break them. This behaviour however, is their way of breaking open many refractory food items, such as snails and hard-shelled eggs. [25]

Apart from the rolling-up behavior of giant pill millipedes, camouflage may be an important defense mechanism against predators that hunt by sight, such as birds.

Sphaerotheriida also are subject to internal parasitism, and several species of nematodes are obligate parasites of particular species of giant pill millipedes. [27] [28]

Taxonomy

Various sphaerotheriidans (and a beetle) illustrated in 1859 Plate VII 1859.png
Various sphaerotheriidans (and a beetle) illustrated in 1859

By 2014, over 326 species in around 34 genera [29] in approximately 20 genera had been described. [3] [30] [31]

Arthrosphaera Pocock, 1895 – 40 species, India, Sri Lanka
Zoosphaerium Pocock, 1895 – 55 species, Madagascar
Sphaeromimus de Saussure & Zehntner, 1902 – 10 species, southeast Madagascar
Microsphaerotherium Wesener & van den Spiegel, 2007 – 2 species, Madagascar
Cyliosoma Pocock, 1895 – 15 species, Australia
Cynotelopus Jeekel, 1986 – monotypic, southwest Australia
Procyliosoma Silvestri, 1917 – 11 species, Eastern Australia, Tasmania, New Zealand
Sphaerotherium Brandt, 1833 – 54 species, South Africa, Zimbabwe
Kylindotherium Attems, 1926 – monotypic, South Africa
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

Phylogeny

The first modern phylogenetic study of Sphaerotheriida (simplified below) was conducted by Wesener and VandenSpiegel in 2009, using morphological data from 36 species in 10 genera. The South African family Sphaerotheriidae was found to be sister to the Madagascar family Arthrosphaeridae. The Australian genus Procyliosoma was found to be distinct from all other genera and placed in its own family, Procyliosomatidae. [3]

Pentazonia
Glomeridesmida

Glomeridesmus

Glomerida

Glomeroides

Sphaerotheriida

Procyliosomatidae

Zephroniidae

Sechelliosoma

Sphaeropoeus

Zephronia

Castanotherium

Sphaerotheriidae

Arthrosphaeridae

Sphaeromimus

Arthrosphaera

Microsphaerotherium

Zoosphaerium

Related Research Articles

<span class="mw-page-title-main">Millipede</span> Class of arthropods

Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a result of two single segments fused together. Most millipedes have very elongated cylindrical or flattened bodies with more than 20 segments, while pill millipedes are shorter and can roll into a tight ball. Although the name "millipede" derives from the Latin for "thousand feet", no species was known to have 1,000 or more until the discovery of Eumillipes persephone, which can have over 1,300 legs. There are approximately 12,000 named species classified into 16 orders and around 140 families, making Diplopoda the largest class of myriapods, an arthropod group which also includes centipedes and other multi-legged creatures.

<span class="mw-page-title-main">Pill millipede</span> Order of millipedes

Pill millipedes are any members of two living orders of millipedes, often grouped together into a single superorder, Oniscomorpha. The name Oniscomorpha refers to the millipedes' resemblance to certain woodlice (Oniscidea), also called pillbugs or "roly-polies". However, millipedes and woodlice are not closely related ; rather, this is a case of convergent evolution.

<span class="mw-page-title-main">Sphaerotheriidae</span> Family of millipedes

Sphaerotheriidae is a family of giant pill millipedes of the class Diplopoda. Millipedes of this family are distributed in southern Africa.

<span class="mw-page-title-main">Glomerida</span> Order of millipedes

Glomerida is an order of pill-millipedes found primarily in the Northern Hemisphere. They superficially resemble pill-bugs or woodlice, and can enroll into a protective ball. They have twelve body segments, 17 to 19 pairs of legs, and males have enlarged rear legs involved in mating. The order includes about 30 genera and at least 280 species, including Glomeris marginata, the common European pill-millipede. The order contains members in Europe, South-east Asia and the Americas from California to Guatemala. Although historically considered closely related with the similar sphaerotheriidans that also enroll, some DNA evidence suggest they may be more closely related to glomeridesmidans, a poorly known order that does not enroll.

<i>Sphaeromimus</i> Genus of millipedes

Sphaeromimus, or the chirping giant pill millipedes, is a genus of giant pill millipedes endemic to southeastern Madagascar. Though described in 1902, the genus was up to 2005 known from a single male specimen, whose appearance was so unusual that the authors suspected a mislabeled giant pill-millipede from India. Their unusual and distinct morphology includes well-developed stridulation organs, probably as devices for courtship. These are the male ‘harp’ and the female ‘washboard’, which contain more stridulation ribs than in other members of the order Sphaerotheriida. They have a closer affinity with the Indian genus Arthrosphaera than to other Malagasy genera, though all belong to the Arthrosphaeridae.

Tridontomidae is a small family of millipedes. Its members are endemic to Guatemala. These millipedes range from 22 mm to 28 mm in length and are uniformly grayish in color; their legs and antennae are unusually long and slender. This family includes the remarkable species Aenigmopus alatus, in which adult males feature no gonopods. This millipede is the only species in the infraclass Helminthomorpha without gonopods.

<span class="mw-page-title-main">Zephroniidae</span> Family of millipedes

Zephroniidae is a family of giant pill millipedes in the taxonomic order Sphaerotheriida. They occur in southeast Asia from the Himalayas and China south and east to Sulawesi and to Australia, and also inhabit some Philippine islands.

<span class="mw-page-title-main">Polyxenida</span> Order of millipedes

Polyxenida is an order of millipedes readily distinguished by a unique body plan consisting of a soft, non-calcified body ornamented with tufts of bristles – traits that have inspired the common names "bristly millipedes" or "pincushion millipedes". There are at least 86 species in four families worldwide, and are the only living members of the subclass Penicillata.

<span class="mw-page-title-main">C. A. W. Jeekel</span> Dutch myriapodologist and entomologist

Casimir Albrecht Willem Jeekel (1922–2010) was a Dutch myriapodologist and entomologist known for his major contributions to the taxonomy of millipedes. His 1971 monograph Nomenclator Generum et Familiarum Diplopodorum is credited as launching the "modern era" of millipede taxonomy, and has been considered the "most important single work ever published on the Diplopoda". He served as director of the Zoological Museum Amsterdam, and authored over 150 works on the taxonomy of millipedes and other myriapods.

<span class="mw-page-title-main">Arthrosphaeridae</span> Family of millipedes

Arthrosphaeridae is a family of giant pill millipedes that are found in Madagascar, the Western Ghats of India, and Sri Lanka. The largest pill-millipede in the world, Zoosphaerium neptunus belongs to this family and is known to swarm at certain times of the year.

<span class="mw-page-title-main">Zephronia</span> Genus of millipedes

Zephronia is a genus of giant pill millipedes in the family Zephroniidae. This genus includes 47 species distributed throughout N. India, Myanmar, Thailand, Malayan Peninsula, Java, Sumatra and Borneo.

<i>Zoosphaerium neptunus</i> Species of millipede

Zoosphaerium neptunus, also known as the giant emerald pill millipede, is a species of millipede within the family Arthrosphaeridae. It is the largest known pill-millipede in the world, with some individuals reaching lengths of 90 mm long. The species is endemic to Madagascar, where it is known to swarm at certain times of the year.

Arthrosphaera attemsi, is a species of pill millipedes in the family Arthrosphaeridae. It is native to India and Sri Lanka.

The Tanzanian Pill Bug Millipede,, is a species of pill millipede in the family Arthrosphaeridae. It is found in many African and Asian countries including India and Sri Lanka. Mature individuals of the species reach 3–4 cm in length. Adults are pale brown in colour with black lines in between the segments, whereas juveniles are dark brown. The species is one of the most commonly kept giant pill millipede species within the exotic pet keeping hobby. An introduced population of Arthrosphaera brandtii exists on the Usambara Mountains of Tanzania.

Arthrosphaera ruginosa, is a species of pill millipedes in the family Arthrosphaeridae. It is endemic to Sri Lanka.

Pygmaeosomatidae is a family of millipedes belonging to the order Chordeumatida. Adult millipedes in this family have 30 or 32 segments.

Trachygonidae is a family of millipedes belonging to the order Chordeumatida. Adult millipedes in this family have 28 or 30 segments.

Glomeridellidae is a family of millipedes belonging to the order Glomerida. The adult males in this family have two pairs of telopods rather than the single pair found in other families in this order.

<i>Zoosphaerium darthvaderi</i> Species of millipede

Zoosphaerium darthvaderi, also known as the Darth Vader giant pill millipede, is a species of giant pill millipede endemic to the island state of Madagascar. It is named after the Star Wars character Darth Vader. The millipede is restricted to the Ambohitantely Reserve and is considered an extremely endangered species.

<i>Zoosphaerium</i> Genus of myriapods

Zoosphaerium is a genus of giant pill millipedes endemic to Madagascar. Some species within this genus express island gigantism. The largest of the known species of giant pill millipede known is Zoosphaerium neptunus, which is capable of growing to lengths of 90 mm.

References

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