Siphoniulus Temporal range: | |
---|---|
Two views of Siphoniulus alba head | |
Scientific classification | |
Kingdom: | |
Phylum: | |
Subphylum: | |
Class: | |
Order: | Siphoniulida Cook, 1895 |
Family: | Siphoniulidae Pocock, 1894 |
Genus: | Siphoniulus Pocock, 1894 |
Species | |
S. alba Pocock, 1894 Contents |
Siphoniulus is a poorly known genus of millipede containing only two living species: S. alba from Indonesia, and S. neotropicus from Mexico and Guatemala. An additional two fossil species are known from Cretaceous amber. Siphoniulus species are the only members of the family Siphoniulidae and order Siphoniulida, making Siphoniulida the smallest millipede order. Few specimens are known, and their classification is contentious, although most recent studies place them as basal members of the Helminthomorpha ("worm-like millipedes").
Siphoniulids are small and eyeless. [1] The head is drawn out into a conical beak, and lacks Tömösváry organs. The body is relatively long and narrow, containing up to 51 segments and reaching up to 7.5 mm (0.30 in) long, [lower-alpha 1] and 0.25 mm (0.01 in) wide. The body color ranges from pure white to tan, and has been described as resembling a nematode. [2] [3] The exoskeleton is smooth and has few setae, and ozopores (defensive glands) are lacking. The third segment is legless. Males possess a single pair of gonopods (modified copulatory legs) consisting of the anterior limbs on the seventh segment, and which are partially recessed into the body. The telson (rear-most segment) possesses small bristle-like structures called spinnerets. [2]
S. alba is only known from a single specimen collected in 1894 near Maninjau on the Indonesian island of Sumatra. S. neotropicus was discovered near the Mayan ruins of Tikal in Guatemala, and is also known from the Mexican states of Chiapas and Veracruz. [2] The extinct species S. muelleri and S. preciosus are known from Late Cretaceous Burmese amber. [4]
Siphoniulus alba (and the family Siphoniulidae) was described in 1894 by R. I. Pocock, from a female specimen collected near Maninjau on the island of Sumatra. [5] The specimen was deposited in the Zoological Museum Amsterdam, but by 1975 the specimen was missing its head and anterior segments. [3] S. neotropicus was described by Richard L. Hoffman in 1979 from two possibly immature females collected in Guatemala. [3] Males of either species were unknown until 2003, when samples of leaf litter from Mexico sorted by researchers at the Field Museum of Natural History revealed new specimens of S. neotropicus, allowing for complete anatomical descriptions and characterization of the gonopods. [2]
Simplified cladogram from Sierwald et al 2003 [2] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
| |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
The incorporation of Siphoniulida anatomy did not clearly resolve relationships of millipede orders. |
Simplified cladogram from Sierwald & Bond 2007 [6] | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Combining morphological with molecular data improved resolution, and also supported a basal helminthomorph placement of Siphoniulida. |
Siphoniulids have been classified in various, conflicting placements within the Helminthomorpha ("worm-like" millipedes) since their initial description, and their relation to the rest of millipedes is still unresolved. When first described, the family Siphoniulidae was placed in the "Suborder" Colobognatha, a group that is now recognized as a larger grouping including the orders Platydesmida, Polyzoniida, and Siphonophorida. [5] The following year, the American entomologist Orator F. Cook considered Stemiulids as "suborder Siphoniuloidea", closely related to Julidans and Spirostreptidans (a grouping termed Diplochaeata). In 1979, Hoffman placed Siphoniulida (now considered an order) as Helminthomorph incertae sedis , meaning the placement within Helminthomorphs was undetermined, due to the absence of male specimens. [7] [lower-alpha 2] With only two known species, Siphoniulida is the smallest order of millipedes, followed by Siphonocryptida with three to six species. [8] [9]
In more recent years, millipedes have been studied by cladistic and modern phylogenetic methods, yet Siphoniulida remains enigmatic. In the first cladistic study of millipedes, Enghoff could only place Siphoniulids as incertae sedis within the Helminthomorpha, but "probably... a specialized subordinate taxon within some juliform or colobognathan order". [10] However, in the first morphological study of millipede phylogeny incorporating full details of Siphoniulus anatomy, Siphoniulida did not appear closely related to the Juliformia nor Colobognatha, but rather appeared as an outgroup to all other helminthomorphs, and the internal classification of Helminthomorpha was poorly resolved and significantly differed from Enghoff's. [2] In a subsequent study combining anatomical data with DNA sequence data from other groups, Siphoniulida again appeared as basal within Helminthomorpha. [6]
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.
Chilognatha is a subclass of the class Diplopoda, which includes the vast majority of extant millipedes, about 12,000 species.
Gonopods are specialized appendages of various arthropods used in reproduction or egg-laying. In males, they facilitate the transfer of sperm from male to female during mating, and thus are a type of intromittent organ. In crustaceans and millipedes, gonopods are modified walking or swimming legs. Gonopods may be highly decorated with elaborate structures which may play roles in sperm competition, and can be used to differentiate and identify closely related species. Gonopods generally occur in one or more pairs, as opposed to the single (un-paired) reproductive organs such as the aedeagus of insects or the penis of harvestmen.
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.
Mammamia profuga is a species of cave-dwelling millipede in the family Julidae. The only known species of the genus Mammamia, it was described in 2011 from a specimen discovered in a cave in Italy.
Polydesmida is the largest order of millipedes, containing approximately 3,500 species, including all the millipedes reported to produce hydrogen cyanide (HCN).
Sphaerotheriida is an order of millipedes in the infraclass Pentazonia, sometimes known as giant pill millipedes. They inhabit Southern Africa, Madagascar, South and Southeast Asia, Australia and New Zealand. 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 or golf ball, but some species from Madagascar can even reach the size of an orange. 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. This order of millipedes is also unique in that some African species are used for medicinal purposes.
Tridontomidae is a small family of millipedes. Its members are endemic to Guatemala. These millipedes range from 22 mm to 28 mm in length. 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.
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.
Richard Lawrence Hoffman was an American zoologist known as an international expert on millipedes, and a leading authority on the natural history of Virginia and the Appalachian Mountains. He was a biology professor at Virginia's Radford College for almost thirty years, and curator of invertebrates at the Virginia Museum of Natural History for another twenty years. He co-founded the Virginia Natural History Society, described over 400 species of millipedes, and produced more than 480 scientific publications. He is commemorated in the scientific and/or common names of over 30 animal species, including the valley and ridge salamander and Hoffman's dwarf centipede.
Callipodida is an order of millipedes containing around 130 species, many characterized by crests or ridges.
Dalodesmidae is a family of millipedes in the order Polydesmida, containing at least 250 species found in the Southern Hemisphere.
Chordeumatida is a large order of millipedes containing some 1200 species with a nearly worldwide distribution. Also known as "sausage millipedes," they possess around 30 body segments behind the head as adults and reach about 25 mm (0.98 in) in length.
Branneria is a genus of small millipedes in the order Chordeumatida and the only genus in the family Branneriidae. Individuals reach about 4 mm (0.16 in) long. There are two species known, found in the southeastern USA: Branneria bonoculus is found in Arkansas and eastern Texas while B. carinata occurs from North Carolina to Florida and Mississippi. The adult B. bonoculus millipede has 28 body segments, and the adult B. carinata millipede has 26, both fewer than the 30 usually found in the order Chordeumatida. In both species, the gonopod complex in adult males includes three leg pairs rather than just the two that are usually modified into gonopods in this order.
Harold Frederick Loomis was an American botanist and myriapodologist known for his contributions to agronomy, plant pathology, and millipede taxonomy. He worked for the U.S. Department of Agriculture for over four decades, studying diseases of crop plants, and was a colleague of Orator F. Cook. He also made major contributions to the natural history of Central America and the West Indies, naming over 500 species of millipedes in total. He co-described with Cook the leggiest animal on earth: Illacme plenipes, with over 700 legs.
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.
Siphonocryptida is an order of millipedes, comprising the sole family Siphonocryptidae. With only seven described species, the Siphonocryptida is the second smallest millipede order, surpassed only by Siphoniulida, with two species.
Metopidiotrichidae is a family of millipedes in the order Chordeumatida. These millipedes range from 4 mm to 17 mm in length. Adult millipedes in this family have 32 segments, not the 30 segments usually found in this order. Adult males in this family often feature a reduced or vestigial leg pair 10 as part of the gonopod complex, in addition to the two leg pairs typically modified into gonopods in this order. There are about 9 genera and at least 70 described species in Metopidiotrichidae.
Speophilosomatidae is a family of millipedes belonging to the order Chordeumatida. These millipedes are found in Japan and range from 4 mm to 6 mm in length. Adult millipedes in this family are notable for being among the few in this order with only 26 segments instead of the 30 usually found in chordeumatidans. The adult males in this family are also notable for involving three leg pairs in the gonopod complex rather than the two pairs typically modified into gonopods in this order.
Peterjohnsiidae is a small family of millipedes belonging to the order Chordeumatida. These millipedes range from 3 mm to 8 mm in length and are found in Australia. Species in this family exhibit sexual dimorphism in segment number: adult males have 30 segments, but adult females have 32 segments. In adult males in this family, the gonopod complex involves three leg pairs rather than just the two usually modified into gonopods in this order.