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INVERTEBRATES Onychophora

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Onychophora


Abstract

This page presents a brief overview of the Onychophora, from Cambrian to Recent.

Keywords: Onychophora, velvet worm, Aysheaia, Hallucinogenia, Microdictyon, Peripatoides, Peripatus, Xenusion

Introduction

Onychophorans, of which there are several endemic species in New Zealand, share a number of characteristics with both annelids (segmented worms) and arthropods, although they are more closely related to the latter and are sometimes, as here, regarded as a subphylum of the Arthropoda. Other authors (e.g. Nielsen 2001) regard the Onychophora as a phylum.

Currently there are around 10 genera and 110 species recognized within two extant families: the Peripatidae (known from the circumtropical regions of Mexico, Central and northern South America, equatorial West Africa, and South East Asia) and the Peripatopsidae (found in Chile, South Africa, Australia including Tasmania, and New Zealand).

 
 

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Phylogeny

Fossil History

A number of fossils from the Cambrian have been described which look more or less like onychophorans. Some, such as the Middle Cambrian form Aysheaia are rather similar to living forms. Others were armored with various plates and spines which, disarticulated, contribute substantially to the "small shelly fauna." All of these Cambrian forms differed from living onychophorans in being marine.

Hallucigenia ...

Xenusion, from early Cambrian sandstones of eastern Europe. This form was also armed with spines, although they were shorter than those of Hallucigenia. Only two specimens have been found so far.

Microdictyon ...

The earliest (and the only) fossil terrestrial onychophoran was found in the Pennsylvanian deposits of external link Mazon Creek, near Chicago, a locality that has yielded a great many fossils of soft-bodied organisms. The fossil is not very different from living onychophorans (Thompson and Jones 1980).

Segmentation

"Onychophorans are thought to be the sister taxon of [eu]arthropods and are segmented. However, onychophorans lack engrailed expression in their dermis. Instead, expression is observed in the posterior half of the developing limb and in a segmental pattern in the lateral mesoderm. The limb staining suggests shared ancestry of the onychophoran and arthropod limbs. However, given the close relationship of Arthropoda and Onychophora, and their segmented body plans, the lack of segmental ectodermal expression in Onychophora suggests that the ancestral role of engrailed was not segmentation; this absence may be a consequence of evolutionary loss of skeletons. Onychophoran dermis lacks a chitinous cuticle; thus Onychophora lack an exoskeleton. Furthermore, Cambrian fossils thought to be stem group onychophorans, such as Microdictyon, Hallucinogenia, and Xenusion, bear skeletal elements above the limb on each segment. Therefore, the absence of engrailed transcription in the ectoderm of modern Onychophora could well be a consequence of evolutionary loss of exoskeletal elements..." (Jacobs et al. 2000, pp. 343-345; full text).

Systematics

Phylum Arthropoda Siebold & Stannius 1895

1895 Arthropoda Siebold & Stannius
1938 Lobopodia Snodgrass
1995 Panarthropoda Nielsen
1997 Lobopodia Snodgrass 1938, Budd
2001 Panarthropoda Nielsen, Nielsen
2001 Arthropoda, Budd

Subphylum Protarthropoda Lankester 1904

1949 Pararthropoda Vandel
1954 Oncopoda Weber

Type: Peripatus Guilding 1825

Also includes the subphyla Pentastomida and Tardigrada.

Class Onychophora Grube 1853

1853 Onychophora Grube
2001 Phylum Onychophora Nielsen

 

... describe gross morphology ...
The body itself is not segmented except for the head, which is divided into three segments. The first contains the two large antennae with an eye at the base (Jamaica and South Africa have cave dwelling species which do not have eyes). Some males also have other appendages believed to be involved in sperm transfer. The second segment contains the jaw like mouth which is used for rasping into prey and then sucking out the nutrients. The third segment holds the first pair of parapodia-like legs.

Order Protonychophora Hutchinson 1930

The Order Protonychophora includes the problematical fossil Aysheaia, first described from the Burgess Shale by Walcott.

Family Aysheaiidae Walcott 1911

Genus Aysheaia Walcott 1911

Aysheaia pedunculata Walcott 1911

This problematical fossil was first described from the Burgess Shale by Walcott in 1911.

Family Uncertain

Genus Xenusion Pompeckj 1927

Order Euonychophora Hutchinson 1930

Anatomy: Contemporary onycophorans are ceolomates and have hemocoel, which means they have a lined body cavity filled with blood, rather than a vascular system. They have a muscular tubular heart which pumps the colorless blood around the body cavity. Locomotion is essentially annelid-like, with the body cavity functioning as a hydrostatic skeleton. The parapodia-like legs are also filled with blood and a valve at the base keeps them firm and muscular coordination can extend them or retract them and make them move forward or make them move backward.
On the end of their legs they bear chitinous claws for gripping, although on smooth substrate they walk on walking pads. Onycophorans have a cuticle with a-chitin but lacking collagen, which is periodically shed to permit growth (ecdysis). New cuticle is secreted underneath the old one by the ectodermal cells which develop microvilli that are subsequently withdrawn. Ecdysteroids have been found in various tissues but their function remains unknown (Hoffmann 1997; Nielsen 2001, p. 198). Unlike insect dermis, their cuticle is non-articulated, thin and soft and covered in hundreds of papillae and sensory hairs giving them a velvety texture, hence the common name ‘velvet worm.’
Like insects the Onycophora breathe through spiracles. Spiracles open out to the environment and oxygen enters through a system of tubules (tracheae) and is absorbed into the tissues across the moist surfaces. However, unlike the insects, Onycophora have no control on the spiracles and they are always open, making the animal extremely vulnerable to desiccation, so high levels of humidity are required.
Contemporary Onycophorans are able to predate organisms several times larger than themselves by immobilizing it with a gluey secretion from glands in its head, projected up to 30cm. The secretion holds the prey while the animal approaches it, bites through the cuticle, and injects a toxic, digestive saliva into the wound.

Onychophorans themselves have few predators, except perhaps insect carnivores such as centipedes, birds and rodents.

Family Peripatidae Evans 1901

Type: Peripatus Guilding 1825

The members of this family are generally coloured red-brown and possess 22 to 43 pairs of legs.

Family Peripatopsidae Bouvier 1907

This family contains organisms which are generally coloured blue-green and possess 14 to 25 pairs of legs.

Peripatoides novaezealandiae

Description: Peripatoides novaezealandiae has 16 pairs of legs with hooks at the end, two robust feelers, and is velvety in appearance and comes in colours of blue, green, grey and brown; and may reach 80 mm in length.

Occurrence: Never abundant but "not uncommon" at Titirangi, near Auckland (Gill 1998).

Habit: The Peripatus preys on small arthropods by squirting and entrapping them in a glue fired from openings beside the mouth. It then injects saliva into the prey, dissolving the inner contents, and thus enabling the Peripatus to suck them out. Peripatus live in damp areas such as under moss, in rotting logs, behind the bark of trees, and in leaf litter.

Discussion: ...

References

Budd, Graham E. 2001: Why are Arthropods Segmented? Evolution and Development, vol. 3 (5): 332-342.

Gill, B.J. (ed.) 1998: Powell's Native Animals of New Zealand. 4th edition. Bateman. 94 pp.

Hoffmann, K. 1997: Ecdysteroids in Adult Females of a 'Walking Worm' Euperipatoides leuckartii (Onychophora, Peripatopsidae). Invert. Reprod. Dev. 32: 27-30.

Jacobs, David K.; Wray, Charles G.; Wedeen, Cathy J.; Kostriken, Richard; DeSalle, Rob; Staton, Joseph L.; Gates, Ruth D.; Lindberg, David R. 2000: Molluscan engrailed Expression, Serial Organization, and Shell Evolution. Evolution & Development, vol. 2 (6): 340-347.

Nielsen, Claus 2001: Animal Evolution. Second Edition. 563 pp. Oxford University Press.

Ramskold, L. 1992: The Second Leg Row of Hallucigenia Discovered. Lethaia 25: 221-224.

Thompson, I. and Jones, D. 1980: A Possible Onychophoran from the Middle Pennsylvanian Mazon Creek beds of Northern Illinois. Journal of Paleontology 54: 588-596.





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