Occiperipatoides
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''Occiperipatoides'' is a
monospecific genus In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...
of velvet worm containing the single species ''Occiperipatoides gilesii''. This genus is
ovoviviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a "bridging" form of reproduction between egg-laying oviparity, oviparous and live-bearing viviparity, viviparous reproduction. Ovoviviparous animals possess embryos that develo ...
and found in
Western Australia Western Australia (WA) is the westernmost state of Australia. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to the south, the Northern Territory to the north-east, and South Australia to the south-east. Western Aust ...
. The genus is part of the ancient
phylum In biology, a phylum (; : phyla) is a level of classification, or taxonomic rank, that is below Kingdom (biology), kingdom and above Class (biology), class. Traditionally, in botany the term division (taxonomy), division has been used instead ...
Onychophora Onychophora (from , , "claws"; and , , "to carry"), commonly known as velvet worms (for their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, ''Peripatus''), is a phylum of el ...
that contains soft-bodied, many-legged relatives of
arthropods Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
known commonly as velvet worms.


Characteristics

''Occiperipatoides gilesii'' has a unique combination of features that differentiate it from other members of the family
Peripatopsidae Peripatopsidae or the Southern Velvet Worms are one of two extant families of velvet worm. This family includes more than 140 described species distributed among 41 genera, but some authorities deem only 131 of these species to be valid. The Fre ...
found in Western Australia. It has 16 pairs of oncopods, unstructured appendages with a stubby appearance, that are mostly uniform in size. The skin of the velvet worm is covered with papillae that are cylindrical in shape and more elongated in comparison to closely related West Australian species in the genus '' Kumbadjena''. These papillae are covered in ribbed scales that give the skin a velvety appearance. ''O. gilesii'' also exhibits a ridge-like structure that segments the papillae at the first oncopod pair. The underside of the body in Australian Onychophorans displays repeated pairs of posteriorly situated
ventral Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
and anteriorly situated pre-ventral organs at each oncopod segment. These pairs arise during development when segmental hardenings occur in the tissue of an
embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
and serve to attach limb depressor muscles to the body in adulthood. Antenna are present and are composed of 30 antennal rings that widen to form sensory pads at their base. Colouration differs between specimens but has been described as ranging from tan to greyish-blue. ''O. gilesii'' is
sexually dimorphic Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, with the larger females ranging in length from 7-46mm and the smaller males between 5-31mm.


Glands

Males of the species possess a posterior
accessory gland This glossary of entomology describes terms used in the formal study of insect species by entomologists. A–C A synthetic chlorinated hydrocarbon insecticide Insecticides are pestic ...
. A long crural gland extending into the
body cavity A body cavity is any space or compartment, or potential space, in an animal body. Cavities accommodate organs and other structures; cavities as potential spaces contain fluid. The two largest human body cavities are the ventral body cavity, a ...
is present in both sexes. In females the crural gland is irregular in its occurrence and does not have a set position, whereas it always occurs in males and is located at the first oncopod pair. Crural glands in Peripatopsidae species play a role in emitting sex
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s. These pheromones allow velvet worms to find potential mates in the absence of sound generating/receiving structures and poorly adapted eyes. Slime glands common to Onychophoran species occur in ''O. gilesii'', where they perform important physiological functions in regards to hunting and feeding. A sticky secretion is produced in the glands that is ejected onto prey via oral papillae, trapping it for the velvet worm to consume. The slime itself is stored in reservoirs that make up a significant amount of the worm's total bodyweight (up to 11% in some Australian species).


Taxonomy

''Occiperipatoides gilesii'' was first identified and named by Baldwin Spencer in 1909 and the genus ''Occiperipatoides'' was later defined by Hilke Ruhberg as monospecific in 1985. Before this reclassification by Ruhberg, ''Occiperipatoides'' was thought to include the species ''Occiperipatoides occidentalis'' (now '' Kumbadjena occidentalis''). Recent genetic studies have shown a similar difference in genetics between the two species to that observed between genera, with 81% of gene loci being incongruent between them. The taxonomic relationships between Australian peripatopsid species are generally not well understood and the discovery of significant levels of interspecific
allozyme Alloenzymes (or also called allozymes) are variant forms of an enzyme which differ structurally but not functionally from other allozymes coded for by different alleles at the same locus. These are opposed to isozymes, which are enzymes that p ...
diversity, as exemplified by the case of the ''Occiperipatoides'' genus, as well as unusual physiological modifications in the family may warrant revision of current classifications.


Distribution

''Occiperipatoides gilesii'' is primarily found in
banksia ''Banksia'' is a genus of around 170 species of flowering plants in the family Proteaceae. These Australian wildflowers and popular garden plants are easily recognised by their characteristic flower spikes, and woody fruiting "cones" and head ...
and
eucalypt Eucalypt is any woody plant with Capsule (fruit), capsule fruiting bodies belonging to one of seven closely related genera (of the tribe Eucalypteae) found across Australia: ''Eucalyptus'', ''Corymbia'', ''Angophora'', ''Stockwellia'', ''Allosyn ...
dominated bush with relatively low rainfall in comparison to areas preferred by other members of the Peripatopsidae family. Most samples of the species have been collected in the Perth Basin, which stretches from the
Swan Coastal Plain The Swan Coastal Plain in Western Australia is the geographic feature which contains the Swan River as it travels west to the Indian Ocean. The coastal plain continues well beyond the boundaries of the Swan River and its tributaries, as a geol ...
to the Darling Scarp. Its occurrence in this area has been noted as widespread by the Environmental Protection Authority of Western Australia. The range of ''O. gilesii'' receives most of its rainfall in winter months and it was in that period that the most specimens were observed during a survey conducted in 2002.


Speciation

''Occiperipatoides gilesii'' is considered to be part of the south-west Australian biodiversity hotspot, an area populated with many varieties of
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
flora and fauna. It is, along with other West Australian Onychophora, believed to represent relictual lineages of species that lived in the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
epoch. Short range endemism in the biodiversity hotspot comes as a result of climatic change in Australia during the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
that shrunk
mesic habitat In ecology, a mesic habitat is a type of habitat with a well-balanced or moderate supply of moisture throughout the growing season (e.g., a mesic forest, temperate hardwood forest, or dry-mesic prairie). The term derives from the Greek ''mesos'' ...
s by the process of
aridification Aridification is the process of a region becoming increasingly arid, or dry. It refers to long term change, rather than seasonal variation. It is often measured as the reduction of average soil moisture content. It can be caused by reduced preci ...
, limiting ecologically diverse faunal communities to the eastern coast and south-west of the continent. This, combined with the geographical isolation of Australia and the large period of time that has elapsed since it detached from
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
, has made ''O. gilesii'' evolutionarily primitive in comparison to Onychophorans found in South America and Asia. Periods of evolutionary isolation have also made populations of Peripatopsidae in south-west Australia highly disjointed although samples of ''O. gilesii'' reflect low rates of
genetic diversity Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species. It ranges widely, from the number of species to differences within species, and can be correlated to the span of survival for a species. It is d ...
.


Threats to habitat

Despite the significant presence of ''O. gilesii'' across its range, the proximity of Perth to major populations of the species presents a threat to their health as urban encroachment destroys forest undergrowth. Debris from logging activities can increase the habitability of an area for ''O. gilesii'' in the short term. Once the initial plant matter from clearing has decayed, however, forest regrowth will not be sufficient to supply a sustainable amount of undergrowth cover for the species. Carefully planned, sustainable forestry practices are therefore recommended to preserve ''O. gilesii'' habitats. The effects of increasingly severe Australian
wildfire A wildfire, forest fire, or a bushfire is an unplanned and uncontrolled fire in an area of Combustibility and flammability, combustible vegetation. Depending on the type of vegetation present, a wildfire may be more specifically identified as a ...
seasons fueled by
climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
, that in 2019-2020 alone wiped out more than 10 million hectares of land of which 80% was native forest, are also potentially harmful to ''O. gilesii'' habitats. The impact of fire on
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s in Australia is not well understood due to a lack of scientific research and a classification rate of only about 30% among relevant taxa. However, a study conducted by the Western Australian Department of Parks and Wildlife found that no ''O. gilesii'' were collected for 17 years after undergrowth in a survey area was burned by wildfire.


Behaviour


Habits

''Occiperipatoides gilesii'', like other West Australian velvet worms, relies heavily on plant litter and forest undergrowth for protection. Surveys of Australian Onychophoran species have shown that water absorbing, decaying logs are a particularly important component of forest floor habitats. A positive correlation exists between increase in age and volume of logs and the likelihood of inhabitance by velvet worms. The health of ''O. gilesii'' populations is also closely tied to rainfall patterns as they have no physiological mechanism for the retention of moisture and are prone to
desiccation Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. The ...
. Although relatively little is known about the behaviour of ''O. gilesii'' specifically, studies conducted on other Australian Peripatopsidae show that their brains include similar "mushroom structures" to those found in arthropods that are believed to play a role in enabling complex behaviors. These behaviours include group hunting, formation of social hierarchies and intraspecific aggression.


Reproduction

Male ''O. gilesii'' lack
spermatophore A spermatophore, from Ancient Greek σπέρμα (''spérma''), meaning "seed", and -φόρος (''-phóros''), meaning "bearing", or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especiall ...
carrying papillae adjacent to the eyes, a feature common in other Australian Ochynophorans, and the method of sperm transfer that they employ is not definitively known. Females possess a large
spermatheca The spermatheca (pronounced : spermathecae ), also called ''receptaculum seminis'' (: ''receptacula seminis''), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, Oligochaeta worms and certain other in ...
and a cross shaped
gonopore A gonopore, sometimes called a gonadopore, is a genital pore in many invertebrates. Hexapods, including insects, have a single common gonopore, except mayflies, which have a pair of gonopores. More specifically, in the unmodified female, it is ...
. The purpose of the spermatheca is to store sperm before it moves through the reproductive tract and fertilizes eggs that go on to form embryotic juveniles in the
uterus The uterus (from Latin ''uterus'', : uteri or uteruses) or womb () is the hollow organ, organ in the reproductive system of most female mammals, including humans, that accommodates the embryonic development, embryonic and prenatal development, f ...
. In other Australian Peripatopsidae the sperm is admitted to the spermatheca by means of deposit of the spermatophore on the skin of the female, where it breaks through into the body cavity. Broods are often of multiple paternity as different males deposit spermatophores on the skin of the female, a process that may increase genetic diversity of species with confined ranges like ''O. gilesii''. Due to ''O. gilesiis lack of spermatophore-transferring head structures, it has been theorized that dermal insemination may instead be achieved by use of a spiked appendage, or that sperm may be deposited via the vaginal opening. Members of the Peripatopsidae family such as ''O. gilesii'' have longer female maturation times, frequently entering into
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
, but higher fecundity than tropical velvet worms. This species exhibits lecithotrophic
ovoviviparity Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop inside eggs that r ...
; that is, mothers in this species produce and retain yolky eggs in their uteri. Juveniles are born in the autumn between March and April. Juveniles have fewer papillae than adults, due to an absence of secondary and accessory papillae that develop by the time of maturation, but are born with their full complement of oncopods.


References

{{Taxonbar, from1=Q19336133, from2=Q3600956 Onychophorans of Australasia Monotypic onychophoran genera Fauna of Western Australia Endemic fauna of Australia