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=== Species description ===
=== Species description ===
As with all Onychophora, Peripatoides novaezealandiae have a long, worm-like body, a head, and an anal cone<ref name="Hutton 1876">{{cite journal |last1=Hutton |first1=F.W. |title=On Peripatus novæ-zealandiæ |journal=Annals and Magazine of Natural History |date=November 1876 |volume=18 |issue=107 |pages=361–369 |doi=10.1080/00222937608682060}}</ref><ref name="Mayer 2015">{{Citation |last1=Mayer |first1=Georg |title=Onychophora |date=2015 |url=http://link.springer.com/10.1007/978-3-7091-1865-8_4 |work=Evolutionary Developmental Biology of Invertebrates 3 |pages=53–98 |editor-last=Wanninger |editor-first=Andreas |place=Vienna |publisher=Springer Vienna |language=en |doi=10.1007/978-3-7091-1865-8_4 |isbn=978-3-7091-1864-1 |last2=Franke |first2=Franziska Anni |last3=Treffkorn |first3=Sandra |last4=Gross |first4=Vladimir |last5=de Sena Oliveira |first5=Ivo}}</ref>. Their heads have three pairs of modified limbs: the antennae, the jaws, and the oral papillae<ref name="Hutton 1876" /><ref name="Mayer 2015" />. Their skin is velvety in texture and the sticky slime projected from their oral papillae does not stick to it<ref name="Hutton 1876" />. They respire via two rows of trachea on the dorsal surface, which alternate with the legs<ref name="Hutton 1876" />.
As with all Onychophora, Peripatoides novaezealandiae have a long, worm-like body, a head, and an anal cone<ref name="Hutton 1876">{{cite journal |last1=Hutton |first1=F.W. |title=On Peripatus novæ-zealandiæ |journal=Annals and Magazine of Natural History |date=November 1876 |volume=18 |issue=107 |pages=361–369 |doi=10.1080/00222937608682060}}</ref><ref name="Mayer 2015">{{Citation |last1=Mayer |first1=Georg |title=Onychophora |date=2015 |url=http://link.springer.com/10.1007/978-3-7091-1865-8_4 |work=Evolutionary Developmental Biology of Invertebrates 3 |pages=53–98 |editor-last=Wanninger |editor-first=Andreas |place=Vienna |publisher=Springer Vienna |language=en |doi=10.1007/978-3-7091-1865-8_4 |isbn=978-3-7091-1864-1 |last2=Franke |first2=Franziska Anni |last3=Treffkorn |first3=Sandra |last4=Gross |first4=Vladimir |last5=de Sena Oliveira |first5=Ivo}}</ref>. Their heads have three pairs of modified limbs: the antennae, the jaws, and the oral papillae<ref name="Hutton 1876" /><ref name="Mayer 2015" />. Their skin is velvety in texture and the sticky slime projected from their oral papillae does not stick to it<ref name="Hutton 1876" />. They respire via two rows of trachea on the dorsal surface, which alternate with the legs<ref name="Hutton 1876" />.
Prior to 1998, Peripatoides novaezealandiae described a species complex containing at least six morphologically similar, but genetically distinct species<ref name="Pripnow a">{{cite journal |last1=Pripnow |first1=Birgit |last2=Ruhberg |first2=Hilke |title=Peripatopsidae (Onychophora) from New Zealand - observations on selected morphs of the 'Peripatoides novaezealandiae-complex' in culture: morphological and reproductive aspects |journal=African Invertebrates |date=31 August 2003 |volume=44 |issue=1 |pages=103–114 |doi=10.5281/zenodo.7666500 |url=https://zenodo.org/records/7666500}}</ref> (Trewick, 1998). P. novaezealandiae s. str. is one of these allozymatic species (Trewick, 1998). Other described species include P. aurorbis, P. kawekaensis, P. morgani and P. sympatrica (Trewick, 2000).
Prior to 1998, Peripatoides novaezealandiae described a species complex containing at least six morphologically similar, but genetically distinct species<ref name="Pripnow a">{{cite journal |last1=Pripnow |first1=Birgit |last2=Ruhberg |first2=Hilke |title=Peripatopsidae (Onychophora) from New Zealand - observations on selected morphs of the 'Peripatoides novaezealandiae-complex' in culture: morphological and reproductive aspects |journal=African Invertebrates |date=31 August 2003 |volume=44 |issue=1 |pages=103–114 |doi=10.5281/zenodo.7666500 |url=https://zenodo.org/records/7666500}}</ref><ref name="Trewick 1998">{{Cite journal|last=Trewick|first=Steven A.|date=1998|title=Sympatric cryptic species in New Zealand Onychophora|journal=Biological Journal of the Linnean Society|language=en|volume=63|issue=3|pages=307–329|doi=10.1111/j.1095-8312.1998.tb01520.x|issn=0024-4066|doi-access=free}}</ref>. P. novaezealandiae s. str. is one of these allozymatic species<ref name="Trewick 1998" />. Other described species include P. aurorbis, P. kawekaensis, P. morgani and P. sympatrica<ref name="Trewick 2000">{{Cite journal|last=Trewick|first=S. A.|date=2000|title=Mitochondrial DNA sequences support allozyme evidence for cryptic radiation of New Zealand Peripatoides (Onychophora)|journal=Molecular Ecology|language=en|volume=9|issue=3|pages=269–281|doi=10.1046/j.1365-294x.2000.00873.x|pmid=10736025|bibcode=2000MolEc...9..269T |s2cid=8637591|issn=1365-294X}}</ref>.
All species in the P. novaezealandiae-complex have fifteen pairs of unjointed, hollow cone-like limbs called lobopods<ref name="Hutton 1876" /> (Trewick, 1998). This distinguishes them from the closely related P. suteri which has sixteen pairs of lobopods<ref name="Gleeson 1996" /><ref name="Gleeson 2010">{{cite book |last1=Gleeson |first1=D. M. |last2=Ruberg |first2=H. |editor1-last=Gordon |editor1-first=D. P. |title=New Zealand inventory of biodiversity |date=2010 |publisher=Canterbury University Press |location=Christchurch, N.Z |isbn=978-1-87725793-3 |pages=36-39 |chapter=Phylum Onychophora: velvet worms, peripatus.}}</ref>. As shown in Figure 1, P. novaezealandiae-complex have three complete spinous pads and a fourth fragmented pad on the ventral side of the legs<ref name="Pripnow a" />. They all have three distal papillae on the feet<ref name="Pripnow a" />(Trewick, 1998).
All species in the P. novaezealandiae-complex have fifteen pairs of unjointed, hollow cone-like limbs called lobopods<ref name="Hutton 1876" /><ref name="Trewick 1998" />. This distinguishes them from the closely related P. suteri which has sixteen pairs of lobopods<ref name="Gleeson 1996" /><ref name="Gleeson 2010">{{cite book |last1=Gleeson |first1=D. M. |last2=Ruberg |first2=H. |editor1-last=Gordon |editor1-first=D. P. |title=New Zealand inventory of biodiversity |date=2010 |publisher=Canterbury University Press |location=Christchurch, N.Z |isbn=978-1-87725793-3 |pages=36-39 |chapter=Phylum Onychophora: velvet worms, peripatus.}}</ref>. As shown in Figure 1, P. novaezealandiae-complex have three complete spinous pads and a fourth fragmented pad on the ventral side of the legs<ref name="Pripnow a" />. They all have three distal papillae on the feet<ref name="Pripnow a" /><ref name="Trewick 1998" />.
Figure 1. Schematic ventral sketch of the location of sole pads and distal papillae on the fifth leg of a male from P. novaezealandiae-complex (Pripnow & Ruhberg, 2003c). CC BY 4.0.
Figure 1. Schematic ventral sketch of the location of sole pads and distal papillae on the fifth leg of a male from P. novaezealandiae-complex<ref name="Pripnow c">{{cite journal |last1=Pripnow |first1=Birgit |last2=Ruhberg |first2=Hilke |title=Figs 3–5 in Peripatopsidae (Onychophora) from New Zealand - observations on selected morphs of the 'Peripatoides novaezealandiae-complex' in culture: morphological and reproductive aspects |journal=African Invertebrates |date=31 August 2003 |volume=44 |issue=1 |pages=103-114 |url=https://zenodo.org/records/7666506}}</ref>. CC BY 4.0.


Colour varies from brownish red to purplish black, but there is usually a thin black dorsal stripe<ref name="Hutton 1876" />. A bulge of orange-coloured papillae near the eyes is characteristic of the P. novaezealandiae-complex<ref name="Pripnow a" /> (see Figure 2).
Colour varies from brownish red to purplish black, but there is usually a thin black dorsal stripe<ref name="Hutton 1876" />. A bulge of orange-coloured papillae near the eyes is characteristic of the P. novaezealandiae-complex<ref name="Pripnow a" /> (see Figure 2).
Figure 2. Location of characteristic orange papillae bulge relative to eye on P. novaezealandiae-complex (Pripnow & Ruhberg, 2003b). CC BY 4.0.
Figure 2. Location of characteristic orange papillae bulge relative to eye on P. novaezealandiae-complex<ref name="Pripnow b">{{cite journal |last1=Pripnow |first1=Birgit |last2=Ruhberg |first2=Hilke |title=Fig. 2 in Peripatopsidae (Onychophora) from New Zealand - observations on selected morphs of the 'Peripatoides novaezealandiae-complex' in culture: morphological and reproductive aspects |journal=African Invertebrates |date=31 August 2003 |volume=44 |issue=1 |pages=103-114 |url=https://zenodo.org/records/7666504}}</ref>. CC BY 4.0.


Length varies from 2.5-5 cm (Trewick, 1998). Males can reach a weight of up to 200 mg while adult female weight varies more widely, with some specimens of over 800 mg (Tutt et al., 2002).
Length varies from 2.5-5 cm<ref name="Trewick 1998" />. Males can reach a weight of up to 200 mg while adult female weight varies more widely, with some specimens of over 800 mg<ref name="Tutt 2002">{{Cite journal|last1=Tutt|first1=Karen|last2=Daugherty|first2=Charles H.|last3=Gibbs|first3=George W.|date=2002|title=Differential life-history characteristics of male and female Peripatoides novaezealandiae (Onychophora: Peripatopsidae)|journal=Journal of Zoology|language=en|volume=258|issue=2|pages=257–267|doi=10.1017/S095283690200136X|issn=1469-7998|doi-access=free}}</ref>.


Morphologically, males can usually be distinguished from females by the presence of a pair of posterior orange papillae which mark the opening of the anal glands (Trewick, 1998; Tutt et al., 2002). These papillae appear before maturity, so cannot be used to identify sexually mature males (Tutt et al., 2002). Males in the P. novaezealandiae-complex do not have the pheromone producing crural glands found in many other Onychophora, such as Euperipatoides rowelli<ref name="Barclay 2000">{{cite journal |last1=Barclay |first1=S. D. |last2=Rowell |first2=D. M. |last3=Ash |first3=J. E. |title=Pheromonally mediated colonization patterns in the velvet worm Euperipatoides rowelli (Onychophora) |journal=Journal of Zoology |date=April 2000 |volume=250 |issue=4 |pages=437–446 |doi=10.1017/s0952836900004027}}</ref> (Watt, 1960).
Morphologically, males can usually be distinguished from females by the presence of a pair of posterior orange papillae which mark the opening of the anal glands<ref name="Trewick 1998" /><ref name="Tutt 2002" />. These papillae appear before maturity, so cannot be used to identify sexually mature males<ref name="Tutt 2002" />. Males in the P. novaezealandiae-complex do not have the pheromone producing crural glands found in many other Onychophora, such as Euperipatoides rowelli<ref name="Barclay 2000">{{cite journal |last1=Barclay |first1=S. D. |last2=Rowell |first2=D. M. |last3=Ash |first3=J. E. |title=Pheromonally mediated colonization patterns in the velvet worm Euperipatoides rowelli (Onychophora) |journal=Journal of Zoology |date=April 2000 |volume=250 |issue=4 |pages=437–446 |doi=10.1017/s0952836900004027}}</ref>.


Newborns of P. novaezealandiae s. str. are pure white with slightly purple antennae<ref name="Hutton 1876" />(Tutt, 1997). Other P. novaezealandiae-complex morphs are born with pigments, possibly differentiating them from P. novaezealandiae s. str.<ref name="Pripnow a" />.
Newborns of P. novaezealandiae s. str. are pure white with slightly purple antennae<ref name="Hutton 1876" /><ref name="Tutt 1997">{{cite book |last1=Tutt |first1=Karen |title=The life history and reproductive cycle of Peripatoides novaezealandiae (Onychophora: Peripatopsidae) |date=1997 |url=https://ir.wgtn.ac.nz/handle/123456789/24036 |language=en-NZ}}</ref>. Other P. novaezealandiae-complex morphs are born with pigments, possibly differentiating them from P. novaezealandiae s. str.<ref name="Pripnow a" />.


== Geographic Distribution and habitat ==
== Geographic Distribution and habitat ==
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=== New Zealand range ===
=== New Zealand range ===
Peripatoides novaezealandiae-complex are found throughout New Zealand<ref name="Hutton 1876" /><ref name="DOC 2014">{{Cite book|title=New Zealand peripatus/ngaokeoke : current knowledge, conservation and future research needs|others=New Zealand. Department of Conservation, New Zealand.|year=2014|publisher=Department of Conservation, Ōtepoti/Dunedin Office |isbn=9780478150094|oclc=994631114}}</ref><ref name="Gleeson 1996">{{cite journal |last1=Gleeson |first1=Dianne M. |title=Onychophora of New Zealand; past, present and future |journal=New Zealand Entomologist |date=January 1996 |volume=19 |issue=1 |pages=51–55 |doi=10.1080/00779962.1996.9722023}}</ref>. The strict allozyme species P. novaezealandiae s. str. are limited to Wellington, Wairarapa, and southern Hawkes Bay regions in places such as Miller reserve, Otari, Akatarawa, Waiohine, Carterton, and Pahiatua (Trewick, 1998).
Peripatoides novaezealandiae-complex are found throughout New Zealand<ref name="Hutton 1876" /><ref name="DOC 2014">{{Cite book|title=New Zealand peripatus/ngaokeoke : current knowledge, conservation and future research needs|others=New Zealand. Department of Conservation, New Zealand.|year=2014|publisher=Department of Conservation, Ōtepoti/Dunedin Office |isbn=9780478150094|oclc=994631114}}</ref><ref name="Gleeson 1996">{{cite journal |last1=Gleeson |first1=Dianne M. |title=Onychophora of New Zealand; past, present and future |journal=New Zealand Entomologist |date=January 1996 |volume=19 |issue=1 |pages=51–55 |doi=10.1080/00779962.1996.9722023}}</ref>. The strict allozyme species P. novaezealandiae s. str. are limited to Wellington, Wairarapa, and southern Hawkes Bay regions in places such as Miller reserve, Otari, Akatarawa, Waiohine, Carterton, and Pahiatua<ref name="Trewick 1998" />.


=== Habitat preferences ===
=== Habitat preferences ===
Peripatoides novaezealandiae-complex are usually found within or beneath rotting logs (Trewick, 1998, 1999; Tutt et al., 2002), though they have occasionally been discovered among leaf litter and beneath objects such as stones and rocks and in crevasses<ref name="Hutton 1876" />. A relative water content over 211% in rotting logs is a minimum for the Australian species Euperipatoides leuckartii (Scott & Rowell, 1991). Sufficient moisture is vital for all Onychophora as they cannot regulate water loss due to a lack of both a waxy cuticle and tracheal spiracles<ref name="Barclay 2000" /><ref name="Gleeson 1996" /><ref name="Hardie 1975">{{cite journal |last1=Hardie |first1=R. |title=The riddle of Peripatus |journal=Australian Natural History |date=1975 |volume=18 |issue=5 |pages=180-185 |url=https://museum-publications.australian.museum/aus-nat-hist-1975-v18-iss5/ |language=en |issn=0004-9840}}</ref>.
Peripatoides novaezealandiae-complex are usually found within or beneath rotting logs<ref name="Trewick 1998" /><ref name="Trewick 1999">{{cite journal |last1=Trewick |first1=S. A. |title=Molecular diversity of Dunedin peripatus (Onychophora: Peripatopsidae) |journal=New Zealand Journal of Zoology |date=January 1999 |volume=26 |issue=4 |pages=381–393 |doi=10.1080/03014223.1999.9518201}}</ref><ref name="Tutt 2002" />, though they have occasionally been discovered among leaf litter and beneath objects such as stones and rocks and in crevasses<ref name="Hutton 1876" />. A relative water content over 211% in rotting logs is a minimum for the Australian species Euperipatoides leuckartii<ref name="Scott 1991">{{cite journal |last1=Scott |first1=Iaw |last2=Rowell |first2=D. |title=Population Biology of Euperipatoides-Leuckartii (Onychophora, Peripatopsidae) |journal=Australian Journal of Zoology |date=1991 |volume=39 |issue=5 |pages=499 |doi=10.1071/zo9910499}}</ref>. Sufficient moisture is vital for all Onychophora as they cannot regulate water loss due to a lack of both a waxy cuticle and tracheal spiracles<ref name="Barclay 2000" /><ref name="Gleeson 1996" /><ref name="Hardie 1975">{{cite journal |last1=Hardie |first1=R. |title=The riddle of Peripatus |journal=Australian Natural History |date=1975 |volume=18 |issue=5 |pages=180-185 |url=https://museum-publications.australian.museum/aus-nat-hist-1975-v18-iss5/ |language=en |issn=0004-9840}}</ref>.


In 1989, several thousand of the P. novaezealandiae-complex were found on a property in Dunedin, living in an old kitchen dump among dry tins cans and sheets of roofing iron, and in a separate pile of bricks<ref name="Harris 1991">{{cite journal |last1=Harris |first1=A.C. |title=A large aggregation of Peripatoides novaezealandiae (Hutton, 1876) (Onychophora: Peripatopsidae) |journal=Journal of the Royal Society of New Zealand |date=December 1991 |volume=21 |issue=4 |pages=405–406 |doi=10.1080/03036758.1991.10420836}}</ref>. The existence of the P. novaezealandiae-complex in a wide range of altitudes, forest, scrub and tussock, suggests that prey availability and moisture are more important than vegetation type when determining habitat suitability (Trewick, 1999).
In 1989, several thousand of the P. novaezealandiae-complex were found on a property in Dunedin, living in an old kitchen dump among dry tins cans and sheets of roofing iron, and in a separate pile of bricks<ref name="Harris 1991">{{cite journal |last1=Harris |first1=A.C. |title=A large aggregation of Peripatoides novaezealandiae (Hutton, 1876) (Onychophora: Peripatopsidae) |journal=Journal of the Royal Society of New Zealand |date=December 1991 |volume=21 |issue=4 |pages=405–406 |doi=10.1080/03036758.1991.10420836}}</ref>. The existence of the P. novaezealandiae-complex in a wide range of altitudes, forest, scrub and tussock, suggests that prey availability and moisture are more important than vegetation type when determining habitat suitability<ref name="Trewick 1999" />.


== Life cycle/Phenology ==
== Life cycle/Phenology ==
All of P. novaezealandiae-complex use lecithotrophic viviparity to reproduce and supply nutrition to their young<ref name="Mayer 2015" />. This means embryos are surrounded by egg membranes and derive nutrition from a yolk while inside the ovary<ref name="Mayer 2015" />. Hatching and birth are simultaneous<ref name="Mayer 2015" />.
All of P. novaezealandiae-complex use lecithotrophic viviparity to reproduce and supply nutrition to their young<ref name="Mayer 2015" />. This means embryos are surrounded by egg membranes and derive nutrition from a yolk while inside the ovary<ref name="Mayer 2015" />. Hatching and birth are simultaneous<ref name="Mayer 2015" />.
Hutton<ref name="Hutton 1876" /> originally claimed that Peripatoides novaezealandiae were hermaphroditic, possibly due to confusion regarding sperm storage sacs found within the female<ref name="Hardie 1975" />. This has since been contested (Trewick, 1999) and more recent literature clearly designates individuals of P. novaezealandiae-complex as male or female<ref name="Pripnow a" /> (Tutt et al., 2002). Sex can be identified in some morphs as young as two months after birth, but sexual differentiation is complete for all members of the P. novaezealandiae-complex by five months<ref name="Pripnow a" />.
Hutton<ref name="Hutton 1876" /> originally claimed that Peripatoides novaezealandiae were hermaphroditic, possibly due to confusion regarding sperm storage sacs found within the female<ref name="Hardie 1975" />. This has since been contested<ref name="Trewick 1999" /> and more recent literature clearly designates individuals of P. novaezealandiae-complex as male or female<ref name="Pripnow a" /><ref name="Tutt 2002" />. Sex can be identified in some morphs as young as two months after birth, but sexual differentiation is complete for all members of the P. novaezealandiae-complex by five months<ref name="Pripnow a" />.


Juveniles go through three stages<ref name="Pripnow a" />:
Juveniles go through three stages<ref name="Pripnow a" />:
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* Stage C: integument fully pigmented and has no lustre. Sensory spines no longer exposed, and papillae resemble adult form. Time to reach this stage varies considerably with the location and morph, suggesting possible diagnostic differences between subspecies of the P. novaezealandiae-complex.
* Stage C: integument fully pigmented and has no lustre. Sensory spines no longer exposed, and papillae resemble adult form. Time to reach this stage varies considerably with the location and morph, suggesting possible diagnostic differences between subspecies of the P. novaezealandiae-complex.


Peripatids grow by moulting the outer cuticle when it becomes too restrictive<ref name="Hardie 1975" />. P. novaezealandiae s. str. is an iteroparous batch breeder, meaning that it produces young in discontinuous batches (Tutt, 1997).
Peripatids grow by moulting the outer cuticle when it becomes too restrictive<ref name="Hardie 1975" />. P. novaezealandiae s. str. is an iteroparous batch breeder, meaning that it produces young in discontinuous batches<ref name="Tutt 1997" />.


Males reach sexual maturity between 40-60 mg body weight, while females reach sexual maturity between 80-95 mg body weight (Tutt et al., 2002). Females with embryos are usually between 114-508 mg but can be as large as 800 mg (Tutt et al., 2002). Prior to the development of the first embryos, female paired uteri are white and thin (Tutt et al., 2002). Subsequently, the uteri are thick-walled, yellow, and baggy for all mature females (Tutt et al., 2002).
Males reach sexual maturity between 40-60 mg body weight, while females reach sexual maturity between 80-95 mg body weight<ref name="Tutt 2002" />. Females with embryos are usually between 114-508 mg but can be as large as 800 mg<ref name="Tutt 2002" />. Prior to the development of the first embryos, female paired uteri are white and thin<ref name="Tutt 2002" />. Subsequently, the uteri are thick-walled, yellow, and baggy for all mature females<ref name="Tutt 2002" />.


Females can store sperm in spermathecae (Ruhberg, 1985, as cited in Tutt et al., 2002), possibly for more than two years<ref name="Pripnow a" />. Dissections show that both males and females contain sperm in all months of the year (Tutt et al., 2002).
Females can store sperm in spermathecae<ref name="Tutt 2002" />, possibly for more than two years<ref name="Pripnow a" />. Dissections show that both males and females contain sperm in all months of the year<ref name="Tutt 2002" />.


In some Onychophora, spermatophores (capsules containing sperm) from the male attach themselves to the integument of the females<ref name="Manton 1938">{{cite journal |last1=Manton |first1=S. M. |title=Studies on the Onychophora, IV - The passage of spermatozoa into the ovary on Peripatopsis and the early developments of the ova |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |date=18 February 1938 |volume=228 |issue=556 |pages=421–441 |doi=10.1098/rstb.1938.0001}}</ref>. Sperm invade the haemolymph (body cavity), making their way to the reproductive tract<ref name="Manton 1938" />. No spermatophores or scars have been found on female P. novaezealandiae-complex, but sperm has been found within the haemolymph (Tutt et al., 2002). Evidence suggests sperm transfer in this species is dermal-haemocoelic (through skin into the body cavity), but this has yet to be confirmed (Tutt et al., 2002).
In some Onychophora, spermatophores (capsules containing sperm) from the male attach themselves to the integument of the females<ref name="Manton 1938">{{cite journal |last1=Manton |first1=S. M. |title=Studies on the Onychophora, IV - The passage of spermatozoa into the ovary on Peripatopsis and the early developments of the ova |journal=Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences |date=18 February 1938 |volume=228 |issue=556 |pages=421–441 |doi=10.1098/rstb.1938.0001}}</ref>. Sperm invade the haemolymph (body cavity), making their way to the reproductive tract<ref name="Manton 1938" />. No spermatophores or scars have been found on female P. novaezealandiae-complex, but sperm has been found within the haemolymph<ref name="Tutt 2002" />. Evidence suggests sperm transfer in this species is dermal-haemocoelic (through skin into the body cavity), but this has yet to be confirmed<ref name="Tutt 2002" />.


== Diet / Prey / Predators ==
== Diet / Prey / Predators ==
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Like others in their family, the Peripatoides novaezealandiae-complex are nocturnal predators<ref name="Hutton 1876" /><ref name="Hardie 1975" />. Reports from Australian species suggest that peripatids are sensitive to daylight and try to move back into cover when exposed<ref name="Hardie 1975" />.
Like others in their family, the Peripatoides novaezealandiae-complex are nocturnal predators<ref name="Hutton 1876" /><ref name="Hardie 1975" />. Reports from Australian species suggest that peripatids are sensitive to daylight and try to move back into cover when exposed<ref name="Hardie 1975" />.


Prey are thought to be mainly arthropods that live in rotten wood (Watt, 1960). Australian peripatids are known to feed on springtails, termites, insect larvae and nymphs and even dead adult insects and rotten meat<ref name="Hardie 1975" />. Hutton<ref name="Hutton 1876" /> sustained captive P. novaezealandiae with flies. Tutt (1997) found an abundance of centipedes in logs that also contain P. novaezealandiae, suggesting a possible predator-prey relationship.
Australian peripatids are known to feed on springtails, termites, insect larvae and nymphs and even dead adult insects and rotten meat<ref name="Hardie 1975" />. Hutton<ref name="Hutton 1876" /> sustained captive P. novaezealandiae with flies. Tutt<ref name="Tutt 1997" /> found an abundance of centipedes in logs that also contain P. novaezealandiae, suggesting a possible predator-prey relationship.


Peripatids use their oral papillae to shoot out sticky slime which thickens upon contact with the air and covers prey in a strong, net-like structure<ref name="Hutton 1876" /><ref name="Hardie 1975" />. They approach and use their jaws to puncture the cuticle of the trapped animal, injecting digestive enzymes and sucking up the liquified remains<ref name="Hutton 1876" /><ref name="Hardie 1975" /><ref name="Mayer 2015" />.
Peripatids use their oral papillae to shoot out sticky slime which thickens upon contact with the air and covers prey in a strong, net-like structure<ref name="Hutton 1876" /><ref name="Hardie 1975" />. They approach and use their jaws to puncture the cuticle of the trapped animal, injecting digestive enzymes and sucking up the liquified remains<ref name="Hutton 1876" /><ref name="Hardie 1975" /><ref name="Mayer 2015" />.


=== Predators, Parasites, and Diseases ===
=== Predators, Parasites, and Diseases ===
Globally there has been very little research on the ecology of Onychophora<ref name="Guerro 2022">{{cite journal |last1=Guerrero-Casado |first1=José |last2=Monge-Nájera |first2=Julián |title=Bibliometrics of velvet worm (Onychophora) research: geographic and historical trends over 150 years, and recommendations for future work |journal=Biologia |date=16 November 2022 |volume=78 |issue=1 |pages=109–117 |doi=10.1007/s11756-022-01221-5}}</ref>. Several species, including spiders and beetles, have been found in rotting logs along with P. novaezealandiae (Tutt, 1997). It is unknown if these species are predators, prey, or perhaps either depending on the specific interaction (Tutt, 1997). Observations on iNaturalist have shown two species eating P. novaezealandiae, a harvestman (Nuncia conjuncta ssp. conjuncta) (Figure 3) and a cave wētā (Miotopus diversus) (Figure 4).
Globally there has been very little research on the ecology of Onychophora<ref name="Guerro 2022">{{cite journal |last1=Guerrero-Casado |first1=José |last2=Monge-Nájera |first2=Julián |title=Bibliometrics of velvet worm (Onychophora) research: geographic and historical trends over 150 years, and recommendations for future work |journal=Biologia |date=16 November 2022 |volume=78 |issue=1 |pages=109–117 |doi=10.1007/s11756-022-01221-5}}</ref>. Several species, including spiders and beetles, have been found in rotting logs along with P. novaezealandiae<ref name="Tutt 1997" />. It is unknown if these species are predators, prey, or perhaps either depending on the specific interaction<ref name="Tutt 1997" />. Observations on iNaturalist have shown two species eating P. novaezealandiae, a harvestman (Nuncia conjuncta ssp. conjuncta) (Figure 3) and a cave wētā (Miotopus diversus) (Figure 4).
Figure 3. Photograph of a harvestman (Nuncia conjuncta ssp. conjuncta) eating P. novaezealandiae (Schneehagen, 2021). CC BY SA 4.0.
Figure 3. Photograph of a harvestman (Nuncia conjuncta ssp. conjuncta) eating P. novaezealandiae<ref name="Schneehagen 2021">{{cite web |last1=Schneehagen |first1=Uwe |title=Subspecies Nuncia conjuncta conjuncta |url=https://www.inaturalist.org/observations/88363312 |website=iNaturalist |language=en |date=24 July 2021}}</ref>. CC BY SA 4.0.
Figure 4. Photograph of a cave wētā (Miotopus diversus) eating P. novaezealandiae (Schneehagen, 2022). CC BY SA 4.0.
Figure 4. Photograph of a cave wētā (Miotopus diversus) eating P. novaezealandiae<ref name="Schneehagen">{{cite web |last1=Schneehagen |first1=Uwe |title=New Zealand peripatus (Peripatoides novaezealandiae) |url=https://www.inaturalist.org/observations/105486552 |website=iNaturalist |language=en |date=24 January 2022}}</ref>. CC BY SA 4.0.


External nematodes have been found behind the oral papillae of P. novaezealandiae, but there is no evidence of parasitism (Tutt, 1997). Mites have also been found on the integument, but without evidence of any feeding (Tutt, 1997). Both relationships may be better characterised as phoresy, where species are transported by the peripatus without causing harm (Tutt, 1997).
External nematodes have been found behind the oral papillae of P. novaezealandiae, but there is no evidence of parasitism<ref name="Tutt 1997" />. Mites have also been found on the integument, but without evidence of any feeding<ref name="Tutt 1997" />. Both relationships may be better characterised as phoresy, where species are transported by the peripatus without causing harm<ref name="Tutt 1997" />.


No internal parasites or damage consistent with fungal infection have been found in P. novaezealandiae (Tutt, 1997).
No internal parasites or damage consistent with fungal infection have been found in P. novaezealandiae<ref name="Tutt 1997" />.


== Other information ==
== Other information ==
=== Conservation status ===
=== Conservation status ===
P. novaezealandiae-complex are classed as ‘not threatened’ according to the 2018 New Zealand Threat Classification System (Trewick et al., 2018). In part, this is because the allozymatic species have not all been described and little is known about their distribution<ref name="DOC 2014" />.
P. novaezealandiae-complex are classed as ‘not threatened’ according to the 2018 New Zealand Threat Classification System<ref name="Trewick 2018">{{cite book |last1=Trewick |first1=S. |last2=Hitchmough |first2=R. |last3=Rolfe |first3=J. |last4=Stringer |first4=I. |title=Conservation status of New Zealand Onychophora (‘peripatus’ or velvet worm), 2018 |date=2018 |publisher=New Zealand Department of Conservation |url=https://www.doc.govt.nz/globalassets/documents/science-and-technical/nztcs26entire.pdf}}</ref>. In part, this is because the allozymatic species have not all been described and little is known about their distribution<ref name="DOC 2014" />.


=== Scientific notes ===
=== Scientific notes ===
Onychophora, including P. novaezealandiae-complex, are very difficult to rear in captivity (Tutt, 1997). A German study of P. novaezealandiae-complex had some success keeping specimens alive long enough to birth young<ref name="Pripnow a" />.
Onychophora, including P. novaezealandiae-complex, are very difficult to rear in captivity<ref name="Tutt 1997" />. A German study of P. novaezealandiae-complex had some success keeping specimens alive long enough to birth young<ref name="Pripnow a" />.


There is currently no known way to age P. novaezealandiae-complex, which makes it very difficult to compare individuals and fully understand their life history (Tutt, 1997).
There is currently no known way to age P. novaezealandiae-complex, which makes it very difficult to compare individuals and fully understand their life history<ref name="Tutt 1997" />.


=== Cultural connections ===
=== Cultural connections ===
The te reo Maori name for velvet worms is ngaokeoke which comes from the word ‘ngaoki’, to crawl (Ryan, 2007).
The te reo Maori name for velvet worms is ngaokeoke which comes from the word ‘ngaoki’, to crawl<ref>{{cite web |last1=Ryan |first1=P. |title=Story: Peripatus |url=https://teara.govt.nz/en/peripatus |website=Te Ara - the Encyclopedia of New Zealand |access-date=2024-03-17}}</ref>.


When considering a new roading project, the presence of P. novaezealandiae-complex in Caversham Valley gave the location high conservation value<ref name="DOC 2014" />. The New Zealand Transport Agency (NZTA) worked with the Department of Conservation, Dunedin City Council and Otago University to translocate any Peripatus impacted by the project, along with ongoing monitoring, management, and habitat enhancement<ref name="DOC 2014" />.
When considering a new roading project, the presence of P. novaezealandiae-complex in Caversham Valley gave the location high conservation value<ref name="DOC 2014" />. The New Zealand Transport Agency (NZTA) worked with the Department of Conservation, Dunedin City Council and Otago University to translocate any Peripatus impacted by the project, along with ongoing monitoring, management, and habitat enhancement<ref name="DOC 2014" />.

Revision as of 09:25, 26 April 2024

Anistasya/sandbox
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Onychophora
Family: Peripatopsidae
Genus: Peripatoides
Pocock, 1894
Species
See text

Peripatoides is a genus of velvet worms in the family Peripatopsidae, whose species are found in New Zealand. Like all velvet worms, these animals are nocturnal predators that spit a sticky slime to trap their prey. Species of Peripatoides have 14, 15, or 16 pairs of legs.[1]

Taxonomy

The Peripatoides novaezealandiae species complex consists of at least five reproductively isolated species described in 1998, each of which has 15 pairs of legs.[2] These species (P. aurorbis, P. kawekaensis, P. morgani, and P. sympatrica) have no morphological characters that distinguish them although they are genetically differentiated.[3][2] Peripatoides novaezealandiae and the cryptic species were considered nomina dubia by de Sena Oliveira et al. (2012) because type localities were identified in the species descriptions rather than holotype specimens. In 2014 the New Zealand Department of Conservation recognised these species.[4] de Sena Oliveira (2023) later located the specimens used in the descriptions and as such no longer considered them nomina dubia.[5] Three additional species were described in 2024.[6]

Species

The genus Peripatoides consists of the following species:[7][6]

Reproduction

This genus exhibits lecithotrophic ovoviviparity; that is, mothers in this genus produce and retain yolky eggs in their uteri.[8] The eggs are fertilized internally, and babies develop inside their mother until large enough to be born, in batches of 4–6, as colourless miniatures of the parents.[1] These live-bearing Peripatoides have dermal-haemocoelic sperm transfer – which means sperm dissolve holes in the skin of the female to enter the body (haemolymph) anywhere on the body wall of the female.[9]

References

  1. ^ a b Trewick, Steven (2019). Wild Life New Zealand. New Zealand: Hand-in-Hand Press. ISBN 978-0-473-48320-3.
  2. ^ a b Trewick, Steven A. (1998). "Sympatric cryptic species in New Zealand Onychophora". Biological Journal of the Linnean Society. 63 (3): 307–329. doi:10.1111/j.1095-8312.1998.tb01520.x. ISSN 0024-4066.
  3. ^ Trewick, S. A. (2000). "Mitochondrial DNA sequences support allozyme evidence for cryptic radiation of New Zealand Peripatoides (Onychophora)". Molecular Ecology. 9 (3): 269–281. Bibcode:2000MolEc...9..269T. doi:10.1046/j.1365-294x.2000.00873.x. ISSN 1365-294X. PMID 10736025. S2CID 8637591.
  4. ^ New Zealand peripatus/ngaokeoke : current knowledge, conservation and future research needs. New Zealand. Department of Conservation, New Zealand. Department of Conservation, Ōtepoti/Dunedin Office. 2014. ISBN 9780478150094. OCLC 994631114.{{cite book}}: CS1 maint: others (link)
  5. ^ Oliveira, Ivo de Sena (2023-11-16). "An updated world checklist of velvet worms (Onychophora) with notes on nomenclature and status of names". ZooKeys (1184): 133–260. Bibcode:2023ZooK.1184..133O. doi:10.3897/zookeys.1184.107286. ISSN 1313-2970. PMC 10680090. PMID 38023768.
  6. ^ a b Trewick, Steven A.; Koot, Emily M.; Morgan-Richards, Mary (2024). "Ngāokeoke Aotearoa: The Peripatoides Onychophora of New Zealand". Insects. 15 (4): 248. doi:10.3390/insects15040248. ISSN 2075-4450.
  7. ^ Oliveira, I. S.; Read, V. M. S. J.; Mayer, G. (2012). "A world checklist of Onychophora (velvet worms), with notes on nomenclature and status of names". ZooKeys (211): 1–70. Bibcode:2012ZooK..211....1O. doi:10.3897/zookeys.211.3463. PMC 3426840. PMID 22930648. Retrieved 16 July 2016.
  8. ^ Mayer, Georg; Franke, Franziska Anni; Treffkorn, Sandra; Gross, Vladimir; de Sena Oliveira, Ivo (2015), Wanninger, Andreas (ed.), "Onychophora", Evolutionary Developmental Biology of Invertebrates 3, Vienna: Springer Vienna, pp. 53–98, doi:10.1007/978-3-7091-1865-8_4, ISBN 978-3-7091-1864-1, retrieved 2023-02-16
  9. ^ Tutt, Karen; Daugherty, Charles H.; Gibbs, George W. (2002). "Differential life-history characteristics of male and female Peripatoides novaezealandiae (Onychophora: Peripatopsidae)". Journal of Zoology. 258 (2): 257–267. doi:10.1017/S095283690200136X. ISSN 1469-7998.

Further reading

New Peripatoides Novaezealandiae Page

Identification

Species description

As with all Onychophora, Peripatoides novaezealandiae have a long, worm-like body, a head, and an anal cone[1][2]. Their heads have three pairs of modified limbs: the antennae, the jaws, and the oral papillae[1][2]. Their skin is velvety in texture and the sticky slime projected from their oral papillae does not stick to it[1]. They respire via two rows of trachea on the dorsal surface, which alternate with the legs[1]. Prior to 1998, Peripatoides novaezealandiae described a species complex containing at least six morphologically similar, but genetically distinct species[3][4]. P. novaezealandiae s. str. is one of these allozymatic species[4]. Other described species include P. aurorbis, P. kawekaensis, P. morgani and P. sympatrica[5]. All species in the P. novaezealandiae-complex have fifteen pairs of unjointed, hollow cone-like limbs called lobopods[1][4]. This distinguishes them from the closely related P. suteri which has sixteen pairs of lobopods[6][7]. As shown in Figure 1, P. novaezealandiae-complex have three complete spinous pads and a fourth fragmented pad on the ventral side of the legs[3]. They all have three distal papillae on the feet[3][4].

Figure 1. Schematic ventral sketch of the location of sole pads and distal papillae on the fifth leg of a male from P. novaezealandiae-complex[8]. CC BY 4.0.

Colour varies from brownish red to purplish black, but there is usually a thin black dorsal stripe[1]. A bulge of orange-coloured papillae near the eyes is characteristic of the P. novaezealandiae-complex[3] (see Figure 2).

Figure 2. Location of characteristic orange papillae bulge relative to eye on P. novaezealandiae-complex[9]. CC BY 4.0.

Length varies from 2.5-5 cm[4]. Males can reach a weight of up to 200 mg while adult female weight varies more widely, with some specimens of over 800 mg[10].

Morphologically, males can usually be distinguished from females by the presence of a pair of posterior orange papillae which mark the opening of the anal glands[4][10]. These papillae appear before maturity, so cannot be used to identify sexually mature males[10]. Males in the P. novaezealandiae-complex do not have the pheromone producing crural glands found in many other Onychophora, such as Euperipatoides rowelli[11].

Newborns of P. novaezealandiae s. str. are pure white with slightly purple antennae[1][12]. Other P. novaezealandiae-complex morphs are born with pigments, possibly differentiating them from P. novaezealandiae s. str.[3].

Geographic Distribution and habitat

Natural global range

Peripatoides novaezealandiae are endemic to New Zealand[6].

New Zealand range

Peripatoides novaezealandiae-complex are found throughout New Zealand[1][13][6]. The strict allozyme species P. novaezealandiae s. str. are limited to Wellington, Wairarapa, and southern Hawkes Bay regions in places such as Miller reserve, Otari, Akatarawa, Waiohine, Carterton, and Pahiatua[4].

Habitat preferences

Peripatoides novaezealandiae-complex are usually found within or beneath rotting logs[4][14][10], though they have occasionally been discovered among leaf litter and beneath objects such as stones and rocks and in crevasses[1]. A relative water content over 211% in rotting logs is a minimum for the Australian species Euperipatoides leuckartii[15]. Sufficient moisture is vital for all Onychophora as they cannot regulate water loss due to a lack of both a waxy cuticle and tracheal spiracles[11][6][16].

In 1989, several thousand of the P. novaezealandiae-complex were found on a property in Dunedin, living in an old kitchen dump among dry tins cans and sheets of roofing iron, and in a separate pile of bricks[17]. The existence of the P. novaezealandiae-complex in a wide range of altitudes, forest, scrub and tussock, suggests that prey availability and moisture are more important than vegetation type when determining habitat suitability[14].

Life cycle/Phenology

All of P. novaezealandiae-complex use lecithotrophic viviparity to reproduce and supply nutrition to their young[2]. This means embryos are surrounded by egg membranes and derive nutrition from a yolk while inside the ovary[2]. Hatching and birth are simultaneous[2]. Hutton[1] originally claimed that Peripatoides novaezealandiae were hermaphroditic, possibly due to confusion regarding sperm storage sacs found within the female[16]. This has since been contested[14] and more recent literature clearly designates individuals of P. novaezealandiae-complex as male or female[3][10]. Sex can be identified in some morphs as young as two months after birth, but sexual differentiation is complete for all members of the P. novaezealandiae-complex by five months[3].

Juveniles go through three stages[3]:

  • Stage A: wet and shiny integument (outer tissue). Needle-like spines of sensory papillae exposed.
  • Stage B: integument becomes more strongly pigmented and loses lustre. Sensory spines still exposed to some extent. This change is up to nine days after birth.
  • Stage C: integument fully pigmented and has no lustre. Sensory spines no longer exposed, and papillae resemble adult form. Time to reach this stage varies considerably with the location and morph, suggesting possible diagnostic differences between subspecies of the P. novaezealandiae-complex.

Peripatids grow by moulting the outer cuticle when it becomes too restrictive[16]. P. novaezealandiae s. str. is an iteroparous batch breeder, meaning that it produces young in discontinuous batches[12].

Males reach sexual maturity between 40-60 mg body weight, while females reach sexual maturity between 80-95 mg body weight[10]. Females with embryos are usually between 114-508 mg but can be as large as 800 mg[10]. Prior to the development of the first embryos, female paired uteri are white and thin[10]. Subsequently, the uteri are thick-walled, yellow, and baggy for all mature females[10].

Females can store sperm in spermathecae[10], possibly for more than two years[3]. Dissections show that both males and females contain sperm in all months of the year[10].

In some Onychophora, spermatophores (capsules containing sperm) from the male attach themselves to the integument of the females[18]. Sperm invade the haemolymph (body cavity), making their way to the reproductive tract[18]. No spermatophores or scars have been found on female P. novaezealandiae-complex, but sperm has been found within the haemolymph[10]. Evidence suggests sperm transfer in this species is dermal-haemocoelic (through skin into the body cavity), but this has yet to be confirmed[10].

Diet / Prey / Predators

Diet and foraging

Like others in their family, the Peripatoides novaezealandiae-complex are nocturnal predators[1][16]. Reports from Australian species suggest that peripatids are sensitive to daylight and try to move back into cover when exposed[16].

Australian peripatids are known to feed on springtails, termites, insect larvae and nymphs and even dead adult insects and rotten meat[16]. Hutton[1] sustained captive P. novaezealandiae with flies. Tutt[12] found an abundance of centipedes in logs that also contain P. novaezealandiae, suggesting a possible predator-prey relationship.

Peripatids use their oral papillae to shoot out sticky slime which thickens upon contact with the air and covers prey in a strong, net-like structure[1][16]. They approach and use their jaws to puncture the cuticle of the trapped animal, injecting digestive enzymes and sucking up the liquified remains[1][16][2].

Predators, Parasites, and Diseases

Globally there has been very little research on the ecology of Onychophora[19]. Several species, including spiders and beetles, have been found in rotting logs along with P. novaezealandiae[12]. It is unknown if these species are predators, prey, or perhaps either depending on the specific interaction[12]. Observations on iNaturalist have shown two species eating P. novaezealandiae, a harvestman (Nuncia conjuncta ssp. conjuncta) (Figure 3) and a cave wētā (Miotopus diversus) (Figure 4).

Figure 3. Photograph of a harvestman (Nuncia conjuncta ssp. conjuncta) eating P. novaezealandiae[20]. CC BY SA 4.0.

Figure 4. Photograph of a cave wētā (Miotopus diversus) eating P. novaezealandiae[21]. CC BY SA 4.0.

External nematodes have been found behind the oral papillae of P. novaezealandiae, but there is no evidence of parasitism[12]. Mites have also been found on the integument, but without evidence of any feeding[12]. Both relationships may be better characterised as phoresy, where species are transported by the peripatus without causing harm[12].

No internal parasites or damage consistent with fungal infection have been found in P. novaezealandiae[12].

Other information

Conservation status

P. novaezealandiae-complex are classed as ‘not threatened’ according to the 2018 New Zealand Threat Classification System[22]. In part, this is because the allozymatic species have not all been described and little is known about their distribution[13].

Scientific notes

Onychophora, including P. novaezealandiae-complex, are very difficult to rear in captivity[12]. A German study of P. novaezealandiae-complex had some success keeping specimens alive long enough to birth young[3].

There is currently no known way to age P. novaezealandiae-complex, which makes it very difficult to compare individuals and fully understand their life history[12].

Cultural connections

The te reo Maori name for velvet worms is ngaokeoke which comes from the word ‘ngaoki’, to crawl[23].

When considering a new roading project, the presence of P. novaezealandiae-complex in Caversham Valley gave the location high conservation value[13]. The New Zealand Transport Agency (NZTA) worked with the Department of Conservation, Dunedin City Council and Otago University to translocate any Peripatus impacted by the project, along with ongoing monitoring, management, and habitat enhancement[13].

The discovery of ngaokeoke (P. novaezealandiae-complex) on their farm inspired Taranaki farming couple Damien and Jane Roper to form a partnership with the regional council to legally protect their 2.5 ha of native bush[24].

References

  1. ^ a b c d e f g h i j k l m n Hutton, F.W. (November 1876). "On Peripatus novæ-zealandiæ". Annals and Magazine of Natural History. 18 (107): 361–369. doi:10.1080/00222937608682060.
  2. ^ a b c d e f Mayer, Georg; Franke, Franziska Anni; Treffkorn, Sandra; Gross, Vladimir; de Sena Oliveira, Ivo (2015), Wanninger, Andreas (ed.), "Onychophora", Evolutionary Developmental Biology of Invertebrates 3, Vienna: Springer Vienna, pp. 53–98, doi:10.1007/978-3-7091-1865-8_4, ISBN 978-3-7091-1864-1
  3. ^ a b c d e f g h i j Pripnow, Birgit; Ruhberg, Hilke (31 August 2003). "Peripatopsidae (Onychophora) from New Zealand - observations on selected morphs of the 'Peripatoides novaezealandiae-complex' in culture: morphological and reproductive aspects". African Invertebrates. 44 (1): 103–114. doi:10.5281/zenodo.7666500.
  4. ^ a b c d e f g h Trewick, Steven A. (1998). "Sympatric cryptic species in New Zealand Onychophora". Biological Journal of the Linnean Society. 63 (3): 307–329. doi:10.1111/j.1095-8312.1998.tb01520.x. ISSN 0024-4066.
  5. ^ Trewick, S. A. (2000). "Mitochondrial DNA sequences support allozyme evidence for cryptic radiation of New Zealand Peripatoides (Onychophora)". Molecular Ecology. 9 (3): 269–281. Bibcode:2000MolEc...9..269T. doi:10.1046/j.1365-294x.2000.00873.x. ISSN 1365-294X. PMID 10736025. S2CID 8637591.
  6. ^ a b c d Gleeson, Dianne M. (January 1996). "Onychophora of New Zealand; past, present and future". New Zealand Entomologist. 19 (1): 51–55. doi:10.1080/00779962.1996.9722023.
  7. ^ Gleeson, D. M.; Ruberg, H. (2010). "Phylum Onychophora: velvet worms, peripatus.". In Gordon, D. P. (ed.). New Zealand inventory of biodiversity. Christchurch, N.Z: Canterbury University Press. pp. 36–39. ISBN 978-1-87725793-3.
  8. ^ Pripnow, Birgit; Ruhberg, Hilke (31 August 2003). "Figs 3–5 in Peripatopsidae (Onychophora) from New Zealand - observations on selected morphs of the 'Peripatoides novaezealandiae-complex' in culture: morphological and reproductive aspects". African Invertebrates. 44 (1): 103–114.
  9. ^ Pripnow, Birgit; Ruhberg, Hilke (31 August 2003). "Fig. 2 in Peripatopsidae (Onychophora) from New Zealand - observations on selected morphs of the 'Peripatoides novaezealandiae-complex' in culture: morphological and reproductive aspects". African Invertebrates. 44 (1): 103–114.
  10. ^ a b c d e f g h i j k l m Tutt, Karen; Daugherty, Charles H.; Gibbs, George W. (2002). "Differential life-history characteristics of male and female Peripatoides novaezealandiae (Onychophora: Peripatopsidae)". Journal of Zoology. 258 (2): 257–267. doi:10.1017/S095283690200136X. ISSN 1469-7998.
  11. ^ a b Barclay, S. D.; Rowell, D. M.; Ash, J. E. (April 2000). "Pheromonally mediated colonization patterns in the velvet worm Euperipatoides rowelli (Onychophora)". Journal of Zoology. 250 (4): 437–446. doi:10.1017/s0952836900004027.
  12. ^ a b c d e f g h i j k Tutt, Karen (1997). The life history and reproductive cycle of Peripatoides novaezealandiae (Onychophora: Peripatopsidae).
  13. ^ a b c d New Zealand peripatus/ngaokeoke : current knowledge, conservation and future research needs. New Zealand. Department of Conservation, New Zealand. Department of Conservation, Ōtepoti/Dunedin Office. 2014. ISBN 9780478150094. OCLC 994631114.{{cite book}}: CS1 maint: others (link)
  14. ^ a b c Trewick, S. A. (January 1999). "Molecular diversity of Dunedin peripatus (Onychophora: Peripatopsidae)". New Zealand Journal of Zoology. 26 (4): 381–393. doi:10.1080/03014223.1999.9518201.
  15. ^ Scott, Iaw; Rowell, D. (1991). "Population Biology of Euperipatoides-Leuckartii (Onychophora, Peripatopsidae)". Australian Journal of Zoology. 39 (5): 499. doi:10.1071/zo9910499.
  16. ^ a b c d e f g h Hardie, R. (1975). "The riddle of Peripatus". Australian Natural History. 18 (5): 180–185. ISSN 0004-9840.
  17. ^ Harris, A.C. (December 1991). "A large aggregation of Peripatoides novaezealandiae (Hutton, 1876) (Onychophora: Peripatopsidae)". Journal of the Royal Society of New Zealand. 21 (4): 405–406. doi:10.1080/03036758.1991.10420836.
  18. ^ a b Manton, S. M. (18 February 1938). "Studies on the Onychophora, IV - The passage of spermatozoa into the ovary on Peripatopsis and the early developments of the ova". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 228 (556): 421–441. doi:10.1098/rstb.1938.0001.
  19. ^ Guerrero-Casado, José; Monge-Nájera, Julián (16 November 2022). "Bibliometrics of velvet worm (Onychophora) research: geographic and historical trends over 150 years, and recommendations for future work". Biologia. 78 (1): 109–117. doi:10.1007/s11756-022-01221-5.
  20. ^ Schneehagen, Uwe (24 July 2021). "Subspecies Nuncia conjuncta conjuncta". iNaturalist.
  21. ^ Schneehagen, Uwe (24 January 2022). "New Zealand peripatus (Peripatoides novaezealandiae)". iNaturalist.
  22. ^ Trewick, S.; Hitchmough, R.; Rolfe, J.; Stringer, I. (2018). Conservation status of New Zealand Onychophora (‘peripatus’ or velvet worm), 2018 (PDF). New Zealand Department of Conservation.
  23. ^ Ryan, P. "Story: Peripatus". Te Ara - the Encyclopedia of New Zealand. Retrieved 2024-03-17.
  24. ^ Nolly, R. (29 August 2021). "Farming journey leads to learning te reo Māori". www.farmersweekly.co.nz. Farmers Weekly. Retrieved 2022-06-16.
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