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Hemiandrus

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Hemiandrus
Hemiandrus pallitarsis
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Orthoptera
Suborder: Ensifera
Family: Anostostomatidae
Genus: Hemiandrus
Ander, 1938
Species

See text

Synonyms

Zealandosandrus Salmon, 1950

Hemiandrus electra male

Hemiandrus is a genus of wētā in the family Anostostomatidae. In New Zealand they are known as ground wētā due to their burrowing lifestyle.[1][2] Hemiandrus wētā are nocturnal,[3] and reside in these burrows during the day.[4] Ground wētā seal the entrance of their burrow during the day with a soil plug or door so that their burrow is concealed.[4] Ground wētā adults are smaller than other types of wētā, with the unusual trait of having either long or short ovipositors, depending on the species. The name of this genus comes from this trait[5] as hemi- mean half and -andrus means male, as the species where the female has a short ovipositor can sometimes be mistaken for a male. This genus has a diverse diet, depending on the species.

Taxonomy

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The genus Hemiandrus was originally described by Kjell Ander in 1938.[6] In 2024 a new genus was created (Anderus) reducing Hemiandrus to eleven described species.[7] Hemiandrus is one of the most species rich genera of the New Zealand Anostostomatidae, and sister to two Australian genera (Penalva and the winged Exogryllacis).[6][7] Ten undescribed species of this genus are said to be in Australia,[6][8] however molecular genetic analyses show that these taxa represent a separate lineage,[7] therefore their Australasian distribution is under debate.[9]

Morphology

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The genus Hemiandrus include the smallest wētā species, with adult individual body size no more than 7 millimetres,[8] and weighing less than a gram.[10] The largest Hemiandrus species has a body length of almost 30mm.[11] These wētā have no typical tympanum, and instead are able to detect sound through their cuticle, which is adaptive for their underground lifestyle.[2] Some species in this genus have unusually short ovipositors for orthopterans, the species with this morphological trait exhibit maternal care,[4] which is uncommon as it occurs only in five families within the order Orthoptera.[12]

Sexual Dimorphism

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The genus Hemiandrus shows sexual dimorphism between male and females in size, behaviour, and potentially abundance.[10] Females’ pronotum and femur are significantly larger than males, but males have a larger pronotum to femur ratio. Abundance has been observed to be higher in males, but has been taken from a single, small dataset. Higher male abundance would coincide with other species in the family Anostostomatidae, and can be reflected in their post-copulation traditions of mate-guarding from males. [13]

Diet

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The diet of these wētā depends on the species, H. maia is omnivorous, eating fruit and invertebrates.[4][14] Other ground wētā species however have been shown to be mainly herbivorous, eating apricots and various grass species.[15]

Distribution

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Diversification

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Wētā in New Zealand are made up of three major groups: Hemideina & Deinacrida (tree-giant wētā), Hemiandrus & Anderus (Ground wētā), Anisoura & Motuweta (tusked wētā). [18] Each genus in New Zealand is sister to genera in Australia and New Caledonia.[18][7] Molecular dating evidence supports divergence of Anostostomatidae occurring at the Gondwanan breakup, with the species later dispersing across the Tasman sea. [18]

Behaviour

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Several Hemiandrus species have been recorded stridulating, despite lacking the tympanum 'ears' present in other wētā genera. The sounds produced are inaudible to humans and is detected through the cuticle.[2][19][20] Many of the 'short ovipositor' species have been found to exhibit maternal care with females laying eggs in their burrows and remaining with the eggs and even nymphs while they develop.[19][4][13]

Courtship Behaviour

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Hemiandrus perform a local courtship behaviour called substrate drumming. [2] A wanting mate will drum its abdomen on a substrate, until they get reciprocating drumming or move to a new location. [13] Wētā of the genus Hemiandrus have tympanal organs in their foreleg tibia, made up of four sensory components (subgenual organ, intermediate organ, crista acustica, and accessory organ) contained within the tympanal membrane. [2] There are 90 scolopidial sensilla in all four legs. These organs are designed to detect cuticular vibrations through the ground, as ground wētā live in underground burrows, the use of a substrate-vibration communication system is necessary.

Copulation

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In Hemiandrus, copulation differs from other wētā in the family Anostostomatidae, with the presence of the female’s secondary copulatory structure. Hemiandrus assume typical ensiferan copulation position, with the female mounted above the male. The male then attaches the dorsal part of his genitalia to the female’s secondary copulatory structure on the underside of the female’s mid-abdomen. [13] Without detaching, the male stretches the ventral part of his genitalia to the female’s primary genitalia, depositing a spermatophore. The male then deposits a spermatophylax onto her abdomen, in front of the secondary structure. Once the spermatophore is deposited, the male detaches from the female, she bends her head toward her abdomen and grasps the spermatophylax. [21] A larger secondary copulatory structure on a female has positive correlation with a larger clutch of eggs, suggesting that these copulatory structures evolved via sexual selection to acquire nuptial gifts from males. [22]

Post-mating behaviour

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The male stays near the female to actively mate guard until she has consumed the entirety of the spermatophylax.[13] The spermatophylax has nutritional value and takes approximately an hour to eat, providing food for the female and ample time for sperm transfer to occur. The larger the spermatophylax is, the longer it takes for the female to eat, and therefore gives more time for mate guarding. This incentivises large investment into the production of the spermatophylax from the male to increase the chances of successful copulation. It is therefore predicted that a larger spermatophylax is a result of sexual selection. [22]

Hemiandrus in New Zealand

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There are approximately 4 undescribed New Zealand species of this genus,[23] with 12 described species.[8][24][11] Ground wētā are eaten by native species such as the wattle birds,[25] kiwi,[26] but also by introduced species such as mice,[27] stoats,[28] cats,[29] and hedgehogs.[30] Of the described species in this genus, half have a restricted range, but most are common and widespread, and are not thought to be endangered (as per the New Zealand Department of Conservation).[23]

Species

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Species of the genus Hemiandrus can be distinguished from similar ground wētā in the genus Anderus by examination of palps

Previously in the genus Hemiandrus:[7]

Possibly also:

  • Hemiandrus monstrosus Salmon, 1950 (listed as valid in the Orthoptera Species File,[31] as a synonym by other secondary sources[6])

References

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  1. ^ Gwynne, D. T. (2014). "A secondary copulatory structure in a female insect: a clasp for a nuptial meal?". Naturwissenschaften. 89 (3): 125–127. doi:10.1007/s00114-002-0298-y. PMID 12046632. S2CID 28093747.
  2. ^ a b c d e Strauß, J.; Lomas, K.; Field, L. H. (2017). "The complex tibial organ of the New Zealand ground weta: sensory adaptations for vibrational signal detection". Scientific Reports. 7 (1): 2031. Bibcode:2017NatSR...7.2031S. doi:10.1038/s41598-017-02132-1. PMC 5435688. PMID 28515484.
  3. ^ Chappell, E. M.; Trewick, S. A.; Morgan-Richards, M. (2012). "Shape and sound reveal genetic cohesion not speciation in the New Zealand orthopteran, Hemiandrus pallitarsis, despite high mitochondrial DNA divergence". Biological Journal of the Linnean Society. 105 (1): 169–186. doi:10.1111/j.1095-8312.2011.01777.x.
  4. ^ a b c d e f Taylor Smith, B. L.; Morgan-Richards, M.; Trewick, S. A. (2013). "New Zealand ground wētā (Anostostomatidae: ): descriptions of two species with notes on their biology". New Zealand Journal of Zoology. 40 (4): 314–329. doi:10.1080/03014223.2013.804422.
  5. ^ Ander, K (1938). "Diagnosen neuer Laubheuschrecken". Opuscula Entomologica. 3: 50–56.
  6. ^ a b c d Johns, P. M. (1997). "The Gondwanaland Weta: Family Anostostomatidae (Formerly in Stenopelmatidae, Henicidae or Mimnermidae): Nomenclatural Problems, World Checklist, New Genera and Species". Journal of Orthoptera Research (6): 125–138. doi:10.2307/3503546. JSTOR 3503546.
  7. ^ a b c d e f Trewick, Steven A.; Taylor-Smith, Briar L.; Morgan-Richards, Mary (2024). "Wētā Aotearoa—Polyphyly of the New Zealand Anostostomatidae (Insecta: Orthoptera)". Insects. 15 (10): 787. doi:10.3390/insects15100787. ISSN 2075-4450. PMC 11508991.
  8. ^ a b c d Taylor-Smith, B. L.; Trewick, S. A.; Morgan-Richards, M. (2016). "Three new ground wētā species and a redescription of". New Zealand Journal of Zoology. 43 (4): 363–383. doi:10.1080/03014223.2016.1205109. S2CID 88565199.
  9. ^ a b Pratt, R. C; Morgan-Richards, M.; Trewick, S. A (2008). "Diversification of New Zealand weta (Orthoptera: Ensifera: Anostostomatidae) and their relationships in Australasia". Philosophical Transactions of the Royal Society B: Biological Sciences. 363 (1508): 3427–3437. doi:10.1098/rstb.2008.0112. PMC 2607373. PMID 18782727.
  10. ^ a b c Chappell, E. M.; Webb, D. S.; Tonkin, J. D. (2014). "Notes on sexual size dimorphism, sex ratio and movements of adult ground weta (Walker) (Orthoptera: Anostostomatidae)". New Zealand Entomologist. 37 (2): 83–92. doi:10.1080/00779962.2013.856377. S2CID 4814697.
  11. ^ a b c Trewick, Steven A. (2021-03-12). "A new species of large Hemiandrus ground wētā (Orthoptera: Anostostomatidae) from North Island, New Zealand". Zootaxa. 4942 (2): 207–218. doi:10.11646/ZOOTAXA.4942.2.4. PMID 33757066.
  12. ^ Gwynne, D. T. (1995). "Phylogeny of the Ensifera (Orthoptera): A Hypothesis Supporting Multiple Origins of Acoustical Signalling, Complex Spermatophores and Maternal Care in Crickets, Katydids, and Weta". Journal of Orthoptera Research (4): 203–218. doi:10.2307/3503478. JSTOR 3503478.
  13. ^ a b c d e Gwynne, Darryl T. (2004). "Reproductive Behavior of Ground Weta (Orthoptera: Anostostomatidae): Drumming Behavior, Nuptial Feeding, Post-copulatory Guarding and Maternal Care". Journal of the Kansas Entomological Society. 77 (4): 414–428. doi:10.2317/E-34.1. ISSN 0022-8567.
  14. ^ Morgan-Richards, M (2008). "When is it coevolution? The case of ground wētā and fleshy fruits in New Zealand". New Zealand Journal of Ecology. Retrieved 2018-10-17.
  15. ^ Basri, W. M. (1978). The biology and economic impact of the weta, Hemiandrus sp. (Orthoptera : Stenopelmatidae) in an apricot orchard, Horotane Valley (Thesis). Christchurch, New Zealand: Lincoln College, University of Canterbury. hdl:10182/3201.
  16. ^ Johns, P.M. 1997: The Gondwanaland weta: family Anostostomatidae (formerly in Stenopelmatidae, Henicidae or Mimnermidae): nomenclatural problems, world checklist, new genera and species. Journal of Orthoptera research, (6): 125-138.
  17. ^ Monteith, G.B.; Field, L.H. 2001: Australian king crickets: distribution, habitats and biology (Orthoptera: Anostostomatidae). Pp. 79-94 in Field, L.H. (ed.) The biology of wetas, king crickets and their allies. CABI publishing.
  18. ^ a b c Pratt, R. C; Morgan-Richards, M; Trewick, S. A (2008). "Diversification of New Zealand weta (orthoptera: ensifera: anostostomatidae) and their relationships in Australasia". Philosophical Transactions of the Royal Society B: Biological Sciences. 363 (1508): 3427–3437. doi:10.1098/rstb.2008.0112. PMC 2607373.
  19. ^ a b Frans, Van Wyngaarden (1995). "The ecology of the Tekapo ground wētā (Hemiandrus new sp., Orthoptera: Anostostomatidae) and recommendations for the conservation of a threatened close relative". University of Canterbury Research Repository. doi:10.26021/6568.
  20. ^ "Meet Bruce, he's a musician". New Zealand Geographic. Retrieved 2023-05-02.
  21. ^ Gwynne, D.T. (2002). "A secondary copulatory structure in a female insect: a clasp for a nuptial meal?". Naturwissenschaften. 89 (3): 125–127. doi:10.1007/s00114-002-0298-y.
  22. ^ a b Gwynne, D. T. (2005). "The secondary copulatory organ in female ground weta (Hemiandrus pallitarsis, Orthoptera: Anostostomatidae): a sexually selected device in females?". Journal of the Linnean Society. 85 (4): 463–469. doi:10.1111/j.1095-8312.2005.00510.x.
  23. ^ a b Johns, P. M. (2001). Distribution and conservation status of ground weta, Hemiandrus species (Orthoptera: Anastostomatidae) (PDF). Wellington, New Zealand: Department of Conservation. Retrieved 12 June 2018.
  24. ^ Trewick, Steven A.; Taylor-Smith, Briar; Morgan-Richards, Mary (2020). "Ecology and systematics of the wine wētā and allied species, with description of four new Hemiandrus species". New Zealand Journal of Zoology. 48: 47–80. doi:10.1080/03014223.2020.1790396. ISSN 0301-4223.
  25. ^ Pierre, J. P. (2000). "Foraging behaviour and diet of a reintroduced population of the South Island Saddleback (Philesturnus carunculatus carunculatus)" (PDF). Notornis. 47: 7–12.
  26. ^ Colbourne, R.; Baird, K.; Jolly, J. (1990). "Relationship between invertebrates eaten by little spotted kiwi, , and their availability on Kapiti Island, New Zealand". New Zealand Journal of Zoology. 17 (4): 533–542. doi:10.1080/03014223.1990.10422951.
  27. ^ Wilson, D. J.; Lee, W. G. (2010). "Primary and secondary resource pulses in an alpine ecosystem: snow tussock grass ( spp.) flowering and house mouse ( ) populations in New Zealand". Wildlife Research. 37 (2): 89. doi:10.1071/WR09118.
  28. ^ Smith, D. H. V.; Jamieson, I. G.; Peach, R. M. E. (2005). "Importance of ground weta (Hemiandrus spp.) in stoat (Mustela erminea) diet in small montane valleys and alpine grasslands". New Zealand Journal of Ecology. 29 (2): 207–214. JSTOR 24058176.
  29. ^ Pascoe, Amelia (1995). "The effects of vegetation removal on rabbits (Oryctolagus cuniculus) and small mammalian predators in braided riverbeds of the Mackenzie Basin" (PDF). University of Otago.
  30. ^ Jones, C.; Moss, K.; Sanders, M. (2005). "Diet of hedgehogs (Erinaceus europaeus) in the upper Waitaki Basin, New Zealand: Implications for conservation". New Zealand Journal of Ecology. 29 (1): 29–35. JSTOR 24056190.
  31. ^ "Species Hemiandrus monstrosus Salmon, 1950: Orthoptera Species File".
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