Haaniella

{{Short description|Genus of stick insects}} {{Automatic taxobox |fossil_range = {{Geological range|35|0}}Eocene to present<ref name="Frontiers" /> |image = Haaniella erringtoniae - pair.JPG |image_caption = ''Haaniella erringtoniae'', pair |taxon = Haaniella |authority = Kirby, 1904 |synonyms = *''Heteropteryx'' Redtenbacher, 1906 *''Miniopteryx'' Zompro, 2004 | type_species = ''Haaniella muelleri'' | type_species_authority = (de Haan, 1842) | subdivision_ranks = Species |subdivision_ref = <ref name = "Phasmida Species File Online" /> | subdivision = {{Species list | hidden = yes | ''Haaniella aculeata''| | ''Haaniella azlini''| | ''Haaniella dehaanii''| | ''Haaniella echinata''| | ''Haaniella erringtoniae''| | ''Haaniella gintingi''| | ''Haaniella glaber''| | ''Haaniella gorochovi''| | ''Haaniella grayii''| | ''Haaniella jacobsoni''| | ''Haaniella kerincia''| | ''Haaniella mecheli''| | ''Haaniella muelleri''| | ''Haaniella parva''| | ''Haaniella rosenbergii''| | ''Haaniella saussurei''| | ''Haaniella scabra''| }} |range_map = Range of Heteropteryginae.jpg |range_map_caption = Range of ''Haaniella''<ref name = "Bank et al. 2021" /> }}

'''''Haaniella''''' is a genus of the Phasmatodea family Heteropterygidae from Southeast Asia.<ref name="Phasmida Species File Online" />

== Characteristics == The species of the genus ''Haaniella'' reach a body length of 4.5 to 16.5 cm in the female sex. The males, which are 2.5 to 9.8 cm in length, are always smaller and have a slimmer abdomen than the females, whose abdomen, especially in the middle, is wider than the rest of the body. In females, as in all members of the subfamily Heteropteryginae, the abdomen ends in a secondary ovipositor, which surrounds the actual ovipositor. The fore wings (tegmina) of the adult females only just reach the abdomen. They also end at this height in the males of most species. Only the wings of males of ''Haaniella aculeata'', ''Haaniella glaber'' and ''Haaniella mecheli'' are significantly longer and have a similar wing construction as those of ''Heteropteryx dilatata''. Their narrow fore wings cover half or part of almost the entire abdomen. The hind wings below are usually even longer. In the short-winged representatives, the also very short hind wings are completely covered by the forewings and are converted into stridulation organs, which are used for defense stridulation.<ref name = "Phasmida Species File Online "/><ref name = "Heteropteryginae Revision" /><ref name =" Phoenix 4 "/>

Characteristic of the species living on Borneo, i.e. ''Haaniella dehaanii'', ''Haaniella echinata'', ''Haaniella grayii'', ''Haaniella saussurei'' and ''Haaniella scabra'' are the strongly colored intermediate membranes in the area of the hind and middle coxae and in some cases the membranes between the abdominal sternites, especially in the youth stages. These are usually colored in a species-specific manner. Many ''Haaniella'' species can be distinguished from each other in particular by the arrangement of their spines (acanthotaxy).<ref name = "Phoenix 4" /><ref name = "Phasmiden" /><ref name = "Heteropterygidae im Terrarium" />

== Distribution area == The main distribution areas of the previously known species are Sumatra, where ten species are endemic<ref name = "Taxonomic Sumatra 1" /> and Borneo with five endemic species. Further representatives can be found on the Malay Peninsula, in Singapore and the island Simeulue. With South Vietnam the distribution area of ''Haaniella gorochovi'' is a specialty.<ref name = "Bank et al. 2021" /><ref name = "Phasmida Species File Online" /><ref name = "Heteropteryginae Revision" /> Most of the species inhabit tropical rainforests from the plains up to an altitude of 800 m. ''Haaniella scabra'', on the other hand, is a mountain species and is native to cooler altitudes between 1000 and 1800 m.<ref name = "Phasmids of Borneo" /> The location of ''Haaniella gintingi'' is also relatively high. The species was discovered by Jimmy Gideon Ginting in November 2010 in northern Sumatra near Mount Sibayak at an altitude of 1400 to 1600 m.<ref name = "bugs 2" /><ref name = "phasmatodea-kneubühler" />

== Way of life and reproduction == thumb|upright|Eggs of eight ''Haaniella'' species [[File:Haaniella males.jpg|thumb|upright|Characteristics of the thoracal spines in the males of ''H. echinata'', ''H. saussurei'' and ''H. grayii'' in comparison]] The females lay their relatively large eggs several centimeters deep in the ground using the ovipositor. With a length of up to 12 mm and a weight of almost 0.3 g, ''Haaniella echinata'' produces the largest known phasmid eggs. The eggs have a diagonal cross-shaped micropylar plate. In the lower angle of which the micropyle is located. In many species, the surface of the eggs is rough and bristled. The eggs of other species, on the other hand, are rather smooth and hairless. The nymphs hatch after 6 to 18 months, depending on the species and environmental conditions. Growing up to imago also takes 6 to 18 months, with some representatives like ''Haaniella erringtoniae'' being the species with the shortest known development time with six months to hatching and a further six months to imago. Many other ''Haaniella'' species not only take longer to develop, but also reach a significantly older age. Oskar V. Conle was able to document an age of more than five years in a female wild caught by ''Haaniella scabra''.<ref name = "Phasmiden" /><ref name = "Phasmatodea" /><ref name="Terrarien Enzyklopädie" />

== Taxonomy == Type species of the genus is ''Haaniella muelleri'', more precisely its basionym ''Phasma muelleri''.<ref name = "Phasmida Species File Online" /> The genus was founded in 1904 by William Forsell Kirby in honor of the Dutch zoologist Wilhem de Haan who had described the type species. Kirby transferred a number of already described species from the genus ''Heteropteryx'' into this genus, which became monotypical.<ref name = "Kirby 1904" /> For many decades ''Haaniella erringtoniae'', ''Haaniella glaber'', ''Haaniella mecheli'' and ''Haaniella rosenbergii'' have been used as synonyms considered to ''Haaniella muelleri''. This synonymization was canceled through an extensive publication by Frank H. Hennemann et al from 2016, in which five more species were newly described. In addition, ''Haaniella parva'' transferred to the genus ''Miniopteryx'' in 2004 was transferred back to ''Haaniella'', whereby ''Miniopteryx'' became a synonym for ''Haaniella''. A classification into three species groups has been proposed for the genus. To the ''Echinata'' species group were assigned ''Haaniella echinata'', ''Haaniella saussurei'' and ''Haaniella scabra'' from Borneo, to the ''Grayii'' species group ''Haaniella grayii'' and ''Haaniella dehaanii'', as well as ''Haaniella gorochovi'' and ''Haaniella parva'' that do not occur in Borneo. All other species, all of which do not come from Borneo, were assigned to the ''Muelleri'' species group.<ref name = "Heteropteryginae Revision" /> This classification could not be confirmed by a paper by Sarah Bank et al. published in 2021. To clarify the phylogeny of the Heteropterygidae four mitochondrial genes and three genes from the cell nucleus were examined. The result showed that the representatives of the Heteropterygini form a common clade, but the genus ''Heteropteryx'' is phylogenetically placed in the middle of several lines of species currently listed in ''Haaniella''. Following this, either ''Haaniella'' would have to be split up into several genera or withdrawn in favor of the previously described genus ''Heteropteryx''.<ref name = "Bank et al. 2021" /> In 2018, Francis Seow-Choen described ''Haaniella parva muiengae'', another subspecies to the nominate subspecies of ''Haaniella parva''. In the same work he described with ''Haaniella azlini'', a species that is very similar to ''Haaniella gintingi''.<ref name = "Taxonomic Sumatra 1" /> Further subspecies described and revalidated Seow-Choen in 2020.<ref name = "Taxonomic Sumatra 2" /> {{cladogram|caption=Relationships of the examined ''Haaniella'' and ''Heteropteryx'' species by Sarah Bank et al (2021)<ref name = "Bank et al. 2021" /> |cladogram={{clade|style=width:350px; |1={{clade |1=''Haaniella saussurei'' 50 px |2={{clade |1={{clade |1={{clade |1=''Haaniella gorochovi'' 50 px |2=''Haaniella gintingi'' 50 px }} |2={{clade |1=''Haaniella erringtoniae'' 50 px |2={{clade |1=''Heteropteryx'' sp. 'Khao Lak' |2=''Heteropteryx dilatata'' 50 px = ''Heteropteryx dilatata'' 'Kuala Boh' }} }} }} |2={{clade |1={{clade |1=''Haaniella scabra'' 50 px |2=''Haaniella grayii'' 50 px }} |2={{clade |1=''Haaniella dehaanii'' 50 px |2={{clade |1={{clade |1=''Haaniella echinata'' 'Similajau' |2=''Haaniella echinata'' 'Mulu' }} |2=''Haaniella echinata'' 'Brunei' = ''Haaniella echinata'' 'Tenom' = ''Haaniella echinata'' 'Kiansom' = ''Haaniella echinata'' 'Luyang' 50 px

}} }} }} }} }} }} }} The following species and subspecies are counted under ''Haaniella'':<ref name = "Phasmida Species File Online" /> * ''Haaniella aculeata'' Hennemann, Conle, Brock & Seow-Choen, 2016 ** ''Haaniella aculeata aculeata'' Hennemann, Conle, Brock & Seow-Choen, 2016 ** ''Haaniella aculeata longipennis'' Seow-Choen, 2020 * ''Haaniella azlini'' Seow-Choen, 2018 * ''Haaniella dehaanii'' (Westwood, 1859) (syn. = ''Heteropteryx dipsacus'' Redtenbacher, 1906) * ''Haaniella echinata'' (Redtenbacher, 1906) * ''Haaniella erringtoniae'' (Redtenbacher, 1906) * ''Haaniella gintingi'' Hennemann, Conle, Brock & Seow-Choen, 2016 * ''Haaniella glaber'' (Redtenbacher, 1906) (syn. = ''Haaniella muelleri simplex'' Günther, 1944) * ''Haaniella gorochovi'' Hennemann, Conle, Brock & Seow-Choen, 2016 * ''Haaniella grayii'' (Westwood, 1859) (syn. = ''Heteropteryx australe'' Kirby, 1896) * ''Haaniella jacobsoni'' Günther, 1944 * ''Haaniella kerincia'' Hennemann, Conle, Brock, Seow-Choen & Gorochov, 2016 * ''Haaniella mecheli'' (Redtenbacher, 1906) ** ''Haaniella mecheli mecheli'' (Redtenbacher, 1906) ** ''Haaniella mecheli macroptera'' Hennemann, Conle, Brock & Seow-Choen, 2016 * ''Haaniella muelleri'' (de Haan, 1842) * ''Haaniella parva'' Günther, 1944 ** ''Haaniella parva parva'' Günther, 1944 ** ''Haaniella parva muiengae'' Seow-Choen, 2020 * ''Haaniella rosenbergii'' (Kaup, 1871) ** ''Haaniella rosenbergii rosenbergii'' (Kaup, 1871) ** ''Haaniella rosenbergii novaeguineae'' Günther, 1930 ** ''Haaniella rosenbergii subulussalama'' Seow-Choen, 2020 * ''Haaniella saussurei'' Kirby, 1904 (syn. = ''Haaniella echidna'' Rehn, J.W.H., 1938) (syn. = ''Heteropteryx grayi'' Saussure, 1869) (syn. = ''Heteropteryx saussurei'' Redtenbacher, 1906) * ''Haaniella scabra'' (Redtenbacher, 1906)

== Human use == Some indigenous peoples on Borneo are known to eat the very large eggs of some ''Haaniella'' species. These are not only valued for their high protein content, but are also said to help against diarrhea when cooked.<ref name = "Phasmatodea" />

From the genus ''Haaniella'' there are some species in breeding by the phasmid enthusiasts. The Phasmid Study Group has seven species in its culture list, the Dutch - Belgian Phasma workgroup eight species, some of them in several origin stocks. After ''Haaniella echinata'' (PSG number 26) was introduced as the first one in 1979, the other species native to Borneo followed. So in 1984 came ''Haaniella scabra'' (PSG number 70), in 1990 ''Haaniella grayii'' (PSG number 125) and ''Haaniella dehaanii'' (PSG number 126) and finally in 1994 ''Haaniella saussurei'' (PSG number 177) in the terrariums of European terrarieaner. Also in the early 1990s, ''Haaniella erringtoniae'' was introduced from the Malay Peninsula, first called ''Haaniella muelleri''. Also ''Haaniella gintingi'', which before it was described being called as ''Haaniella'' sp. 'Sibayak' was introduced in 2011 and was successfully bred by Bruno Kneubühler. In 2014 ''Haaniella gorochovi'', first known as ''Haaniella'' sp. 'Bidoup Nui Ba' from Vietnam as the eighth species introduced to Europe and in breeding under PSG number 404.<ref name = "bugs 2" /><ref name = "Phasmatodea" /><ref name="PSG Culture List" />

The breeding of ''Haaniella'' species is generally considered to be a bit tricky due to its slow development. Although the insects accept most forage plants without any problems, they tend to have diarrhea in spring and early summer when feeding on leaves that are too fresh, which can lead to the total loss of entire breeding stocks. In addition, the long development time of the brittle and sometimes shock-sensitive eggs often leads to low hatching rates, since constant or optimal conditions for incubation must be guaranteed over a long period of time.<ref name = "Phoenix 4" /><ref name = "bugs 2" />

== Gallery == <gallery>

Haaniella dehaanii female.JPG|''Haaniella dehaanii'', fresh adult female Stick Insects (Haaniella echinata) mating (23171676042).jpg|''Haaniella echinata'', pair Haaniella sp Sibayak pair.jpg|''Haaniella gintingi'' 'Sibayak', pair Haaniella gorochovi mating.JPG|''Haaniella gorochovi'' 'Bidoup Nui Ba', pair Haaniella grayii Pärchen 2.jpg|''Haaniella grayii'', pair Haaniella sp? (36877193371).jpg|''Haaniella parva'', male Haaniella saussurei Männchen.jpg|''Haaniella saussurei'', male Haaniella scabra - pair.JPG|''Haaniella scabra'', pair </gallery>

== References == <references> <ref name="Frontiers">Robertson, J. A.; Bradler, S. & Whiting, M. F. (2018). ''[https://www.frontiersin.org/articles/10.3389/fevo.2018.00216/full Evolution of Oviposition Techniques in Stick and Leaf Insects (Phasmatodea)]'', ''Frontiers in Ecology and Evolution''</ref>

<ref name="Bank et al. 2021">Bank, S.; Buckley, T. R.; Büscher, T. H.; Bresseel, J.; Constant, J.; de Haan, M.; Dittmar, D.; Dräger, H.; Kahar, R. S.; Kang, A.; Kneubühler, B.; Langton-Myers, S. & Bradler, S. (2021). [https://onlinelibrary.wiley.com/doi/full/10.1111/syen.12472 Reconstructing the nonadaptive radiation of an ancient lineage of ground-dwelling stick insects (Phasmatodea: Heteropterygidae)], ''Systematic Entomology'', DOI: 10.1111/syen.12472</ref>

<ref name="Phasmida Species File Online">Brock, P. D.; Büscher, T. H. & Baker, E. W.: [https://phasmida.speciesfile.org/otus/852504/overview Phasmida Species File Online]. (accessdate 11 August 2025)</ref>

<ref name="Phoenix 4">Dräger, H. (Dec 2011). Gespenstschrecken der Familie Heteropterygidae Kirby, 1896 (Phasmatodea) – ein Überblick über bisher gehaltene Arten, Teil 1: Die Unterfamilie Heteropteryginae Kirby, 1896, ''ZAG Phoenix'', Nr. 4 Jahrgang 2(2), pp. 38–61, {{ISSN|2190-3476}}</ref>

<ref name="Phasmiden">Seiler, C; Bradler, S. & Koch, R. (2000). Phasmiden – Pflege und Zucht von Gespenstschrecken, Stabschrecken und Wandelnden Blättern im Terrarium. bede, Ruhmannsfelden, {{ISBN|3-933646-89-8}}</ref>

<ref name="Heteropterygidae im Terrarium">Zompro, O. (2000). Gespenstschrecken der Familie Heteropterygidae im Terrarium, ''Reptilia - Terraristik Fachmagazin'', Nr. 24, August/September 2000, Natur und Tier - Verlag, Münster</ref>

<ref name="Taxonomic Sumatra 1">Seow-Choen, F. (2018). A Taxonomic Guide to the Stick Insects of Sumatra Vol. 1, Natural History Publikations (Borneo) Sdn. Bhd., Kota Kinabalu, Sabah, Malaysia, pp. 603–650, {{ISBN|978-983-812-190-3}}</ref>

<ref name="Phasmids of Borneo">Bragg, P. E. (2001).Phasmids of Borneo, Natural History Publikations (Borneo) Sdn. Bhd., Kota Kinabalu, Sabah, Malaysia, {{ISBN|983-812-027-8}}</ref>

<ref name="bugs 2">Dräger, H. & Kneubühler, B. (2013). Nachzucht einer neuen Gespenstschrecke aus Sumatra: Haaniella sp. "Sibayak", ''Bugs - Das Wirbellosenmagazin'', Nr. 2, June/July/August 2013, Natur und Tier Verlag, Münster, pp. 50–53 {{ISSN|2195-8610}}</ref>

<ref name="phasmatodea-kneubühler">Kneubühler, B. auf [http://www.phasmatodea.com/haaniella-gintingi-sibayak Phasmatodea.com] about ''Haaniella gintingi'' 'Sibayak'</ref>

<ref name="Terrarien Enzyklopädie">Bruins, E. (2006). Illustrierte Terrarien Enzyklopädie, Dörfler Verlag, Eggolsheim, pp. 75–77, {{ISBN|978-3-89555-423-0}}</ref>

<ref name="Phasmatodea">[http://www.phasmatodea.com/taxonomy/term/181 Phasmatodea.com] von Oskar V. Conle Frank H. Hennemann, Bruno Kneubühler & Pablo Valero</ref>

<ref name="Kirby 1904">Kirby, W. F. (1904). [https://archive.org/details/insectbookpopula00howauoft A synonymic catalogue of Orthoptera. 1. Orthoptera Euplexoptera, Cursoria et Gressoria. (Forficulidae, Hemimeridae, Blattidae, Mantidae, Phasmidae)], pp. 396–400</ref>

<ref name="Taxonomic Sumatra 2">Seow-Choen, F. (2020). A Taxonomic Guide to the Stick Insects of Sumatra Vol. 2, Natural History Publikations (Borneo) Sdn. Bhd., Kota Kinabalu, Sabah, Malaysia, pp. 321–380, {{ISBN|978-983-812-198-9}}</ref>

<ref name="Heteropteryginae Revision">Hennemann, F. H.; Conle, O. V.; Brock, P. D. & Seow-Choen, F. (2016). Revision of the Oriental subfamiliy Heteropteryginae Kirby, 1896, with a re-arrangement of the family Heteropterygidae and the descriptions of five new species of Haaniella Kirby, 1904. (Phasmatodea: Areolatae: Heteropterygidae), ''Zootaxa'' 4159 (1), Magnolia Press, Auckland, New Zealand, {{ISSN|1175-5326}}</ref>

<ref name="PSG Culture List">[http://phasmidstudygroup.org/index.php/phasmids/psg-culture-list Culture List] on the Phasmid Study Group website</ref>

</references>

==External links== *{{Wikispecies inline|Haaniella|''Haaniella''}} *{{Commons category-inline|Haaniella|''Haaniella''}}

Category:Phasmatodea Category:Phasmatodea genera Category:Phasmatodea of Asia