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  • 2 Am Freischütz 16, 47058 Duisburg, Germany.; Email:
  • 3 The Natural History Museum, Cromwell Road, London, SW7 5BD, U.K.; Email:
  • 4 Lee Kong Chian Natural History Museum, Singapore.; Email:
Zootaxa, 2016 Sep 01;4159(1):1-219.
PMID: 27615907 DOI: 10.11646/zootaxa.4159.1.1


The areolate Oriental family Heteropterygidae Kirby, 1893 is critically reviewed and the results of the present study contradict the arrangement suggested by Zompro (2004), but in most aspects agree with a molecular study presented by Whiting et al (2003) and a phylogenetic study presented by Bradler (2009). The family is critically discussed and new hypotheses are presented for the phylogeny and intra-familiar relationships, placing the subfamily Dataminae Rehn & Rehn, 1939 as the basalmost clade of Heteropterygidae. The subfamilies Obriminae Brunner v. Wattenwyl, 1893 and Heteropteryginae Kirby, 1893 together represent the sister-group of Dataminae. Arguments and a tree are presented to support this hypothesis. New diagnoses and lists of genera are provided for all three subfamilies contained in Heteropterygidae, along with keys to distinguish between them.        The subfamily Obriminae is critically reviewed and the distinction between the three tribes Obrimini Brunner v. Wattenwyl, 1893, Eubulidini Zompro, 2004 and Miroceramiini Zompro, 2004 introduced by Zompro (2004) is shown to be poorly supported. While Obrimini sensu Zompro, 2004 is generally accepted (but now also contains genera that were placed in Eubulidini or Miroceramiini by Zompro (2004)), the tribes Eubulidini and Miroceramiini are not supported. A new arrangement is introduced, which is based on morphological characters neglected or overlooked by Zompro (2004) but were partly discussed by Bradler (2009). The genus Mearnsiana Rehn & Rehn, 1939 is removed from Miroceramiini and transferred to Obrimini. The genera Eubulides Stål, 1877, Heterocopus Redtenbacher, 1906, Theramenes Stål, 1875 and Stenobrimus Redtenbacher, 1906 are removed from Eubulidini and also transferred to Obrimini. Consequently, Eubulidini is synonymised with Obrimini (n. syn.). Miroceramiini is a monotypical tribe and only includes the Wallacean genus Miroceramia Günther, 1934. The new tribe Tisamenini n. trib. is established for the three basal genera Tisamenus Stål, 1875, Ilocano Rehn & Rehn, 1939 and Hoploclonia Stål, 1875 all of which were placed in Eubulidini by Zompro (2004). The latter genus differs from the other two genera by the morphology of the female genitalia, which is unique amongst the entire family. Three generic groups are recognized within Obrimini, the Obrimus-group, Stenobrimus-group and Theramenes-group. Keys are presented to distinguish between the three tribes now contained in the Obriminae, i.e. Obrimini, Tisamenini n. trib. and Miroceramiini. The genus Hennobrimus Conle, 2006 is synonymised with Mearnsiana Rehn & Rehn, 1939, based on the fact that the type-species of both genera are conspecific (n. syn.). Hennobrimus hennemanni Conle, 2006, the type-species of Hennobrimus, and Trachyaretaon manobo Lit & Eusebio, 2005 are synonymised with Mearnsiana bullosa Rehn & Rehn, 1939, the type-species of Mearnsiana (n. syn.). Theramenes dromedarius Stål, 1877 from the Philippines is removed from synonymy with the Wallacean Theramenes olivaceus (Westwood, 1859) and re-established as a valid species (rev. stat.).        The subfamily Heteropteryginae Kirby, 1896 is revised at the species-level and a new diagnosis is presented. Keys to the two genera and all 16 known species are provided along with new descriptions, differential diagnoses, lists of examined material, detailed information on the known distributions, measurements and illustrations of the insects and eggs. The intra-subfamiliar and intra-generic relationships are discussed and a cladogram is presented. Heteropteryginae contains two genera: Heteropteryx Gray, 1835 (Type-species: Phasma dilatatum Parkinson, 1798) and Haaniella Kirby, 1896 (Type-species: Phasma (Heteropteryx) muelleri de Haan, 1842). The distribution of this subfamily is restricted to Sundaland with the exception of a single species that is found in Vietnam. All other species are distributed in Borneo, Sumatra, the Mentawai Islands, Singapore, Peninsular Malaysia and Thailand. Heteropteryginae contains the largest and most striking members of the entire family Heteropteryginae, some of which are amongst the heaviest insects known. The subfamily is characterized by apomorphies such as the presence of wings, having a tympanal area (= stridulatory organ) in the basal portion of the alae, straight profemora, strongly shortened tarsi, lack of rough sensory-areas on the prosternum and typically X-shaped micropylar plate of the eggs. The sister-group of Heteropteryginae is represented by the Obriminae, with which it shares a beak-like secondary ovipositor in the females and presence of a medio-apical spine on the area apicalis. Both features are synapomorphies of Heteropteryginae + Obriminae.        The genus Haaniella Kirby, 1904 contains 16 known species, five of which are newly described herein. The genus Miniopteryx Zompro, 2004 (Type-species: Haaniella parva Günther, 1944) is synonymised with Haaniella on the basis that the distinguishing feature mentioned in the original description is a character that is frequently found throughout the genus (n. syn.). The type-species H. parva Günther, 1944 is automatically retransferred to Haaniella (rev. stat.). Haaniella aculeata n. sp. from western Sumatra is described from the male. Haaniella macroptera n. sp. from Singapore and the Johor state in southern Peninsular Malaysia is described from both sexes and the eggs. Haaniella gintingi n. sp. from Central Sumatra is described from both sexes and the eggs and Haaniella kerincia n. sp. from Western Sumatra is described from the insects only, the eggs being still unknown. One new species, Haaniella gorochovi n. sp., is the only representative of the genus and subfamily Heteropteryginae known from Vietnam and both sexes as well as the eggs are described. Haaniella erringtoniae (Redtenbacher, 1906) is endemic in Peninsular Malaysia, here removed from synonymy with H. muelleri (de Haan, 1842) and re-established as a valid species (rev. stat.). The Sumatran Haaniella glaber (Redtenbacher, 1906) is removed from synonymy with H. muelleri (Haan, 1842) and re-established as a valid species (rev. stat.). Leocrates glaber Redtenbacher, 1906 and Haaniella muelleri simplex Günther, 1944 are removed from synonymy with H. muelleri (Haan, 1842) (rev. stat.) and synonymised with H. glaber. Haaniella mecheli (Redtenbacher, 1906) and H. rosenbergii (Kaup, 1871) are removed from synonymy with H. muelleri (Haan, 1842) and re-established as valid species (rev. stat.). Haaniella erringtoniae novaeguineae Günther, 1934 and Haaniella muelleri var. b. (Haan, 1842) are synonymized with H. rosenbergii (Kaup, 1871) (n. syn.). The type-species Haaniella muelleri (Haan, 1842) is shown to be a fairly rare species that is restricted to Sumatra. All subsequent records of H. muelleri from outside Sumatra and references to captive breeding of stock originating from Peninsular Malaysia in Europe relate to H. erringtoniae (Redtenbacher, 1906). The previously unknown males and eggs of H. rosenbergii (Kaup, 1871) as well as the previously unknown females and eggs of H. parva Günther, 1944 are described and illustrated for the first time. Based on morphological characters of the insects and eggs three distinct species-groups are recognized within Haaniella. The muelleri species-group contains nine species that are distributed throughout Sumatra, the Mentawei Islands, Singapore and Peninsular Malaysia. These are characterized by the smooth ventral surface of the meso- and metafemora and lemon-shaped eggs which entirely lack the setae seen in the two other species-groups. The grayii species-group comprises four species, two of which are endemic in Borneo, one endemic in Sumatra and the fourth species being the only known representative of the subfamily in Vietnam. These species are characteristic for the prominent pair of spines on the abdominal tergites II-IV of males and long apically multidentate epiproct of females. The echinata species-group contains three exceptionally Bornean species, which are characterized by the long and apically pointed subgenital plate of females, which clearly projects beyond the epiproct, as well as the sub-basal lateral tooth of the anal segment of males. The muelleri species-group is sister to the remainder two species-groups.        Heteropteryx Gray, 1853 is a monotypical genus and only contains the type-species H. dilatata (Parkinson, 1798), which is found throughout Peninsular Malaysia, Thailand, Sumatra and Northeastern Borneo. This genus differs from Haaniella by the strongly conically elevated head, which posteriorly projects over the anterior margin of the pronotum, females being bright green or yellow in colour with plain and translucent pink alae and having distinct spines on the abdominal tergites, and males having a strongly shortened mesothorax and dull pink alae.        Lectotypes are designated for Haaniella parva Günther, 1944, Heteropteryx echinata Redtenbacher, 1906, Heteropteryx saussurei Redtenbacher, 1906 and Heteropteryx scabra Redtenbacher, 1906 to guarantee stability of these names.        Information on the habitats, host-plants, biology, life cycle, parasitism and captive breeding of the species of Heteropteryginae is presented and a list summarising all taxonomic changes presented herein.

* Title and MeSH Headings from MEDLINE®/PubMed®, a database of the U.S. National Library of Medicine.