Revision of the Oriental subfamily Heteropteryginae Kirby, 1896, with ...

Zootaxa 4159 (1): 001–219 http://www.mapress.com/j/zt/ Copyright © 2016 Magnolia Press

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http://doi.org/10.11646/zootaxa.4159.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:B9B69D54-BD32-4EF3-B684-A0812B098531

ZOOTAXA 4159

Revision of the Oriental subfamily 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) FRANK H. HENNEMANN*¹, OSKAR V. CONLE², PAUL D. BROCK³ & FRANCIS SEOW-CHOEN4 ¹ Reiboldstrasse 11, 67251 Freinsheim, Germany. E-mail: [email protected], Website: www.Phasmatodea.com ² Am Freischütz 16, 47058 Duisburg, Germany. E-mail: [email protected], Website: www.Phasmatodea.com ³ The Natural History Museum, Cromwell Road, London, SW7 5BD, U.K. E-mail: [email protected], Website: http://phasmida.speciesfile.org 4 Lee Kong Chian Natural History Museum, Singapore. E-mail: [email protected] * Corresponding author

Magnolia Press Auckland, New Zealand

Accepted by B. Montovani: 7 Jun. 2016; published: 1 Sept. 2016

FRANK H. HENNEMANN, OSKAR V. CONLE, PAUL D. BROCK & FRANCIS SEOW-CHOEN Revision of the Oriental subfamily 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) 219 pp.; 30 cm. 1 Sept. 2016 ISBN 978-1-77557-519-1 (paperback) ISBN 978-1-77557-520-7 (Online edition)

FIRST PUBLISHED IN 2016 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/j/zt

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2 · Zootaxa 4159 (1) © 2016 Magnolia Press

HENNEMANN ET AL.

Table of contents 1. 2. 2.1. 2.2. 3. 3.1 3.2 3.2.1 3.2.2 3.2.3 3.3 3.4 4. 4.1 4.1.1 4.1.2 4.1.3 4.2 5. 6. 6.1 6.2 6.3 7. 7.1 7.2 7.3 7.4 7.5 7.6 8. 8.1 9. 10.

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Acanthotaxy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Systematic background - the family Heteropterygidae Kirby, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Subfamily Dataminae Rehn & Rehn, 1939. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Subfamily Obriminae Brunner v. Wattenwyl, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Tribe Obrimini Brunner v. Wattenwyl, 1893. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Tribe Tisamenini n. trib.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Tribe Miroceramiini Zompro, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Subfamily Heteropteryginae Kirby, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Classification of Heteropterygidae Kirby, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Revision of Heteropteryginae Kirby, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Genus Haaniella Kirby, 1904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 The muelleri species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 The grayii species-group. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 The echinata species-group. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80 Genus Heteropteryx Gray, 1835 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96 Arguments for the inter- and intrageneric systematization of Heteropteryginae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106 Distribution of Dataminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108 Distribution of Obriminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 Distribution of Heteropteryginae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111 Biology & Life history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 Behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 Life cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 Parasitism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 Habitats & host plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 Rearing in captivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 Local beliefs / medicinal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115 Summary.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 List of taxonomic changes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120

Abstract 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 REVISION OF HETEROPTERYGINAE

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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.


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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. Key words: Phasmatodea, Heteropterygidae, Heteropteryginae, Obriminae, Dataminae, Heteropteryx, Haaniella, taxonomic revision, classification, new tribe, new species, new subspecies, new synonyms, lectotypes, keys, differentiations, descriptions, illustrations, eggs, habitats, ecology, biogeography, distribution, Oriental Region, Southeast Asia, Borneo, Sumatra, Singapore, Peninsular Malaysia, Thailand, Vietnam.

1. Introduction Besides the giant stick insects of the subfamily Clitumninae and the tribe Pharnaciini in particular (see Hennemann & Conle, 2008), the Oriental family Heteropterygidae Kirby, 1896 includes some of the most striking insects of the Old World region, some of which are amongst the world’s heaviest extant insects. This and the fact that most species are fairly easy to culture are the main reasons why so much attention has been given to this particular group of phasmids in the past decades. Several species of Heteropterygidae have been successfully cultured in Europe and have become favourite pets amongst phasmid enthusiasts, usually the larger members of the subfamily Heteropteryginae Kirby, 1896. The well known “Jungle Nymph” Heteropteryx dilatata (Parkinson, 1798) in particular is one of the main attractions at entomological exhibitions or zoological gardens world-wide. Numerous papers on the Heteropterygidae have since been published, most covering the biology, life cycle and breeding of certain species (e.g. Cocking, 1985 & 1986; D’Hulster, 1990; Abercrombie, 1993; Dräger, 2011). The first comprehensive taxonomic treatment of species in the family Heteropterygidae was in the first part of the monograph “Die Insektenfamilie der Phasmiden” by Brunner v. Wattenwyl & Redtenbacher (1906–1908). Subsequent taxonomic works on the family related to single taxa of the group or were restricted to limited faunistic regions. The most comprehensive treatments were the revision of Obriminae Brunner v. Wattenwyl, 1893 of the Philippine Islands by Rehn & Rehn (1939), and revision of the Heteropterygidae of Borneo by Bragg (1998 & 2001). Günther (1944) published a paper that dealt with members of the subfamily Heteropteryginae and described two new species. He also proposed a re-arrangement of the genus Haaniella Kirby, 1904 and provided keys to the species of this genus. Seow-Choen (2000a) provided an illustrated key to the species of Haaniella. Lastly, Zompro (2004) provided keys to all genera, phylogenetic hypotheses and a new arrangement of the subordinate taxa of the family Heteropterygidae. This work had since been critically discussed in subsequent molecular and phylogenetic studies (Whiting et al., 2003; Bradler, 2009). Although much attention has been drawn to this striking group of Oriental Phasmatodea in recent years, there are still questions concerning the currently supposed relationships between the subordinate taxa and arrangement of taxa within the subfamily Obriminae. The present work clarifies many of these uncertainties. Detailed studies of the two genera in the subfamily Heteropteryginae, i.e. Heteropteryx Gray, 1835 and Haaniella Kirby, 1904, were started as early as 1994. These include visiting museums that contain type specimens and other important material of the subfamily and have resulted in several new taxa being discovered. Furthermore examination of several hundred specimens had shown there still was considerable confusion concerning the identity and taxonomic status of several taxa from outside Borneo. Hence, a detailed taxonomic revision of the group appears warranted and is presented herein. The present study includes a new diagnosis of the subfamily and new descriptions of all known species along with distinguishing keys and information on the intraspecific variability, distribution, habitats and biology.

2. Material & Methods The present study is based on examination of all necessary type material and has received tremendous support from curators of all corresponding museums and institutions housing type specimens. In addition to the types all other material available from various public and private collections has been examined and determined, making a total of more than 950 specimens of the subfamily Heteropteryginae studied for the present revision. The specimens used for this study are almost without exception dried and pinned. Material in the first author's collection (FH) was preserved as described by Zompro (1996), with acetone or with silica-gel, material in the

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second author's collection (OC) with acetone or silica-gel. Certain specimens have been eviscerated and stuffed with cotton to maintain the natural colouration. Insects and eggs were examined using an entomological lens with 4x magnification and a Russian MBC-10 binocular microscope. Measurements were taken using a long ruler or a digital calliper and are given to 0.1 mm. If not differently cited, measurements include all examined and listed material as well as the type specimens, although measurements of type specimens are in some cases listed separately. For most species the description of the colouration was based on live material or colour images of live insects. If no live specimens were available the colouration is described from the dried specimens examined. In these cases attention needs to be drawn because the insects may have changed colour due to preservation. Only eggs which were already laid or removed from the female’s ovipositor were examined and hence all descriptions herein are exclusively based on fully developed eggs. The terminology used for the spination and armature of the insects is an extended and updated version of the acanthotaxy established for the subfamily Obriminae by Rehn & Rehn (1939), which has already been applied to Heteropteryginae by Bragg (1998, 2001; → 2.1). The terminology used to describe external and internal egg structures follows that of Sellick (1997 & 1998). The cladogram in Fig. 97b and phenogram in Fig. 355 have been constructed manually and are partly been based on character states that have already been discussed and used in previous molecular and phylogenetic studies by other authors (e.g. Bradler, 2009; Bradlet et al., 2014). For the discussion of taxa within the subfamily Heteropteryginae, the following format is used. Genera and species are listed in alphabetical order. For each species, the data of type-material is included in the synonymic lists and follows the corresponding citation. The section “Material examined” lists all examined non-type material, arranged in geographical order. The data are reproduced exactly as on the original labels attached to the concerned specimens and labels are separated by a semicolon. The differential diagnosis is followed by an etymology section, descriptions of the adult insects, nymphs and eggs, notes on the intraspecific variability, and comments which provide additional information on the type specimens, synonymies, host plants, biology or culture status of the species. The section “Distribution” provides a list of all known localities for each species with the references, i.e. specimens at hand or published sources, included in square brackets. Finally, the complete number of specimens examined for this study, which also includes the types, is given at the end of each column. Lectotypes are designated for four species in order to guarantee stability of the concerned names. The revision of the subfamily Heteropteryginae at the species level justifies the designation of lectotypes according to Article 74 of the International Code of Zoological Nomenclature (ICZN, 1999).

2.1. Acanthotaxy The arrangement, size and shape of the cephalic, thoracic spines as well as the abdominal spines are important and useful features for the identification of species within the subfamily Heteropteryginae. Most species show relatively minor variabilities in these features and many species may easily be recognized by these differences of their body armature. Names for the major spines in the subfamily Obriminae were first proposed by Rehn & Rehn (1939). They suggested that their terminology could be modified to apply to other spiny phasmatodeans. This was already done by Bragg (1998, 2001) for the subfamily Dataminae but no definite terminology has been introduced for Heteropteryginae to date. In general there is much homology between structures in Obriminae and Heteropteryginae, so in most cases the equivalent or closest terms of Rehn & Rehn (1939) have been selected for naming the corresponding spines in Heteropteryginae. New terms are only introduced for spines present in Heteropteryginae but not present in Obriminae. A list of the terms used in the descriptions of species of Heteropteryginae and an explanatory illustration are presented below (Fig. 1). Armature of the head Supra antennals = a pair of tubercles or small spines situated a short distance behind the bases of the antennae. These may be rudimentary or absent. Supra-orbitals = a pair of often strong spines placed slightly medial and posterior of the eyes. Sometimes rudimentary or absent. Anterior occipitals = a pair of medium to large spines at the anterior portion of the vertex, just in front of the coronals and always smaller than the former. Anterior coronals = a pair of strong and large spines roughly in the central portion of the vertex. While mostly simple, these


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may be strongly enlarged in some species to form laterally compressed, crest-like structures with 2–5 apical spines. These spines correspond to occipitals according to Rehn & Rehn (1939) but since there is only one pair in Heteropteryginae, which together with the posterior coronals forms a “crown-like” element on the raised portion of the vertex in Heteropteryginae, they are here regarded as coronals. Posterior coronals = a pair of large and strong spines close to the posterior margin of the head. Usually positioned as two corners of a square together with the two anterior coronals. Lateral coronals = a pair of strong more or less decidedly laterad directed spines near the postero-lateral angles of the head.

Armature of the pronotum Antero-lateral pronotals = a pair of laterally directed spines near the antero-lateral angles of the pronotum. Anterior mesal pronotals = a small and closely spaced pair of tubercles or spines near the anterior margin of the pronotum. Often rudimentary or absent. Pre-median pronotals = the large pair of spines in the anterior portion of the pronotum just in front of the transverse median sulcus. Usually by far the largest spines on the pronotum. Inter-posterior pronotals = a pair of more or less distinct spines some distance in front of the posterior margin of the pronotum. May be rudimentary or absent. Postero-lateral pronotals = a pair of laterally directed spines near the postero-lateral angles of the pronotum. Usually more or less equal in size to the antero-lateral pronotals.

Armature of the mesonotum Antero-lateral mesonotals = a pair of laterally directed spines near the antero-lateral angles of the mesonotum. These are very variable in size, either only slightly larger than the other lateral mesonotals or much enlarged and very prominent. Anterior mesal mesonotals = a closely spaced pair of tubercles or spines near the anterior margin of the pronotum. Often rudimentary or absent. Anterior mesonotals = the strong and distant pair of spines some distance behind the anterior margin of the mesonotum. Usually lying in a transverse line together with the antero-lateral mesonotals and anterior mesal pronotals. Pre-median mesonotals = a pair of often strong closely spaced spines in the anterior portion of the mesonotal disk slightly in front of the centre of the mesonotum. Often the largest spines on the entire mesonotum. Post-median mesonotals = a more distant pair of spines in the posterior portion of the mesonotal disc. These are variable in size and sometimes rudimentary or absent. Lateral mesonotals = a series of often unequally sized small to median tubercles or spines along the lateral margins of the mesonotum. Tegminal = a single spine positioned on the joint of each tegmen. Posteromedial mesonotals = a very closely spaced pair or cluster of 4–6 spines positioned near the posterior margin of the mesonotum and between the joints of the tegmina. These are either separate spines or positioned on a common base, forming a crown-like element. In species with four such spines, these are usually positioned as the four corners of a square. Their size is variable, ranging from small spinules to very large and slender spines that might represent the largest of all mesonotal spines. Some intraspecific variability may be seen in certain species.

Armature of the meso- and metapleura Laterals = a variable number of often unequally sized tubercles or spines along the lateral margins of the pleurae. Supra-coxal = a single often very strong laterad directed spine in the posterior portion of the pleural disc above the coxae.

2.2. Abbreviations ANSP: BMKP: BPBM: ETHZ: HLMD: LKCNHM: MAMU: MCZC: MHNG: MMUM: MNHN: MNHU:

Academy of Natural Sciences, Philadelphia, Pennsylvania / U.S.A. Brunei Museum, Kota Batu / Brunei. Bernice P. Bisoph Museum, Honolulu, Hawaii / U.S.A. Eidgenössische Technische Hochschule, Zürich / Switzerland. Hessisches Landesmuseum Darmstadt / Germany. Lee Kong Chian Natural History Museum / Singapore. MacLeay Museum, University of Sydney, Sydney, NSW / Australia. Museum of Comparative Zoology, Harvard University, Cambridge / U.S.A. Museum d´Histoire Naturelle, Geneve / Switzerland. Manchester Museum, University of Manchester / U.K. Museum National d´Histoire Naturelle, Paris / France. Museum für Naturkunde der Humboldt-Universität, Berlin / Germany.



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MIZT: MSNG: NHMB: NHMM. NHMUK: NHMW: NMSC: RBINS: RMNH: ROM: SFDK: SMTD: SMNS: SMSM: UMB: UMO: ZIN: ZMUH: ZSMC: CLC: FH: FSC: OC: PDB: PEB: HT: AT: PT: ST: LT: PLT:

Museo Regionale de Scienze Naturali, Torino / Italy. Museo Civico di Storia Naturale „Giacomo Doria“, Genova / Italy. Naturhistorisches Museum, Basel / Switzerland. Natuurhistorisch Museum, Maastricht / Netherlands. Natural History Museum, London / U.K. Naturhistorisches Museum, Vienna /Austria. National University of Singapore / Singapore. Institute Royal des Sciences Naturelles de Belgique, Bruxelles / Belgium. Nationaal Natuurhistorisch Museum, Leiden / Netherlands. Royal Ontario Museum / Canada. Sarawak Forestry Department, Kuching, Sarawak / Malaysia. Staatliches Museum für Tierkunde, Dresden / Germany. Staatliches Museum für Naturkunde, Stuttgart / Germany. Sarawak Museum, Kuching, Sarawak / Malaysia. Überseemuseum Bremen / Germany. University Museum, Hope Entomological Collections, Oxford / U.K. Zoological Institute, Russian Academy of Sciences, St. Petersburg / Russia. Zoologisches Museum und Institut, Hamburg / Germany. Zoologische Staatsammlung München, Munich / Germany. Private collection of C. L. Chan, Kota Kinabalu / Malaysia. Private collection of Frank Hennemann, Freinsheim / Germany. Private collection of Francis Seow-Choen / Singapore. Private collection of Oskar V. Conle, Bolsterlang / Germany. Private collection of Paul D. Brock, Brockenhurst / U.K. Private collection of Phil E. Bragg, Nottingham / U.K. Holotype Allotype Paratype Syntype Lectotype Paralectotype

3. Systematic background—the family Heteropterygidae Kirby, 1896 Heteropteryginae was originally introduced as a subfamily by Kirby (1896: 472). He only included the genus Heteropteryx Gray, 1835, as did Rehn (1904: 89). In his “A Synonymic Catalogue of Orthoptera, Vol. 1” Kirby (1904a: 396) included several other Oriental genera in this subfamily, which were subsequently attributed to the two areolate tribes Heteropterygini and Obrimini by Brunner v. Wattenwyl & Redtenbacher (1906: 36, 162). Redtenbacher however was not aware of Kirby’s catalogue and included the two Oriental genera Leocrates Stål, 1875 and Heteropteryx Gray, 1835 as well as the Madagascan Anisacantha Redtenbacher, 1906 and Parectatosoma Wood-Mason, 1879 in the tribe Heteropterygini. However he did not recognize the close relation between the first two genera and the taxa he listed in Obrimini. In his catalogue of Philippine Phasmatodea Bruner (1915: 228) treated the Obrimini sensu Redtenbacher (1906: 36) as a family. Rehn & Rehn (1939: 400) regarded the Obrimini sensu Redtenbacher as a subfamily and distinguished two tribes within Obriminae, i.e. the Obrimini and Datamini. Günther (1953: 551) united the Heteropterygini sensu Redtenbacher and Obrimini sensu Redtenbacher in the subfamily Heteropteryginae, hence accepted Kirby’s subfamily and included the four tribes Heteropterygini, Obrimini, Datamini and Anisacanthini, the latter tribe containing the Madagascan genera that had been placed in Heteropterygini by Redtenbacher (1906: 162). The Obrimini and Datamini were generally accepted by Günther (1953: 551) as defined by Rehn & Rehn (1939). Subsequent authors have generally accepted the arrangement introduced by Günther (Beier, 1957, 1968; Klante, 1976; Bradley & Galil, 1977; Bragg, 1998, 2001). The possible phylogenetic relationships within Heteropteryginae sensu Günther (1953) were briefly discussed by Klante (1976), who presented a preliminary cladogram.


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Zompro (1996: 161) raised Kirby’s Heteropteryginae to family rank. Subsequently, Zompro (2004) excluded the Madagascan Anisacanthini from the Heteropterygidae and raised the traditional tribes to subfamilies, the Heteropteryginae, Obriminae and Dataminae. Furthermore, he subdivided the Obriminae into the three tribes Obrimini, Eubulidini and Miroceramiini but these as well as the inter-familiar relationships proposed by Zompro (2004) are poorly supported as discussed by Bradler (2009: 95 ff). A new arrangement of the subfamilies and tribes of Heteropterygidae, which mostly agrees with molecular data provided by Whiting et al. (2003) and a phylogenetic study based on morphology by Bradler (2009, Fig. 97a) is presented below (Fig. 97b). The two Madagascan genera Anisacantha Redtenbacher, 1906 and Parectatosoma Wood-Mason, 1879, which had traditionally been affiliated with the Oriental Heteropterygidae (e.g. Redtenbacher, 1906; Beier, 1957, 1968; Klante, 1976; Bradley & Galil, 1977; Bragg, 1998), are in fact not closely related and were placed in the family Anisacanthidae by Zompro (2004: 240). The latter placed the Heteropterygidae and Anisacanthidae as subordinate taxa of the supra-family Bacilloidea. Subsequent molecular studies strongly contradict these hypotheses (e.g. Bradler et al., 2015). The Heteropterygidae are currently regarded as monophyletic and the monophyly of the clade seems to be fairly well supported. For instance, Bradler et al. (2015: 2) revealed a Bayesian posterior probability (BPP = 0,99) for Heteropterygidae (referred to as the subfamily Heteropteryginae). Bradler (2009: 95) regarded the paired sensory-areas on the prosternum and the occasional presence of a sensory-area on the profurcasternum as an autapomorphy. The latter was stated to be lacking in the tribe Heteropteryginae (Zompro, 2004; Bradler, 2009) but this is not true for certain representatives of that subfamily. Examination has shown that in fact Heteropteryginae do possess a central sensory-area on the profurcasternum (Figs. 78–83). Furthermore, Bradler (2009: 95) suggested the stridulatory organ in the basal portion of the alae of certain Heteropteryginae as an autapomorphy of Heteropterygidae and that this belongs in the ground pattern of the family. However, this feature is of poor value since most taxa are apterous. Furthermore, we believe that the wings have been gained two times independently in the tribe Heteropteryginae and in a single genus of Obriminae. Hence, the presence of wings is unlikely to be plesiomorphic and consequently does not support the stridulatory organ to belong in the ground pattern of Heteropterygidae. This generally agrees with the results of a molecular study by Whiting et al. (2003). Zompro (2004: 193) postulated that the ground pattern of Heteropterygidae includes straight and not basally compressed profemora, the presence of wings and a short mesothorax. According to Zompro the straight profemora, medio-apical spine of the area apicalis and the secondary ovipositor of ♀♀ are autapomorphies of Heteropteryginae, which have been reduced in the Dataminae. It is however much better supported that these features are primarily lacking in Dataminae and have been secondarily developed in Heteropteryginae (Bradler, 2009: 95). Furthermore, Zompro (2004: 193) postulated that the beak-like secondary ovipositor of ♀♀ to be a synapomorphy of all representatives of the family Heteropterygidae. However this is not possible since it is primarily lacking in Dataminae and was secondarily developed in the Obriminae and Heteropteryginae. Consequently, the ♀♀ ovipositor cannot belong in the ground pattern of Heteropterygidae but is a synapomorphy of Obriminae + Heteropteryginae. The medio-apical spine of the area apicalis is also present in the Madagascan Anisacanthidae and was interpreted contrarily by Zompro (2004). Zompro either regarded it as an autapomorphy of the Heteropterygidae (Zompro, 2004: 192, fig. 119) or as an autapomorphy of Aniscanthidae + Heteropterygidae + Bacillidae (Zompro, 2004: 26, fig. 4), with the spine again reduced in the Bacillidae. Both scenarios are unlikely, since the spine is lacking in Dataminae and hence is not present in all members of Heteropterygidae. It appears more likely to have been gained independently in Obriminae + Heteropteryginae as well as the Madagascan Anisacanthidae, hence it represents a homoplasy. The results here presented strongly contradict the relationships postulated by Zompro (2004) but mostly agree with the suggestions and results presented by Klante (1976), Whiting et al. (2003), Bradler (2009) and Bradler et al. (2014, 2015). According to Zompro (2004) the Heteropteryginae are a basal lateral lineage of Heteropterygidae since they possess wings, which a priori is regarded as plesiomorphic. This would mean the loss of wings is a synapomorphy of Obriminae + Dataminae. A basal position of Dataminae however is much more likely and better supported, which consequently means that the wings were gained independently in the Heteropteryginae and within a single genus in the Obriminae (Fig. 97). This hypothesis is supported by further features that are shared by Obriminae + Heteropteryginae but are lacking in Dataminae, i.e. the presence of a secondary beak-like ovipositor in ♀, presence of a medio-apical spine on the area apicalis and increasingly elongated antennae. Hence these three features are most certainly synapomorphies of Obriminae and Heteropteryginae (Fig. 97b). Furthermore,



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Dataminae possesses paired sensory-areas on the prosternum and an additional central sensory-area on the profurcasternum. The former have been reduced in Heteropteryginae and the latter has been reduced in Obriminae, but is present in Heteropteryginae. Hence, the loss of either sensory-area must be regarded as apomorphic while their presence must be regarded as the plesiomorphic character for Heteropterygidae. As a result Dataminae appears to be the basalmost clade of the family Heteropterygidae. This is in contrast to the relationships suggested by Zompro (2004), who postulated Heteropteryginae to be the basal-most clade. Reconstructing the ground pattern of Heteropterygidae based on the results of the present study suggests the common ancestor was a small, stocky and robust, wingless, ground-dwelling phasmid with moderate sexual dimorphism, a short mesothorax hardly longer than the prothorax, short and robust antennae shorter than the fore legs, an area apicalis on all six tibiae, basally compressed profemora, a well developed vomer in ♂♂ and no ovipositor in ♀♀.

Key to the subfamilies of Heteropterygidae Adults 1. 2. -

Area apicalis with a spine medio-apically; epiproct and subgenital plate of ♀♀ elongated and forming a beak-like ovipositor* . .. .. ... .. .. .. ... .. .. .. .. ... .. .. .. ... .. .. .. ... .. .. .. .. ... .. .. .. . .. .. .. .. .. ... .. .. .. ... .. .. .. .. ... .. .2 Area apicalis without a spine medio-apically; no ovipositor, epiproct and subgenital plate of ♀♀ not elongated, subgenital plate cup-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dataminae Prosternum with two rough sensory areas (Figs. 41, 42, 64, 75) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Obriminae Prosternum without rough sensory areas (Figs. 78-83). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heteropteryginae

* in Hoploclonia Stål, 1875 (subfamily Obriminae) formed by an elongated anal segment and subgenital plate (Fig. 66)

Eggs 1. 2. -

Capsule more or less bullet-shaped and longer than wide; either smooth or with short setae . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Capsule more or less spherical or laterally compressed; setae of capsule long and with hook-like apices, adhesive . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dataminae Micropylar plate quadri-lobate and principally X-shaped, with two anterior and two posterior expansions; no median line in internal micropylar plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heteropteryginae Micropylar plate basically tri-lobate, only with two posterior expansions; internal micropylar plate with a median line. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Obriminae

3.1. Subfamily Dataminae Rehn & Rehn, 1939 Type-genus: Datames Stål, 1875 (= Pylaemenes Stål, 1875) Datamini Rehn & Rehn, 1939: 401, 405, 484. Günther, 1953: 552. Klante, 1976: 74, 76. Bradley & Galil, 1977: 198. Sellick, 1998: 208, fig. 9. Bragg, 1998: 46. Bragg, 2001: 124. Bradler, 2009: 97. Dataminae, Zompro, 2004: 218. Seow-Choen, 2016: 374.

Diagnosis (♂, ♀): Small to very small (body length 18.5–64.0 mm), apterous Heteropterygidae. Body robust, stocky and ± verruculose, occasionally spinose (less frequently in ♀♀). Sexual dimorphism modest, with ♂♂ just slightly shorter but considerably more slender and more spinose than ♀♀. Colouration various shades of brown and grey, ♂♂ may be more varied in colour. No ocelli. Antennae robust and at best equal in length to fore legs, often shorter. Scapus with spines on outer lateral margin (exception Epidares Redtenbacher, 1906). Antennae robust and shorter than fore legs with antennomeres perlamorph or hardly longer than wide and the median antennomeres at


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best 3x longer than wide (exception Epidares Redtenbacher, 1906); with 13–26 antennomeres. Antennomere III ± elongate and considerably longer than IV. No gula. Pronotal foramina displaced towards posterior by roughly its own diameter. Pronotum occasionally with a pair of pre-median spines. Prosternum with a pair of rough sensoryareas. Proforcasternum with a single antero-median sensory-area. Mesonotum either unarmed or with anterior- and/ or posterior mesonotal spines (latter may be composite); occasionally with antero-lateral spines. Metanotum occasionally with posterior metanotal spines, which may be composite. Meso- and metapleurae sometimes with a supra-coxal spine. Meso- and metasternum simple, not carinate and without foramina on metasternum. Median segment much shorter than metanotum, transverse. Abdomen ± widened and swollen medially in ♀♀ (segments IV and V widest), segments II–VII roughly parallel-sided in ♂♂. Segments II–VII transverse in ♀♀, transverse or at best slightly longer than wide in ♂♂. Sternites ± tectiform or with a longitudinal median carina in ♀♀, simple in ♂♂. Praeopercular organ on sternum VII of ♀♀ indistinct, at best formed by one or two blunt projections near posterior margin. Tergites VIII–X in ♂♂ wider than previous segments; IX often with a median projection or laterally compressed crest or carina (♀♀ in particular). Posterior margin of anal segment variable. ♀♀ without a beak-like ovipositor, epiproct and subgenital plate not elongated; the latter bulgy, strongly convex and often carinate longitudinally, never projecting over apex of anal segments. Gonapophyses VIII longer than gonapophyses IX. Gonangulum present. Poculum of ♂♂ strongly convex, often conical. Vomer of ♂♂ well developed and roughly triangular with a single terminal hook. Cerci very small. Profemora curved and compressed basally. Femora rectangular in cross-section; no median carina on ventral surface. Armature of femora reduced to single blunt teeth (more distinct on dorsal carinae). Tibiae unarmed, at best with some very minute blunt teeth in basal portion of dorsal carinae. All tibiae with a distinct area apicalis; no medio-apical spine. Tarsi very short, basitarsus at best slightly longer than following tarsomere. Eggs (Figs. 11–20): Large in relation to size of insects. Shape ± spherical, only in one case (i.e. Spinodares Bragg, 1998, Fig. 20a–c) somewhat elongate and compressed laterally. Operculum flat. Entire capsule and operculum set with conspicuous, ± long and apically hooked or multiply branched setae (these are easily shed and only present in newly laid eggs in certain species of Dares Stål, 1875). Micropylar plate large and very variable in shape, either tri-lobate with only one anterior expansion or quadri-lobate with two anterior and two posterior expansions. Internal micropylar plate with a very short and indistinct median line. Comments: The subfamily Dataminae currently contains eight genera and almost 40 species, all of which are small to very small (body length < 65 mm), very stocky and robust, apterous and mostly ground-dwelling taxa distributed from Japan, Taiwan and China in the north over entire Sundaland as far southeast as Wallacea (Fig. 343). No members have so far been recorded from the Philippines. The monotypical Woodlarkia Günther, 1932 (Type-species: Karabidion scorpionides Montrouzier, 1855) is the only representative that is known from Woodlark Island east of New Guinea (Otte & Brock, 2005: 340), but the absence of Dataminae on New Guinea and original description of this taxon, rather suggest it to belong in what is currently regarded as the subfamily Eurycanthinae. Unfortunately, the type specimens of Woodlarkia scorpionides (Montrouzier, 1855) are not traceable, hence any decision on its systematic position is impossible. Anyway, Woodlarkia is very unlikely to belong in Dataminae and consequently excluded in the list of genera below. The predominantly ground-dwelling Dataminae are characterized by the primary lack of an ovipositor in ♀♀, presence of paired sensory-areas on the prosternum and an additional central sensory-area on the profurcasternum (Bradler, 2009: 95, 97; Fig. 10), primary lack of a medio-apical spine on the area apicalis, short and robust often perlamorph antennae that are no longer than the fore legs and more or less spherical eggs, which possess specialized apically hooked setae. The profemora are compressed and curved basally. The scapus mostly bears spines on the exterior lateral carina. The subgenital plate of ♀♀ is strongly convex, roundly cup-shaped and never projects over the posterior margin of the anal segment. Since there is no ovipositor in ♀♀ the eggs are not laid into a substrate as in Obriminae and Heteropteryginae, but are simply dropped to the ground or ♀♀ use the fore legs to bury their eggs (Abercrombie, 1992). This strategy of egg deposition is regarded as the plesiomorphic state for Phasmatodea and believed to be part of the ground pattern of the order (Bradler, 2009; Goldberg et al., 2015). The eggs are more or less spherical in shape (Figs. 11–20) and the conspicuous hook-like structure of the setae of the eggs make them adhere to various surfaces. The micropylar plate is basically Y-shaped or tri-lobate, with two posterior expansions, which vary considerably in shape and may sometimes even reach as far as to the ventral surface of the egg-capsule. More rarely quadri-lobate shapes occur, which possess two lateral expansions (Fig. 14). Previous authors (e.g. Zompro, 2004, Bradler, 2009, Bradler et al., 2015) are in agreement that the Dataminae are



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the sister-group of the remaining Heteropterygidae, i.e. Obriminae + Heteropteryginae. The lack of a beak-like ovipositor in ♀♀ and the strategy of egg deposition, curved and compressed profemoral base, lack of wings and lack of a median line in the eggs seem to be plesiomorphic and might belong in the ground pattern of Heteropterygidae (see also Bradler, 2009: 97, fig. 42). This is also true for the medially widened and swollen abdomen of ♀♀, which is also found in Obriminae and Heteropteryginae as well as numerous other Phasmatodea. The lack of a spine on the area apicalis also appears to be plesiomorphic because this is typical for most areolate Phasmatodea, the medio-apical spine found in Obriminae and Heteropteryginae otherwise only occurring in Phylliinae Kirby, 1904 and members of the Madagascan family Anisacanthidae Günther, 1953. The multiply branched or hook-like apices of the setae of the egg-capsule, lateral spines on the exterior carina of the scapus, displaced foramina of the pronotum and tectiform or longitudinally carinate abdominal sternites of ♀♀ appear to be autapomorphies of Dataminae. A sensory-area on the profurcasternum is lacking in Obriminae and within the Heteropterygidae otherwise only found in certain members of Heteropteryginae, in which however it is increasingly reduced (→ see below). The specialisations of the setae of the eggs and exterior spines on the scapus are peculiar to Dataminae and represent autapomorphies of this subfamily (Fig. 97b). The micropylar plate of the eggs comprises tri- and quadri-lobate shapes, the tri-lobate type also being found in the Obriminae, while the quadri-lobate type is typical for Heteropteryginae. The basalmost taxon of Dataminae seems to be Planispectrum Rehn & Rehn, 1939, which in addition to the very small size (body length < 35 mm) is very characteristic for the very short, robust and perlamorph antennae. These are considerably shorter than the fore legs and consist of no more than 21 antennomeres, with some species having only 13 antennomeres (e.g. P. bakiense Zompro, 1998 and P. bengalensis (Redtenbacher, 1906), Fig. 3). Epidares Redtenbacher, 1906 appears to be the sister-group of the remainder genera of Dataminae (see Bragg, 1998: 113, fig 113 & Bragg, 2001: 128, fig. 34). This is supported by the elongated and slender antennae, which are unique in Dataminae, being almost as long as the fore legs and having the median antennomeres as much as 3x longer than wide. Furthermore, the scapus lacks the spines on the outer lateral margin seen in all other genera. While all other genera have the body surface more or less verrucose or rugulose, Epidares is the only genus in which the dorsal body surface is merely granulose (Fig. 7). Genera included: 1. Dares Stål, 1875: 51, 93. (Type-species: Dares validispinus Stål, 1875: 93, by subsequent designation of Kirby, 1904b: 400). 2. Epidares Redtenbacher, 1906: 53. (Type-species: Phasma (Acanthoderus) nolimetangere de Haan, 1842: 135, by subsequent designation of Bragg, 1998: 78). 3. Hainanphasma Ho, 2013: 203 (Type-species: Hainanphasma cristata Ho, 2013: 203, by original designation). 4. Orestes Redtenbacher, 1906: 47. (Type-species: Orestes verruculatus Redtenbacher, 1906: 47 (= Orestes mouhotii (Bates, 1865)), by monotypy). 5. Planispectrum Rehn & Rehn, 1939: 484. (Type-species: Platymorpha cochinchinensis Redtenbacher, 1906: 46, by original designation). [Replacement name for preoccupied Platymorpha Redtenbacher, 1906] = Platymorpha Redtenbacher, 1906: 46. (Type-species: Platymorpha cochinchinensis Redtenbacher, 1906: 46, by subsequent designation of Rehn & Rehn, 1939: 484). [Preoccupied by Platymorpha Jacoby, 1888 (Coleoptera), replaced by Planispectrum Rehn & Rehn, 1939] = Platyphasma Uvarov, 1940: 116. (Type-species: Platymorpha cochinchinensis Redtenbacher, 1906: 46, by indication). [Unnecessary replacement name for preoccupied Platymorpha Redtenbacher, 1906. Synonymised by Bradley & Galil, 1977: 198] 6. Pylaemenes Stål, 1875: 51, 93. (Type-species: Phasma (Pachymorpha) coronatum de Haan, 1842: 137, by subsequent designation of Kirby, 1904b: 400). = Datames Stål, 1875: 51, 93. (Type-species: Acanthoderus oileus Westwood, 1859: 53, by subsequent designation of Rehn, 1904: 89). [Synonymised by Hennemann, 1998: 125] = Pyloemenes Bradley & Galil, 1977: 198. [Subsequent misspelling of Pylaemenes Stål, 1875] 7. Spinodares Bragg, 1998: 86. (Type-species: Spinodares jenningsi Bragg, 1998: 86, by original designation).

3.2. Subfamily Obriminae Brunner v. Wattenwyl, 1893 Type-genus: Obrimus Stål, 1875b: 49. Obrimi Brunner v. Wattenwyl, 1893: 98. Obrimini, Redtenbacher, 1906: 36 (in part). Günther, 1953: 552 Klante, 1976: 74.


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Bradley & Galil, 1977: 199. Bragg, 1998: 31. Sellick, 1998: 208, fig. 8. Bragg, 2001: 102. Bradler, 2009: 97. Obriminae, Rehn & Rehn, 1939: 400. Kevan, 1982: 381. Zompro, 2004: 199. Otte & Brock, 2005: 31. Seow-Choen, 2016: 412. Obriminidae, Bruner, 1915: 228. Therameninae Karny, 1923: 232. Shiraki, 1935: 23, 24.

Diagnosis (♂, ♀): Small to medium sized (body length 30.0–115.0 mm) Heteropterygidae, mostly apterous and very rarely winged. Body ranging from robust and stocky to slender and fairly stick-like. Body either unarmed or to a variable degree spinose; head and thorax in particular. Sexual dimorphism distinct, with ♂♂ considerably smaller and much slenderer than ♀♀. Colouration mostly various shades of brown and grey, rarely green. No ocelli. Antennae elongate, slender and at least equal in length to fore legs; with 24–28 antennomeres. Scapus simple, unarmed. Antennomere III elongate and longer than IV. Remaining antennomeres ± elongate and median antennomeres longer than wide. Head variable in shape and ranging from fairly flattened to having the vertex conically raised; either unarmed or spinose. No gula (exception Hoploclonia Stål, 1875). Pronotal foramen at anterior margin on pronotum. Pronotum unarmed or spinose. Prosternum with a pair of rough sensory-areas. No sensory on profurcasternum. Mesothorax at least slightly longer than pronotum, usually considerably longer and elongate; ± widening towards posterior or more rarely expanded medially in ♀♀, the anterior 2/3 usually ± parallel-sided in ♂♂. Meso- and metanotum ranging from unarmed to strongly spinose; occasionally with a ± raised trigonal area in anterior portion. Meso- and metapleurae usually with ± distinct sometimes composite suparcoxal spines. Meso- and metasternum simple or carinate medio-longitudinally. Metasternum often with lateral formina (= slit-like to ± round invaginations). If tegmina and alae present (only Miroceramia Günther, 1934), then tegmina elongate and roughly 2/3 the length of alae and anal fan plain transparent grey. Median segment shorter than metanotum. Abdomen of ♀♀ usually somewhat widened and swollen medially with segments IV and V widest and the remaining segments gradually narrowing; segments II–VII roughly parallel-sided in ♂♂ with VIII– X considerably broader than previous. Tergites unarmed or II–V spinose. Sternites simple or with a fine longitudinal median carina; smooth or minutely spinose. Praeopercular organ of ♀♀ indistinct. Epiproct of ♀♀ elongated and together with the elongated subgenital plate forming a beak-like ovipositor (one genus, i.e. Hoploclonia Stål, 1875, has the epiproct short but the anal segment elongated apically to form an ovipositor, Fig. 66). Subgenital plate carinate longitudinally. Gonapophyses VIII longer than gonapophyses IX. Gonangulum present. Cerci of both sexes very small and laterally compressed. Poculum of ♂♂ ± convex and cup-shaped. Vomer well developed, roughly triangular in shape and with a single terminal hook. Profemora compressed and curved basally. All femora rectangular in cross-section, ventral surface without a median carina. Carinae to a variable degree armed with teeth. Meso- and metafemora conspicuously broadened or thickened in certain taxa. Tibiae trapezoidal in cross-section with anterior carinae nearing; dorsal carinae smooth, ventral carinae ± dentate. Area apicalis with a ± distinct medio-apical spine. Tarsi elongate with basitarsus considerably longer than tarsomere II. Eggs: Variable in size and shape, capsule ± bullet-shaped and up to 4x longer than wide; occasionally anterior portion narrowed and polar-area roundly evaginated. Operculum flat to very gently conical; rarely with specializations like a central pit or rim of hairy structures. Surface ranging from almost smooth over punctate to rugulose; in certain taxa irregularly setose or with hairy-net-like sculpturing. Micropylar plate elongate, covering much of dorsal egg surface and tri-lobate, the two posterior expansions very variable in shape. Internal plate with outer margin lightly notched and with a distinct median line. Comments: The Obriminae was first recognized by Brunner v. Wattenwyl (1893) as Obrimi and was subsequently considered a tribe of the areolate Phasmatodea by Brunner v. Wattenwyl & Redtenbacher (1906). Bruner (1915) treated them as the family Obriminidae and Karny (1923) used the synonymic subfamily name Therameninae. All of these workers included within this group, the taxa now regarded as separate subfamily, namely the Dataminae (Zompro, 2004). Kirby (1904: 396) was the first to recognize the close relation between



13

these two taxa and the genus Heteropteryx Gray, 1835 and united them in the subfamily Heteropteryginae. This was overlooked by Brunner v. Wattenwyl & Redtenbacher (1906) who separated the Obrimini and Heteropterygini. Rehn & Rehn (1939) sub-divided the Obriminae into the two tribes Obrimini and Datamini. Günther (1953: 551) placed these two tribes in the subfamily Heteropteryginae. Zompro (2004) raised the tribes to subfamily level and united them in the family Heteropterygidae. The subfamily Obriminae was sub-divided into three tribes by Zompro (2004: 201), the Obrimini Brunner v. Wattenwyl, 1893, the Eubulidini Zompro, 2004 and the Miroceramiini Zompro, 2004, but the monophyly of the Eubulidini and Miroceramiini in particular is poorly supported. This sub-division based on the characters used by Zompro, resulted in three tribes, which appear highly arbitrary, are clearly paraphyletic or polyphyletic respectively and are no natural groups (→ see comments under 3.2). A new arrangement of Obriminae is proposed below. At present 18 genera and some 61 species are attributed to the Obriminae, which are predominantly distributed throughout the Philippines and Borneo with one genus each in northern Micronesia (Palau Islands) and in the Wallacea (Sulawesi and Seram). With the exception of Hoploclonia Stål, 1875 (→ 3.2.2, Fig. 66) ♀♀ have a beaklike secondary ovipositor, that is formed by an elongated subgenital plate and epiproct (Figs. 43–46, 65), a feature shared with Heteropteryginae and hence a synapomorphy of Obriminae + Heteropteryginae (Bradler, 2009: 97). Another synapomorphy of Obriminae + Heteropteryginae is the presence of a medio-apical spine on the area apicalis, which however is poorly developed in certain Obriminae-genera (→ tribe Tisemenini n. trib.). The profemora are compressed and curved basally but are increasingly straight in the only winged taxon, the Wallacean Miroceramia Günther, 1934. The prosternum bears a pair of sensory-areas, but in contrast to the Dataminae the central sensory-area of the profurcasternum is secondarily reduced and lacking, which is an autapomorphy of Obriminae (Figs. 41–42, 64, 75). The pronotal foramina are placed at the anterior margin of the pronotum. The antennae are very variable in shape and length, ranging from being hardly as long as the fore legs with the antennomeres just moderately elongate in the Tisamenini n. trib. (→ 3.2.2) to very long, slender and filiform with the median antennomeres extemely elongated and much longer than the fore legs (e.g. Stenobrimus Redtenbacher, 1906 in the tribe Obrimini, → 3.2.1). Eggs are more or less bullet-shaped and increasingly elongated and have a trilobate micropylar plate, which is principally shaped like an inverted “Y” and has two posterolateral expansions, and there is a distinct median line Only few genera possess hairy structures on the egg capsule and operculum (→ 3.2.2 tribe Tisemenini n. trib., Figs. 67–71), the eggs of all other genera being destitute of hairs of setae (Figs. 47– 56, 76–77). They are laid into soil or a substrate, for which ♀♀ make use of their beak-like ovipositor. Molecular analyses based on the two genera Aretaon Rehn & Rehn, 1939 and Sungaya Zompro, 1996 molecular analyses have confirmed the monophyly of the tribe Obrimini (Bradler, 2009: 97), but the monophyly of the subfamily as a whole deserves further evaluation. Based on the two genera Trachyaretaon Rehn & Rehn, 1939 and Pterobrimus Redtenbacher, 1906 the molecular study by Bradler et al. (2015: 2) also provides strong for the monophyly of Obriminae (Maximum likelyhood bootstrap [MLB] = 98, Bayesian posterior probability [BPP] = 0,99). Bradler (2009: 97) stated that no confirmed autapomorphy was currently known for Obriminae. However, the secondary lack of a central sensory-area on the profurcasternum (Figs. 41–42, 64, 75) and the elongated tarsi, which have the basitarsus considerably longer than the 2nd tarsomere, are quite certainly autapomorphies of Obriminae (Fig. 97b). Zompro (2004) regarded the Obriminae as a monophylum based on the presence of paired sensory-areas on the prosternum, but at the same time interpreted this feature as a synapomorphy of Dataminae + Obriminae (Zompro, 2004: Fig. 119). Close relation of the Obriminae with the Heteropteryginae is supported by synapomorphies such as the presence of a beak-like secondary ovipositor in ♀♀, increasingly elongated antennae and presence of a medio-apical spine on the area apicalis of the tibiae. Consequently, Obriminae + Heteropteryginae are the sister-group of Dataminae (Zompro, 2004: 192; Bradler, 2009: 97 → Fig. 97a–b). The monophyly of Zompro’s Eubulidini is not supported as it contains genera that differ fundamentally in a variety of characters. Zompro (2004: 205) distinguished the Eubulidini from the other two tribes of Obriminae merely by the lack of composite posterior mesonotal and metanotal spines. As a result Eubulidini includes genera with and without a raised triangular area on the mesonotum, with and without strongly developed or reduced body armature, with and without a medio-apical spine on the area apicalis, with and without complex, composite, branched or antler-like pronotal spines and taxa that either have a strongly shortened or much elongated ovipositor in ♀♀ and taxa with shortened, barrel-shaped and setose eggs as well as ones that have increasingly elongated and bullet-like eggs that lack any traces of setae or hairy structures. Furthermore, since Eubulidini contains the genus Theramenes Stål, 1875 the tribe would have to be regarded a synonym of Theramenini Karny, 1923.


HENNEMANN ET AL.

The monophyly of Miroceramiini as treated by Zompro (2004: 203) is also not supported, since the second genus included, the Philippine Mearnsiana Rehn & Rehn, 1939, differs fundamentally from the Wallacean Miroceramia Günther, 1934 by a good number of characters. The only feature that Zompro (2004: 203) used for separating Miroceramiini from the remainder of Obriminae is the short mesothorax, which he stated to be quadrate and not longer than wide. This is however not true for Mearnsiana and this genus differs furthermore from Miroceramia by the lack of wings, much shorter tarsi, which are hardly 1/3 the length of the corresponding tibiae (> ½ the length of the tibiae in Miroceramia), straight ovipositor of ♀♀ (strongly down-curving in Miroceramia) and ovoid, not curved eggs (very elongate and conspicuously curved in Miroceramia). Consequently, Mearnsiana is not closely related to Miroceramia but instead shows affinity to genera here placed in the Theramenes-group of the tribe Obrimini (→ 3.2.1). Consequently, it is hereby re-transferred back to that tribe.

Tribes included: 1. Obrimini Brunner v. Wattenwyl, 1893: 98 (Type-genus: Obrimus Stål, 1875b: 49). 2. Tisamenini n. trib. (Type-genus: Tisamenus Stål, 1875a: 59, 92). 3. Miroceramiini Zompro, 2004: 203 (Type-genus: Miroceramia Günther, 1934a: 283).

Key to the tribes of Obriminae Brunner v. Wattenwyl, 1893 Adults 1. 2.

-

Apterous; mesonotum > 1.5x longer than pronotum; tarsi short, < 1/3 the length of corresponding tibiae . . . . . . . . . . . . . . . . . 2 Winged (Figs. 72–74); mesonotum very short and only 1.2x longer than pronotum; tarsi very elongate and > ½ the length of corresponding tibiae; Sulawesi and Seram . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Miroceramiini Medium to large insects (body length ♂♂ > 40.0 mm, ♀♀ > 55.0 mm); no raised triangular area in anterior portion of mesonotum; pronotum unarmed or with simple spines; medio-apical spine of area apicalis distinct; antennae elongate and longer than fore legs; Philippines, Palawan, Borneo, Talaud Islands and Palau Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Obrimini Smaller, stocky insects (body length ♂♂ < 50.0 mm, ♀♀ < 70.0 mm); mesonotum with a ± elevated triangular area in anterior portion; pronotum with composite, dichotomic, antler-like spines or denticulate crests in anterior half; medio-apical spine of area apicalis indistinct or obsolete; antennae shorter than fore legs; Philippines & Borneo . . . . . . . . . . . . . .Tisemenini n. trib.

Eggs 1. 2. -

Capsule ± bullet-shaped, up to 4x longer than wide; destitute of setae or hairy structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Capsule barrel-shaped, < 1.6x longer than wide; with setae or hairy structures in anterior portion of capsule and on operculum (Figs. 67–71) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Tisamenini n. trib. Capsule curved in lateral aspect with dorsal surface convex and ventral surface concave (Figs. 76–77). . . . . . . . Miroceramiini Capsule not curved (Figs. 47–56) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Obrimini

3.2.1. Tribe Obrimini Brunner v. Wattenwyl, 1893 Type-genus: Obrimus Stål, 1875b: 49. Obrimini Brunner v. Wattenwyl, 1893: 98. Redtenbacher, 1906: 36 (in part). Günther, 1953: 552 (in part). Klante, 1976: 74 (in part). Bradley & Galil, 1977: 199 (in part). Bragg, 1998: 31 (in part). Sellick, 1998: 208, fig. 8 (in part). Bragg, 2001: 102 (in part). Bradler, 2009: 97 (in part). Obriminae, Rehn & Rehn, 1939: 400 (in part). Kevan, 1982: 381. Obriminidae, Bruner, 1915: 228 (in part). Eubulidini Zompro, 2004: 205. n. syn. Otte & Brock, 2005: 31 (in part).



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Diagnosis (♂, ♀): Medium-sized to large (body length ♂♂ 44.0–80.0 mm, ♀♀ 57.0–115.0 mm), moderately robust to slender, always apterous Obriminae. Sexual dimorphism distinct with ♂♂ much more slender and often differently coloured than ♀♀. Colouration ranging from various shades of grey and brown to green. Head ranging in shape from flattened and roundly rectangular to having the vertex strongly conically raised; vertex ± spinose. Antennae slender, ± filiform and considerably longer than fore legs; with 24–28 antennomeres. Median antennomeres elongated and much longer than wide. No gula. Pronotum ranging from sub-rectangular to subtrapeziform in shape and usually longer than wide (exception Mearnsiana Rehn & Rehn, 1939), armature ranging from minutely tuberculose to spinose; usually a pair of simple pre-medial spines present. Mesothorax elongate and at least 1.5x longer than prothorax, ± widening or widened posteriorly. Meso- and metanotum ranging from minutely granulose or tuberculose to spinose, with or without a pair of simple or composite posterior mesonotal spines (a posteromedian swelling in certain ♂♂ of Theramenes Stål, 1875). Mesopleurae slender, tuberculose to spinose (somewhat expanded in ♀♀ of Mearnsiana). Metapleurae ± expanded in posterior portion, often with a distinct supra-coxal spine. Meso- and metasternum simple or with a fine longitudinal median carina, smooth to sparsely tuberculate. Metasternum occasionally with prominent invaginations (= foramina) near lateral margins. Median segment shorter than metanotum. Abdomen of ♀♀ with terminal five segments ± tapering, tergites II–VII ranging from sub-rectangular to transverse, mostly unarmed but II–V occasionally with single spines; lateral margins often somewhat expanded posteriorly and rarely with single minute teeth. Abdomen of ♂♂ slender with segments II–VII roughly equal and uniform in width, armature as in ♀♀; segments VII–X broadened and giving apex of abdomen a club-like appearance. Sternites ranging from smooth to very sparsely tuberculose or spinulose. Praeopercular organ on sternum VII of ♀♀ a posteromedian invagination or two tubercles near posterior margin. Ovipositor of ♀♀ long and either straight if seen in lateral aspect or up-curving; subgenital plate usually projecting over apex of epiproct. Poculum of ♂♂ strongly convex, bulgy and cup-shaped. Legs mostly elongated, metafemora ± incrassate in members of the Theramenes-group. Profemora distinctly curved and compressed basally. All carinae of femora to a variable degree set with teeth or spines; tibiae either unarmed or with single teeth on ventral carinae. Area apicalis with a distinct medio-apical spine. Tarsi moderately elongate and less than ½ the length of corresponding tibiae; basitarsi longer than following two tarsomeres combined. Eggs (Figs. 47–56): Capsule ± bullet shaped and ranging from barrel to bullet-like in general shape, up to 4x longer than wide. Polar-area mostly rounded but occasionally evaginated and anterior portion of capsule may be conspicuously narrowed. Capsule surface minutely punctured and mostly plain pale to dark grey; sometimes with anterior portion darker than rest of capsule. Micropylar plate tri-lobate and variable in shape. Median line distinct. Operculum flat to slightly conical and ± circular in outline; may be inserted into capsule at a positive angle of up to 45°. Differentiation: The various genera within this tribe characteristically have a more or less slender appearance with the mesothorax considerably longer than the pronotum. Some genera have composite posterior meso- and metanotal spines and possess distinctive metasternal features not being found in the other two tribes. From the Tisemenini n. trib. these genera differ by: the generally larger size and more slender appearance; lack of a raised triangular area on the mesonotum; obsolete or at best very faint longitudinal median carina on the meso- and metasternum; simple armature of the pronotum; longer and more slender antennae, which have the median antennomeres considerably elongated and longer than wide, and relatively longer tarsi of both sexes. Females furthermore differ by the longer ovipositor and not conspicuously broadened metathorax. Males may also be distinguished from those of the Tisamenini n. trib. by the larger, much more bulgy poculum. The eggs of Obrimini frequently differ by being more elongate and lacking the setae or hairy structures on the capsule and operculum seen in Tisamenini n. trib.. From the tribe Miroceramiini both sexes of Obrimini can readily be separated by the lack of wings; longer mesothorax, which is at least 1.5x longer than the pronotum, and shorter tarsi, which are by far less than half the length of the corresponding tibiae. Females furthermore differ by having the ovipositor either straight or up-curving (down-curving in Miroceramiini) and the eggs are never curved in the lateral aspect. Comments: The tribe Obrimini may be divided into three marked generic groups. Several of the genera here included in Obrimini were placed in either Eubulidini or Miroceramiini by Zompro (2004) and are here retransferred. The distinction of the tribes as proposed by Zompro (2004: 200) is questionable (→ see comments under 3.2) and the Eubulidini in particular are not a natural assemblage, hence this tribe is here synonymised with Obrimini (n. syn.). Only four of the genera formerly placed in Eubulidini by Zompro (2004: 205) form a clade here recognized as the Tisamenini n. trib. (→ 3.2.2), i.e. Tisamenus Stål, 1875, Ilocano Rehn & Rehn, 1939,


HENNEMANN ET AL.

Pterobrimus Redtenbacher, 1906 and Hoploclonia Stål, 1875, the remainder genera are all re-transferred to Obrimini. The true obrimoids are here termed the Obrimus-group (Figs. 21–28, 47–53) and are the dominant generic group of the tribe, by far containing the largest number of known taxa all of which are distributed in the Philippines, Palawan and Borneo. The six genera here placed in the Obrimus-group form a rather diverse but at the same time a monophyletic assemblage. This is supported by a phylogenetic study published by Bradler (2009: 97), who stated that based on the two genera Aretaon Redtenbacher, 1906 and Sungaya Zompro, 1996 the Obrimini was monophyletic. Members of the Obrimus-group are characterised by the presence of composite posterior meso- and metanotals and most taxa are moderately slender and have a distinctly granulose or rugulose and to a variable degree spinose body surface and head. The head ranges in shape from subquadrate and flattened (Brasidas Rehn & Rehn, 1939 and Euobrimus Rehn & Rehn) over roundly convex and fairly globose (Obrimus Stål, 1875) to having the vertex strongly conically raised (Aretaon Redtenbacher, 1906, Trachyaretaon Rehn & Rehn, 1939 and Sungaya Zompro, 2006). The ovipositor of ♀♀ is long and both the subgenital plate and epiproct are roughly straight if seen from the lateral aspect. The poculum of ♂♂ is large, bulgy and strongly convex. All genera have fairly long and slender legs with the metafemora never incrassate. The supposedly more derived genera possess distinctive structures in the lateral portions of the metasternum (Figs. 38–40). In the genus Obrimus the lateral portions of the metasternum are somewhat invaginated and the metapleurae are folded under so that that this area appears to be partially covered by a flap, giving the appearance of slight slit-like invaginations along the lateral margin (Rehn & Rehn, 1939: 402). In Brasidas and Euobrimus the metasternum has deep, open pits or holes which are somewhat removed from the lateral margins. While these pits are semi-cingulate in Brasidas they are completely cingulate in Euobrimus, but the distinction of these two very closely related genera based on this character alone appears questionable and warrants further evaluation. Hence it is likely that Brasidas and Euobrimus are synonymous. Regardless, these metasternal foramina are obviously apomorphic. No such specializations of the metasternum are seen in Aretaon, Trachyaretaon and Sungaya or any other members of the tribe all of which possess the “normal type” of metasternum. The eggs are bullet-shaped and usually considerably longer than wide, but may have the anterior portion narrowed and the polar-area roundly evaginated (Obrimus and Sungaya). In these two genera the capsule is less elongate and much more globose. While the operculum is somewhat convex to very slightly conical in Aretaon and Trachyaretaon it is entirely flat in the other genera. A second group contains only the fairly distinctive Philippine genus Stenobrimus Redtenbacher, 1906, hence here termed the Stenobrimus-group (Figs. 31–33, 54). The conically elevated back of the head capsule and lack of invaginations on the metasternum suggest that the most closely related genera are Aretaon Redtenbacher, 1906, Trachyaretaon Rehn & Rehn, 1939 and Sungaya Zompro, 1996. Stenobrimus however readily differs from these and all other genera of the Obrimini by the very slender body and legs, distinctive body armature, very long and filiform antennae, which have the median antennomeres in particular extremely elongated, as well as the morphology of the eggs. The metasternum is plain unlike some genera of the Obrimus-group. The eggs have the capsule fairly short with the polar-area conically evaginated, posterolateral expansions of the micropylar very long and surrounding most of the capsule surface and the operculum inserted in the capsule at an angle of almost 45° (Figs. 54a–b). While a conically evaginated polar-area is also seen in the eggs of Sungaya (Figs. 53A–b) and the genus Obrimus Stål, 1875, the displaced operculum is unique within the entire subfamily Obriminae. From the three genera mentioned above Stenobrimus furthermore differs by the simple paired posterior mesonotal and metanotal spines, lack of any composite spines or armature, presence of a distinct spine at the posterolateral angle of abdominal tergites II–VI, entirely smooth tibiae, and short ovipositor of ♀♀ in which the epiproct is shorter than the anal segment and the apex of the subgenital plate multi-dentate (Fig. 33). All of these features are unique amongst the entire subfamily Obriminae and at least the slender body and legs, the distinctive genital morphology and displaced operculum of the eggs appear to be autapomorphies of Stenobrimus. The exact position and true relationships of Stenobrimus are as yet difficult to define, but the genus appears to represent a rather derived lateral specialization of the Obrimini. This is also indicated by the dietary habits, since at least the type-species Stenobrimus bolivari Redtenbacher, 1906) is known to feed predominantly or perhaps even exclusively on ferns (Polypodiaceae). In contrast, most other representatives of the tribe appear to be fairly polyphagous. The third generic group, the Theramenes-group (Figs. 29–30, 34–37, 55–56), contains the remaining four genera. Two of these are distributed in the Philippines (Eubulides Stål, 1875 and Mearnsiana Rehn & Rehn, 1939), one is represented in the Philippines and the Talaud Islands (Theramenes Stål, 1875) and one genus is only known



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from the Palau Islands (Heterocopus Redtenbacher, 1906). The latter genus represents the most eastward distribution of the entire family Heteropterygidae. The genera here placed in the Theramenes-group are characterized by the increasing simplicity of the body armature, lack of posterior mesonotal and metanotal spines, flattened and weakly armed to unarmed head, and increasingly thickened metafemora (♂♂ in particular). The metasternum is plain unlike some genera of the Obrimus-group. The eggs of most genera in this group are very elongate and up to 4.5x longer than wide (e.g. Figs. 51a–b and 55–56a–b). The genus Theramenes Stål, 1875 is characteristic for the specialization of the thoracic armature and strongly incrassate metafemora of ♂♂ (Fig. 35), which readily distinguishes this genus from all other representatives of the entire subfamily. Hence, the posteromedian hump-like elevations of the meso- and metanotum are certainly an autapomorphy of Theramenes. Females on the other hand, show a much closer affinity to Eubulides and the other genera of the Theramenesgroup, and they share with the former genus the strongly up-curving ovipositor (Fig. 34), which might be a synapomorphy of Theramenes + Eubulides. Heterocopus and Mearnsiana have the ovipositor of ♀♀ straight if seen laterally, but while the former genus appears to be more closely related to Eubulides and shows closer affinity to genera of the Obrimus-group, the latter genus has more characteristics in common with Theramenes, i.e. green colouration (not always true for Theramenes), ♂♂ having the composite posterior mesonotal and metanotal spines very blunt, rounded at the point and contrasting black, the abdominal tergites short and transverse and the poculum very large and bulgy, and ♀♀ having an entirely unarmed abdomen. The unarmed abdomen is likely to be a synapomorphy of Theramenes + Mearnsiana. The eggs of Mearnsiana however rather resemble those of Heterocopus than Theramenes in being fairly short and only about 2.5x longer than wide, instead of remarkably elongate. The short eggs of these two genera show affinity to certain genera of the Obrimus-group, i.e. Brasidas and Euobrimus. As a result the intergeneric relationships within the Theramenes-group certainly deserve more extensive study and remain as yet unclarified. Distribution: Philippines, Palawan and Borneo with single representatives on the Talaud and the Palau Islands.

Genera included: Obrimus-group: 1. Aretaon Rehn & Rehn, 1939: 419. (Type-species: Obrimus asperrimus Redtenbacher, 1906: 41, by original designation). 2. Brasidas Rehn & Rehn, 1939: 430. (Type-species: Brasidas samarensis Rehn & Rehn, 1939: 432, by original designation). 3. Euobrimus Rehn & Rehn, 1939: 445. (Type-species: Euobrimus atherura Rehn & Rehn, 1939: 446, by original designation). 4. Obrimus Stål, 1875b: 49. (Type-species: Phasma (Acanthoderus) bufo Westwood, 1848: 77, by subsequent designation of Kirby, 1904: 398). 5. Sungaya Zompro, 1996: 450. (Type-species: Sungaya inexpectata Zompro, 1996: 450, by original designation). 6. Trachyaretaon Rehn & Rehn, 1939: 422. (Type-species: Obrimus echinatus Stål, 1877: 68, by original designation).

Stenobrimus-group: 7. Stenobrimus Redtenbacher, 1906: 37. (Type-species: Stenobrimus bolivari Redtenbacher, 1906: 37, by monotypy).

Theramenes-group: 8. Eubulides Stål, 1877: 68. (Type-species: Eubulides alutaceus Stål, 1875: 68, by monotypy). 9. Heterocopus Redtenbacher, 1906: 42. (Type-species: Heterocopus leprosus Redtenbacher, 1906: 42, by monotypy). 10. Mearnsiana Rehn & Rehn, 1939: 458. (Type-species: Mearnsiana bullosa Rehn & Rehn, 1939: 459, by original designation) = Hennobrimus Conle, 2006: 44. (Type-species: Hennobrimus hennemanni Conle, 2006: 45, by original designation). n. syn.* 11. Theramenes Stål, 1875b: 46. (Type-species: Eurycantha olivacea Westwood, 1859: 65, by monotypy).** *Comment: The genus Mearnsiana Rehn & Rehn, 1939 was originally described from a ♀ nymph in USNM. Since immature specimens differ considerably from the adult insects in being various shades of brown and grey, much more stocky in appearance with the mesothorax relatively shorter and more distinctly expanded laterally and having the body and leg-armature much more developed, two synonyms have been described recently. Adult insects of Mearnsiana are of a fairly unusual colouration for the tribe Obrimini, being bright green with ♂♂ having the lateral surfaces of the abdomen bright red. Hence, Hennobrimus hennemanni Conle, 2006 and Trachyaretaon manobo Lit & Eusebio, 2005 both described from the island of Mindanao (Philippines) represent synonyms of Mearnsiana bullosa Rehn & Rehn, 1939 (n. syn.).


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**Comment: Theramenes olivaceus (Westwood, 1859) was originally described from “Ceylon” but there can be no doubt the typelocality is wrong (Günther, 1934b: 76; Rehn & Rehn, 1939: 414). All subsequent records have been from the Talaud Islands north of Sulawesi. Theramenes dromedarius Stål, 1877 was described from the Philippines. Günther (1935: 75) erroneously synonymised T. dromedarius with T. olivaceus and doubted the type-locality given by Stål (1877: 68). Rehn & Rehn (1939: 414) removed T. dromedarius from synonymy with T. olivaceus and Zompro & Eusebio (2001: 23) synonymised Stål’s species yet again. Examination of the ♂ holotypes of both species in NHMUK and NHRS however shows that T. olivaceus and T. dromedarius are definitely two distinct species, which differ considerably in a number of morphological characteros of both sexes. T. olivaceus is restricted to the Talaud Islands, ♂♂ are dull green, have abdominal segments II–VI slightly but gradually shortening and the metapleurae armed with several blunt tubercles. T. dromedarius is from the Philippines, ♂♂ being brown, having abdominal tergites II–VI very unequal in length with II almost twice the length of III and IV, V and VI much shortened and transverse and III–IV with the posterior margin strongly concave, and the metapleurae armed with several prominent spines. In ♀♀ of T. dromedarius the meso- and metapleurae are strongly spinose, whereas they are merely bluntly tuberculate in T. olivaceus. Males of T. dromedarius are very similar to those of the Philippine T. mandirigma Zompro & Eusebio, 2002 only differing by the larger size, lack of an anterior swelling in the anterior portion of the mesonotum and more prominent armature of the meso- and metapleurae. Consequently, T. dromedarius Stål, 1877 is here reestablished as a valid species (rev. stat.). The genus now contains four valid species, the type-species T. olivaceus (Westwood, 1859) from the Talaud Islands, and three species distributed throughout the Philippines, i.e. T. dromedarius Stål, 1877 rev. stat. (presumably from Mindanao Island), T. exiguus Hennemann & Conle, 2003 (Panay Island) and T. mandirigma Zompro & Eusebio, 2002 (Cebu Island).

3.2.2. Tribe Tisamenini n. trib. Type-genus: Tisemenus Stål, 1875a: 59, 92. Obrimini Brunner v. Wattenwyl, 1893: 98 (in part). Redtenbacher, 1906: 36 (in part). Günther, 1953: 552 (in part). Klante, 1976: 74 (in part). Bradley & Galil, 1977: 199 (in part). Bragg, 1998: 31 (in part). Sellick, 1998: 208, fig. 8 (in part). Bragg, 2001: 102 (in part). Bradler, 2009: 97 (in part). Eubulidini Zompro, 2004: 205 (in part). Otte & Brock, 2005: 31 (in part).

Diagnosis (♂, ♀): Small to moderately sized (body length ♂♂ 25.0–50.0 mm, ♀♀ 35.0–68.0 mm) exceptionally apterous Obriminae. General form rather stocky and robust. Sexual dimorphism distinct with ♂♂ much more slender and more heavily armed than ♀♀. In ♂♂ thorax considerably broader than abdomen and in ♀♀ metathorax broadest part of body. Colouration various shades of brown and grey. Head roundly rectangular, vertex usually armed either with spines, tubercles or with dentate crests. Antennae robust, ± perlamorph and at best equal in length to fore legs; with 22–26 antennomeres. Median antennomeres short and at best 3x longer than wide. No gula (exception Hoploclonia Stål, 1875). Pronotum varying from sub-rectangular or sub-trapeziform to rather broadly transverse; anterior portion usually with a pair of large often dichotomic or antler-like spines, compound spines, granulose carinations or dentate crests. Mesonotum fairly short and at best 3x longer than pronotum in ♂♂ of some taxa, usually much shorter in ♀♀. Mesonotum with an elevated triangular area in anterior portion which have the anterior angles variously modified and ranging from indistinct nodes or tubercles to large serrate or denticulate, laterally expanded crests. Posterior portion of mesonotum and entire metanotum with a distinct longitudinal median carina or keel. Lateral margins of mesonotum ranging from tuberculate to distinctly spinose. Meso- and metapleurae ranging from almost smooth over denticulate to prominently spinose, with large often composite supra-coxal present in certain taxa; metapleurae strongly widening towards posterior. Meso- and metasternum with a distinct and acute longitudinal median carina or keel; the latter without foramina. In one case (Pterobrimus Redtenbacher, 1906, Fig. 61) with small, scale-like tegmina. Median segment transverse and usually at least with a trace of a longitudinal median carina. Abdomen broad and ± narrowing towards the apex in ♀♀, segments II–VII roughly parallel-sided in ♂♂. Tergites much wider than long in ♀♀, subquadrate and at best very slightly longer than wide in ♂♂; either unarmed or with various elements of armature. Praeopercular organ of ♀♀ obsolete or represented by a pair of low median tubercles or swellings on sternum VII. Ovipositor of ♀♀ short, either formed by an elongated subgenital plate and epiproct (Tisamenus Stål, 1875 and Ilocano Rehn & Rehn, 1939, Fig. 65), or REVISION OF HETEROPTERYGINAE


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an apically elongated anal segment (Hoploclonia Stål, 1875, Fig. 66). Poculum of ♂♂ fairly small and roundly cup-shaped. Legs fairly short and ranging from slender to robust; the metafemora often incrassate. All carinae of femora to a variable degree set with denticles or spines. Tibiae somewhat shorter than corresponding femora and at best with minute denticles or teeth on two outer ventral carinae of meso- and metatibiae. Medio-apical spine of area apicalis obsolete or indistinct, usually most decided on metatibia (only present on all tibiae in Hoploclonia). Tarsi fairly short with basitarsus roughly equal in length to tarsomeres II and III combined. Eggs (Figs. 67–71): Small, barrel-shaped with dorsal surface strongly convex; at best 1.6x longer than wide. Capsule and operculum to a variable degree set with setae or irregular hairy structures, or with net-like sculpturing. Operculum either flat or with a rim of hairy structures. Micropylar plate simply Y-shaped. Differentiation: Members of this tribe differ from those of Obrimini by: the generally smaller size and more stocky appearance (Figs. 57–63); raised triangular and variously modified area of the mesonotum; distinct longitudinal median carina or keel of the meso- and metasternum; complex armature of the pronotum, which includes composite, dichotomic or antler-like spines or denticulate crests; shorter and more robust antennae and relatively shorter tarsi of both sexes. Females furthermore differ by the comparatively shorter ovipositor and broadened metathorax, which is the widest part of the body. Males may also be distinguished from those of the tribe Obrimini by the smaller, considerably less bulgy poculum. The eggs frequently differ by being less elongate and possessing setae or hairy structures on the capsule and operculum (Figs. 67–71). Comments: The four genera of this tribe are here interpreted as the basalmost forms amongst the subfamily Obriminae with the two very closely related and exclusively Philippine Tisamenus Stål, 1875 and Ilocano Rehn & Rehn, 1939 being more basal than the Bornean Hoploclonia Stål, 1875. The latter genus is the sister-group of Tisamenus + Ilocano + Pterobrimus, differing by the distinctive morphology of the ♀♀ ovipositor, which is formed by the elongated subgenital plate and elongation of the anal segment, with the small epiproct entirely concealed by the anal segment. The other three genera as well as all genera of the tribe Obrimini and subfamily Heteropteryginae have the ovipositor formed by the subgenital plate and an elongated epiproct. Bradler (2009: 97) was unsure about the evolutionary state of this feature, but the most likely scenario seems to be that it was evolved only in Hoploclonia, hence most certainly is an autapomorphy of this genus. Furthermore, Hoploclonia is the only genus of the entire Heteropterygidae which has a gula (see also Bradler, 2009: 66, Table 1). While the medio-apical spine of the area apicalis is obsolete on the pro- and mesotibiae of Tisamenus and Ilocano it is well developed on all tibiae in Hoploclonia. Since it is missing in the subfamily Dataminae, which is here interpreted as the basalmost clade of the Heteropterygidae family, this must be an apomorphic character and places Hoploclonia as the most derived genus amongst Tisemenini n. trib.. This would however mean that the medio-apical spine of the area apicalis was evolved independently in Hoploclonia, the tribe Obrimini and the subfamily Heteropteryginae, hence might be a homoplastic character trait. The small general size, stocky and robust body, fairly short tarsi, short and fairly robust antennae, strict lack of wings and predominantly ground-dwelling habits of the four genera here assembled in the tribe Tisamenini n. trib., resemble the subfamily Dataminae. This also applies to the hairy structure of the egg capsule and operculum, which are entirely lacking in Obrimini and Miroceramiini, while specialized hook-like hairs are typical for Dataminae. All of these characters are plesiomorphic and support the basal position of Tisamenini n. trib. within the Obriminae here revealed (Fig. 97b). The raised triangular and diversely modified area in the anterior portion of the mesonotum and armature of the pronotum, which includes composite, dichotomic or antler-like spines or denticulate crests, appear to be autapomorphies of Tisamenini n. trib. and readily distinguish this tribe from Obrimini. Rehn & Rehn (1939: 464) erroneously synonymized Tisamenus with Hoploclonia, stating that the characters used to separate these two genera by Stål (1875) and Redtenbacher (1906) were insufficient and subject to too great variation to be of diagnostic value for genera. Zompro (2004: 207) re-established Tisamenus and merely distinguished this genus from Hoploclonia by the flattened and longitudinally carinate mesonotum and eggs having the operculum not inserted in the capsule “at an angle”. While the latter character holds true for all taxa here examined, the previous feature is only partly true, a median carina on the mesonotum being also present in Hoploclonia. Bradler (2009: 97) was the first author to determine the true generic difference between these two genera, i.e. the remarkably different morphology of the ♀♀ ovipositor with that of Hoploclonia being unique in the entire family Heteropterygidae (→ see above). Furthermore, the distribution of these two genera clearly separates them, Hoploclonia being endemic on Borneo and Tisamenus restricted to the Philippines.


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The genus Pterobrimus Redtenbacher, 1906 (Fig. 61) is questionable in some aspects and the original placement has already been doubted by Rehn & Rehn (1939: 463). Zompro (2004: 210) placed it in his Eubulidini and merely mentioned the distribution was noteworthy. In fact, the type-locality that was given as “Fiji” by Redtenbacher (1906: 43) must be doubted and, although both type specimens in NHMW and ZMUH bear a label stating “Fiji” or “Viti Levu” respectively, is most certainly erroneous. The presence of a beak-like secondary ovipositor in ♀♀, presence of a spine on the area apicalis of all six tibiae, lack of a central sensory area on the profurcasternum and position of the prothoracal foramen at the anterior angle of the pronotum place Pterobrimus in the Obriminae. Moreover, the small size, stocky body (♀♀ in particular), strong anterior pronotals and indicated raised triangular area in the anterior portion of the mesonotum place the genus in Tisamenini n. trib.. As pointed out by Rehn & Rehn (1939: 463) the apical exterior spine of the scapus would affiliate Pterobrimus to the subfamily Dataminae and distinguishes this genus from the three other representatives of Tisamenini n. trib.. Also the denticulate apex of the epiproct of ♀♀ and presence of small, scale-like tegmina in both sexes distinguish Pterobrimus from the other genera of Tisamenini n. trib.. The latter feature in particular is unique within this tribe and makes the systematic position of Pterobrimus somewhat questionable. Since a distribution of a member of the tribe Tisamenini n. trib., subfamily Obriminiae and even family Heteropterygidae in Fiji is more than unlikely, the true distribution of Pterobrimus remains unknown but the genus is likely to originate from the Philippines or perhaps the northern Maluku Islands. It must be hoped that future collections will include fresh material of Pterobrimus to confirm the true distribution of this peculiar genus. Distribution: Philippines and Borneo. So far not recorded from the intervening Palawan. Genera included: 1. Hoploclonia Stål, 1875b: 8. (Type-species: Acanthoderus gecko Westwood, 1859: 52, by original designation). 2. Ilocano Rehn & Rehn, 1939: 460. (Type-species: Ilocano hebardi Rehn & Rehn, 1939: 461, by original designation). 3. Pterobrimus Redtenbacher, 1906: 43. (Type-species: Pterobrimus depressus Redtenbacher, 1906: 43, by monotypy). 4. Tisamenus Stål, 1875a: 50, 92. (Type-species: Tisamenus serratorius Stål, 1875a: 92, by subsequent designation of Kirby, 1904: 399).

3.2.3. Tribe Miroceramiini Zompro, 2004 Type-genus: Miroceramia Günther, 1934a: 283. Miroceramiini Zompro, 2004: 203. Otte & Brock, 2005: 31.

Diagnosis (♂, ♀): Medium sized (body length ♂♂ 46.0 mm, ♀♀ 70.0–73.0 mm) winged Obriminae with moderate sexual dimorphism; ♂♂ smaller with abdomen much more slender than ♀♀. General form rather robust with the mesothorax conspicuously shortened. Colouration various shades of brown, ochre, black and green. Head roundly rectangular, indistinctly longer than wide, vertex flattened and spinose. Antennae long, slender and considerably longer than fore legs; with 26 antennomeres. All antennomeres distinctly longer than wide. No gula. Pronotum trapezoidal with anterior margin much narrower than posterior margin; surface spinose and with a prominent pair of pre-medial spines. Mesothorax short and only about 1.2x longer than prothorax, sub-quadrate in ♂♂ and somewhat transverse in ♀♀. Mesonotum minutely spinose and with somewhat enlarged anterior and antero-lateral spines; no composite posterior mesonotals. Mesopleurae strongly expanded medially, convex and armed with several prominent, long spines. Metanotum smooth; metapleurae strongly expanded and armed with prominent spines. Meso- and metasternum unarmed, the latter without foramina. Tegmina and alae well developed, the tegmina roughly 2/3 the length of the alae and the latter reaching to abdominal tergum VI or VII. Tegmina elongate, broadened and strongly convex sub-basally, the sub-basal hump tuberculose and armed with two distinct spines. Alae with anal region plain ochre and very slightly translucent. Median segment roughly equal in length to metanotum; smooth. Abdomen of ♀♀ with segments II–IV roughly equal in width and about 2x wider than long and remaining segments gradually tapering towards the apex; tergites II–IV smooth dorsally, V–X rugose, all tuberculose laterally and II–VII with three blunt teeth posterolaterally; tergites VI–X with a longitudinal median carina. Abdomen of ♂♂ almost parallel-sided with segments II–VII somewhat wider than long and almost equal in width; tergites all rugulose with the posterolateral angles acute and in II–V armed with a slender spine; VII–X REVISION OF HETEROPTERYGINAE


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carinate longitudinally and IX somewhat deflexed laterally. Sternites II–VII of both sexes irregularly carinate and sparsely rugulose. Praeopercular organ on sternum VII of ♀♀ indistinct. Ovipositor of ♀♀ long and strongly down-curving, the epiproct shorter than the subgenital plate with the apex obtuse; apex of subgenital plate acutely pointed. Poculum moderately convex and cup-shaped. Legs fairly long with the metafemora slightly (♀♀) or decidedly incrassate (♂♂). Profemora gently curved and compressed basally. All carinae of femora multi-dentate, ventral surface smooth. Tibiae with the apex roundly swollen dorsally and all with a very few indistinct teeth. Area apicalis with a distinct medio-apical spine. Tarsi very long and more than ½ the length of corresponding tibiae; basitarsi elongate and longer than following two tarsomeres combined. Eggs (Figs. 76–77): Elongate, 2.6x longer than wide, bullet-shaped with the polar portion somewhat narrowed and curved if seen in lateral aspect with the dorsal surface convex and the ventral surface concave. Capsule minutely punctured, plain dark grey. Micropylar plate elongate and covering most of dorsal egg-surface, the posterior expansions almost straight and directed towards polar-area. Median line distinct and almost reaching to apices of posterior expansions of plate. Operculum flat, slightly oval and with a central pit; inserted into capsule at an angle of about –45°. Differentiation: The only genus contained in this tribe differs from all other members of the Obriminae by the presence of wings (Figs. 72–74), very short and medially expanded mesothorax, which is hardly longer than the pronotum, and very elongate tarsi, which are more than half as long as the corresponding tibiae. Females furthermore differ by the conspicuously down-curving ovipositor and eggs can be distinguished from those of other Obriminae by being noticeably curved, the capsule being convex dorsally and concave ventrally (Fig. 77). Comments: Miroceramiini was established by Zompro (2004: 203) to comprise the Wallacean Miroceramia Günther, 1934 and the Philippine Mearnsiana Rehn & Rehn, 1939. The main key-character on which Zompro based the tribe was the short mesothorax, which he stated to be “transverse to quadrate” (Zompro, 2004: 201) in both genera. This is however only true for Miroceramia, the mesothorax being considerably longer than the pronotum in Mearnsiana. The main reason for this false statement was the fact that at that time Mearnsiana was only known from an immature specimen. Adult specimens of Mearnsiana now available show the short mesothorax is only characteristic in nymphs of this genus. Furthermore, there are numerous characters that clearly distinguish Mearnsiana from Miroceramia and place the former genus in close relation to the genera here assembled in the Theramenes-group of the tribe Obrimini (→ 3.2.1). Consequently, Mearnsiana is here removed from Miroceramiini and transferred to the Obrimini. Miroceramia, the only genus now contained in Miroceramiini, differs from all other Obriminae by such fundamental characters that a separate tribe appears justified. Bradler (2009: 97) questioned the placement of this genus in Obriminae. However, the distinct paired sensory-areas of the prosternum (Fig. 75), gently curved and compressed basal portion of the profemora, elongate tarsi and elongated basitarsi, lack of large ventral spines on the metatibiae and egg-morphology clearly place Miroceramia in the Obriminae. All these features do not support a close relationship to Heteropteryginae as suggested by Bradler (2009: 97). The shortened and medially expanded mesothorax resembles Mearnsiana and features such as the broad, roundly rectangular head and fairly incrassate metafemora of ♂♂ indicate a closer relationship to the Theramenes-group of Obriminae, rather than the Obrimusgroup. The only characters that indicate affinity to the latter group are the fairly well developed armature of the head and thorax as well as the presence of teeth along the lateral margins of the abdominal tergites. The most distinctive features of Miroceramia, i.e. the presence of wings, strongly shortened mesothorax, which is hardly longer than the pronotum, very elongate tarsi, which are more than half as long as the corresponding tibiae, in both sexes and the conspicuously down-curving ovipositor of the ♀♀ as well as the longitudinally curved eggs may be regarded as autapomorphies of Miroceramia. This favours the scenario that the wings were gained in Miroceramia and developed independently from the subfamily Heteropteryginae, consequently indicating that the wings were gained twice within the Heteropterygidae family and represent a homoplasy (→ 3.4). The profemora were erroneously stated to be straight in Miroceramia (Zompro, 2004: 203), but in fact they are gently curved and noticeably compressed basally. A note by M.J.D. Brendell on a label attached to a ♂ from Mansuela National Park, Seram in NHMUK states “This insect stridulates from the wing bases” (Bragg, 2001: 102), but this deserves evaluation once live specimens of M. westwoodii (Bates, 1865) become available. If this observation by the collector of the specimen proves to be true, this would mean that not only the wings but even the presence of a stridulatory organ or tympanal area in the basal portion of the alae, which is otherwise typical for the subfamily Heteropteryginae, was developed independently in Miroceramia.


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The derived position of Miroceramia within the Obriminae here suggested is supported by its geographic distribution, which is far off the main distribution of the Obriminae. While the Obriminae have mainly radiated throughout the Philippines and Borneo with only one representative each on the Palau and Talaud Islands, Miroceramia is peculiar to Wallacea and besides having been recorded from Northern Sulawesi is known from as far southwest as the island of Seram in the southern Maluku Islands (→ 6.3). Distribution: Wallacea (N-Sulawesi and Seram). Genus included: 1. Miroceramia Günther, 1934a: 283. (Type-species Miroceramia pterobrimus Günther, 1934a: 283 (= Heteropteryx westwoodii Bates, 1865: 345), by original designation).

3.3. Subfamily Heteropteryginae Kirby, 1896 A complete revision of the Heteropteryginae at the species-level including a detailed diagnosis of the subfamily are presented below (→ 4.) and a more detailed discussion of the inter- and intrageneric relationships of Heteropteryginae is provided at the end of the present work (→ 5.). The subfamily is here interpreted as the most derived subordinate clade of the Heteropterygidae family. The two genera included in this subfamily, i.e. Haaniella Kirby, 1904 and Heteropteryx Gray, 1835, are distributed throughout Sundaland with one representative as far north as Vietnam (Fig. 345). All other taxa are found on Borneo, Sumatra and the Mentawei Islands, Singapore, Peninsular Malaysia and Thailand. The subfamily is not represented on Java and in the Philippines, hence Borneo is the only island where the distributions of Heteropteryginae and Obriminae overlap. The Heteropteryginae are more closely related to the Obriminae rather than the Dataminae (→ 3.4, Fig. 97), which is in compilance with the molecular studies by Bradler et al. (2014, 2015) and based on the two genera Haaniella Kirby, 1904 and Heteropteryx Gray, 1835 here is also strong support for the monophyly of Heteropteryginae (Maximum likelyhood bootstrap [MLB] = 99, Bayesian posterior probability [BPP] = 0,99). This is supported by synapomorphies such as the presence of a beak-like secondary ovipositor in ♀♀, presence of a medio-apical spine on the area apicalis of the tibiae (Bradler, 2009: 97) and increasingly elongated antennae, which are considerably longer than the fore legs with the median antennomeres much elongated. Consequently, Heteropteryginae + Obriminae are considered to be the sister-group of Dataminae (Zompro, 2004: 192; Bradler, 2009: 97; → Fig. 97). Most members of the Heteropteryginae are characterised by their large size with some species achieving body lengths of more than 12 cm, often strongly spinose body and legs and presence of wings in both sexes. Most taxa are brachypterous but ♂♂ of five species possess fully developed alae and elongated tegmina that are more than half the length of the alae. Brachypterous taxa have a stridulatory organ in the basal portion of the alae, which is formed by a number of fused wing-cells positioned between the radial and cubital vein, the former vein being set with numerous minute teeth dorsally. The alae have the anal fan translucent grey to black or pink with all major veins marked darker, hence producing a reticulate appearance. The sensory-areas of the prosternum have been fully secondarily reduced but there is a more or less distinct central sensory-area on the profurcasternum (Figs. 78–83). Sexual dimorphism is distinct, with ♂♂ being considerably smaller and much slenderer than ♀♀, often having the body armature more developed and the colouration different from that of the corresponding ♀♀. The abdominal sternites II–IV of most species have conspicuous lateral invaginations, that are not seen elsewhere within Heteropterygidae and still of unknown function (Figs. 85–92). There is a prominent praeopercular organ on abdominal sternum VII of ♀♀, which is usually represented by a swollen area that bears a pair of spines. As in Obriminae ♀♀ possess a beak-like ovipositor, that is formed by elongation of the subgenital plate and the epiproct. The epiproct shows a variety of differentiation with the apex either bifid or multi-dentate. The legs are strongly armed and in contrast to the Obriminae there are particularly strong spines on the ventral carinae of the metatibiae, which are used by the insects in active defence against predators. The tarsi are very short with the basitarsus being hardly longer than the 2nd tarsomere. All six tibiae bear a distinct medio-apical spine on the area apicalis (Figs. 93–96). The profemora are straight and not compressed or curved basally as in Obriminae or Dataminae. Eggs of Heteropteryginae are very large, barrel- or lemon-shaped and are the largest and heaviest known eggs of the entire Phasmatodea. They usually possess a more or less setose capsule surface and operculum



23

and a very characteristic for the complex shape of the micropylar plate, which is principally saltire or X-shaped having two anterior and two posterior expansions. The median line is completely reduced and lacking. The straight profemora, presence of wings, secondary development of a stridulatory organ in the basal portion of the alae, secondary lack of sensory-areas on the prosternum, lateral invaginations of abdominal sternites II–IV of several taxa as well as the characteristically X-shaped micropylar plate of the eggs and the lack of a median line are herein regarded as autapomorphies of Heteropteryginae.

3.4. Classification of Heteropterygidae Kirby, 1896 The monophyly of Heteropterygidae is fairly well supported and the molecular study of Bradler et al. (2015) show a high Bayesian posterior probability (BPP = 0,99) for the clade (referred to as Heteropteryginae). The sister-taxon of Heteropterygidae has not yet been identified with confirmation but several previous molecular and phylogenetic studies reveal the family Phylliidae Karny, 1923 and the New World family Pseudophasmatidae Rehn, 1904 as likely candidates (e.g. Bradler et al., 2014, 2015). According to Bradler et al. (2015, fig. 2) the Heteropterygidae appear to have diverged from the remaining Euphasmatodea approximately 42.5 mya ago during the early Eocene. This study also provides strong support for the monophyly of the subafmilies Heteropteryginae (Maximum likelyhood bootstrap [MLB] = 99, Bayesian posterior probability [BPP] = 0,99) and Obriminae (Maximum likelyhood bootstrap [MLB] = 98, Bayesian posterior probability [BPP] = 0,99) and furthermore shows Dataminae to be the sister-taxon of Heteropteryginae + Obriminae, which is in compilance with the results here presented (see Fig. 97b). Although Heteropterygidae are very likely to be monophyletic a confirmed autapomorphy is still not known (see Bradler, 2009: 95). The presence of rough sensory-areas on both or either the prosternum and profurcasternum and the presence of hairy structures on the egg-capsule however are very likely to represent autapomorphies but this assumption deserves further evaluation. The stridulatory organ in the alae of most Heteropteryginae and one genus of Obriminae (→ 3.2.3) has been regarded an autapomorphy and a possible character of the ground plan of Heteropterygidae by Bradler (2009: 95), but as already stated by Bradler is not of much significance since by far most members of Obriminae and the entire Dataminae are wingless. A development of wings within the Heteropterygidae is here favoured and certainly the most likely scenario, which is in accordance to the results of Whiting et al. (2003) and Bradler (2009). The arguments for the classification and intra-familiar relationships of Heteropterygidae proposed herein are summarized and discussed in more detail below and shown in Figure 97. The present study reveals the predominantly ground-dwelling Dataminae as the basalmost clade of Heteropterygidae (Fig. 97b), which is compilance with previous molecular and phylogenetic studies (e.g. Bradler, 2009, Bradler et al., 2015). Plesimorphic characters which support the basal position of Dataminae are the primary lack of an ovipositor, strict lack of wings, presence of rough sensory-areas on the prosternum and profurcasternum (Fig. 97b), basally curved and compressed profemora, lack of a medio-apical spine on the area apicalis, short and perlamorph antennae and mostly ovoid to spherical eggs. In most genera the eggs are simply dropped to the ground by the ♀♀, an behaviour which is the most widespread within Phasmatodea and regarded as the plesiomorphic strategy of egg-deposition, as no particular advanced development of the ♀♀ genitalia is necessary (Bradler, 2009; Goldberg et al., 2015). The presence of an area apicalis at the apex of the tibiae is believed to be a plesimorphic character state and part of the ground plan of Phasmatodea (e.g. Bradler, 1999: 43; Tilgner, 2001: 477; Bradler, 2009: 34), since it is also present in other lower Neoptera such as certain basal Embioptera (Tilgner, 2002; Zompro, 2005a) and some Plecoptera (Zompro, 2005). Hence, any further development of this structure, e.g. the presence of a medio-apical spine, as in Obriminae and Heteropteryginae (and members of Phylliinae Kirby, 1904 and Anisacanthidae Günther, 1953), must be regarded as the apomorphic state and its presence in three distant clades suggests that it has been evolved independently in these taxa, hence is homoplasy. Consequently, the lack of a medio-apical spine on the area apicalis in the Dataminae is quite certainly a plesiomorphic state. The presence of sensory-areas on the prosternum and profurcasternum is obviously an autapomorphy of Heteropterygidae, but since either those of the prosternum (Heteropteryginae) or that of the profurcasternum (Obriminae) are reduced, the presence of both in Dataminae must be interpreted as the plesimorphic state within Heteropterygidae. Traditionally the presence of wings has a priori been regarded to be plesimorphic and it was believed that Phasmatodea have lost their wings during the course of their evolution without the possibility of reacquisition. Recent molecular studies


HENNEMANN ET AL.

(Whiting et al., 2003) have however hypothesized an apterous stem species for the whole Phasmatodea and a secondary recovery of wings in several subgroups independently, suggesting the wing did not re-evolve de novo in Phasmatodea, but rather represents a “re-expression” of the basic wing which was lost in ancestral stick-insects and that the wing development pathways are conserved in wingless taxa. Since the invariably wingless Dataminae are here revealed as the basalmost taxon of Heteropterygidae, the presence of wings in the other two subordinate taxa, i.e. Heteropteryginae and one genus of Obriminae, provide good support for the hypothesis of Whiting et al. (2003). Consequently, the winglessness of Dataminae must be regarded as the plesiomorphic state for Heteropterygidae and means the wings were gained independently in Heteropteryginae and one genus of Obriminae, hence their presence in these two taxa represents a homoplasy. This assumption is supported by an undoubtedly apomorphic specialization of the wings in members of the Heteropteryginae, which possess a stridulatory organ in the basal portion of the alae, that is formed by a number of fused wing-cells positioned between the radial and cubital vein (→ 3.3, Fig. 84). Apomorphies of Dataminae are the position of the pronotal foramina, which are not positioned at the anterolateral angles of the pronotum as usual in Phasmatodea but conspicuously displaced towards the posterior, frequent presence of spines on the exterior lateral margin of the scapus and specialization of the hairy structures of the eggs, which have hook-like apices and make them adhesive to various surfaces (→ 3.1, Figs. 11–20). Alongside these morphological characters the basal position of Dataminae is supported by biogeography and the resulting distributional pattern of the subfamily. Although predominantly ground-dwelling and implying likely slow dispersal Dataminae has the by far most widespread distribution of the subordinate taxa of Heteropterygidae. In addition to Sundaland the present day distribution of Dataminae expands to as far north as China, Taiwan and even Japan and as far east as great parts of Wallacea (→ 6.1), thus also covers the entire distributional ranges of the other two subfamilies, i.e. Obriminae and Heteropteryginae, except for the Philippine Islands (Fig. 343). With five of the seven known genera the by far greatest number of known Dataminae is represented in Borneo, which is the only island that harbours representatives of all three subfamilies of Heteropterygidae (Fig. 342). Hence Borneo seems to be the centre of the present day distribution and radiation of the family. The wide distribution of Heteropterygidae, but Dataminae in particular, indicates a high evolutionary age of the clade and perhaps suggests a Gondwanan origin. The Obriminae and Heteropteryginae are characterized by the presence of a medio-apical spine on the area apicalis of the tibiae and presence of a secondary beak-like ovipositor of ♀♀, which is formed by elongation of the epiproct and subgenital plate. Both features are synapomorphies of Obriminae + Heteropteryginae and suggest these two subfamilies are the sister-group of Dataminae (Fig. 97b), which agrees with the results of other authors (e.g. Zompro, 2004, Bradler, 2009, Bradler et al., 2015). The Heteropteryginae are the most derived subordinate clade of Heteropterygidae and well characterized by autapomorphies such as the secondary reduction of the paired sensory-areas on the prosternum, increasingly straight profemora and the X-shaped micropylar plate of the eggs. The presence of wings and a tympanal area (= stridulatory organ) in the basal portion of the alae of Heteropteryginae is undoubtedly apomorphic, but both characters are also present in a single genus of the subfamily Obriminae, i.e. Miroceramia Günther, 1934. Hence, both characters are believed to have been gained two times independently in these taxa. Nevertheless, they are definitely apomorphic character states because the more basal Dataminae are exclusively wingless (→ see above). The central sensory-area of the profurcasternum is fairly well developed in most Bornean taxa of Haaniella Kirby, 1904 but increasingly reduced in members of the muelleri species-group of the genus, that occur throughout Sumatra and Peninsular Malaysia, and akmost fully reduced in Heteropteryx Gray, 1835. The single known species of the latter genus is the only representative of Heteropteryginae, that shows increasing adaption to arboreal habits by ♀♀ being bright green and having the abdominal tergites strongly laterally deflexed, giving the insects a leaflike general appearance and thereby supporting prolific camouflage in the foliage of shrubs and trees. Males of Heteropteryx are fully winged and capable of short active flight, which also supports arboreal habits. The averaging smaller and more stocky, almost always brown Haaniella-species are mostly brachypterous in both sexes and show no particular adaptions to arboreal habits, emphazising their derivation from mainly ground-dwelling ancestors, i.e. the Dataminae. The distribution of Heteropteryginae is restricted to Sundaland with a single species represented in Southern Vietnam (Fig. 345). The Obriminae are the by far most diverse subordinate clade of Heteropterygidae and divide into three distinct tribes, the Obrimini, Tisamenini n. trib. and Miroceramiini (→ 3.2). Autapomorphies of the Obriminae are the



25

secondary reduction of the central sensory-area on the profurcasternum (Figs. 41–42, 64, 75) and the elongated basitarsus, which is considerably longer than the 2nd tarsomere. While few members occur in Borneo, the subfamily is mainly distributed and most diverse throughout the Philippine Islands and also has single representatives in the Palau Islands and Wallacea (Fig. 344). Amongst the subfamilies members, the Tisamenini n. trib. are here regarded as the most basal forms since they possess several plesiomorphic character states such as the predominantly ground-dwelling habits and strict winglessness and due to their small size, stocky appearance and short, robust antennae most closely resemble Dataminae-taxa (→ 3.2.) The medio-apical spine of the area apicalis, an apomorphic character that is frequently present throughout Obriminae, is indistinct or obsolete on the meso- and metatibiae of most Tisamenini-members. Whereas the hairy structures of the eggs are mostly or fully reduced in the tribes Obrimini and Miroceramiini, eggs of Tisamenini n. trib. to a variable degree possess hairy structures, which however lack the conspicuous specializations seen in Dataminae (→ 3.1). The composite spines of the pronotum and raised triangular area of the mesonotum are autapomorphies of Tisamenini n. trib., readily distinguish this tribe from all other Obriminae and place Tisamenini n. trib. as the sister-group of Obrimini + Mirocermiini (Fig. 97b). The single known genus of the tribe Miroceramiini, i. e. Miroceramia Günther, 1934, is the only member of Obriminae that is represented in Wallacea, thus represents the southeasternmost distribution of the entire subfamily. Also, it is here interpreted as the most derived taxon of Obriminae, since it is the only winged member of the subfamily. As shown above, the secondary gain of wings in both sexes of Miroceramiini is an apomorphic character state and an autapomorphy of the tribe, which readily distinguishes it from all other Obriminae (Fig. 97b). The lack of hairy structures in the eggs and presence of composite posterior spines on the mesonotum and metanotum support close relation to Obrimini and are synapomorphies of Obrimini + Miroceramiini (Fig. 97b). The majority of Obriminae-taxa belongs in the tribe Obrimini, which is the by far most diverse subordinate taxon of the subfamily and divides into three fairly well marked generic groups, here defined as the Obrimus-group, Stenobrimus-group and Theramenes-group (→ 3.2.1). The inter-generic relationships however still deserve comprehensive study and clarification and the only character that might represent a possible autapomorphy of Obrimini is the presence of pseudoforamina on the metasternum of certain genera (Figs. 38–40). These so-called pseudoforamina are slit-like invaginations or semi-cingulate to cingulate pits or holes in the lateral portion of the metasternum and are certainly apomorphic developments, whose function is not yet known. The phylogenetic value of these structures however might be limited since they are only present in some genera of the Obrimusgroup (→ 3.2.1), but lack in all other taxa. It is hoped that future phylogenetic or molecular studies will clarify many of these inter-generic relationships within Obrimini and provide more information on the function of these metasternal structures.

4. Revision of Heteropteryginae Kirby, 1896 Type-genus: Heteropteryx Gray, 1835: 32. Heteropteryginae Kirby, 1896: 472 Rehn, 1904: 89. Kirby, 1904a: 396 (in part). Günther, 1953: 546, 551 (in part). Bragg, 1998: 12 (in part). Bragg, 2001: 69 (in part). Zompro, 2004: 195. Otte & Brock, 2005: 17. Seow-Choen, 2016: 405. Heteropterygini Redtenbacher, 1906: 162 (in part). Günther, 1953: 551 (in part). Bradley & Galil, 1977: 199 (in part). Sellick, 1998: 208, fig. 7. Bragg, 1998: 13 (in part). Bragg, 2001: 70 (in part). [Not: Heteropteryginae Bruner, 1915: 37 (=Phylliidae)].

Description (♂, ♀): Medium sized to very large (body length 34.3–166.0 mm), mostly stocky and robust (♀♀ in


HENNEMANN ET AL.

particular), often strongly spinose Heteropterygidae. Sexual dimorphism distinct with ♂♂ much more slender and elongate than ♀♀. ♂♂ roughly cylindrical, ♀♀ with body ± compressed dorsoventrally. Both sexes with wings. All brachyterous taxa with a tympanal area (= stridulatory organ) formed by a number of fused wing-cells in basal portion of alae positioned between the radial and cubital vein; radial vein set with minute teeth dorsally. Radial sector of tegmina branched. ♀♀ brachypterous with tegmina covering alae completely; ♂♂ mostly brachypterous, more rarely with fully developed alae and elongated tegmina, which reach up to ¾ the length of the alae. Alae of brachyterous taxa without a distinct separation between costal region and anal fan. Head hardly longer than wide, vertex ± globose. No ocelli. Antennae long, >½ of body length and much longer than the fore legs; with 22–28 antennomeres but number of antennomeres unequal in ♂♂ and ♀♀ with ♂♂ having two less antennomeres than ♀♀. Antennomeres strongly elongated and much longer than wide; III very elongate and longer than pedicellus and following three antennomeres. No gula. Pronotal foramen at anterior margin on pronotum. No sensory areas on prosternum. Profurcasternum often with a ± distinct central sensory area. Meso- and metasternum simple and ± spinose; no foramina. Head, nota and pleurae of thorax, lateral margins of abdominal tergites II–VII and surface of abdominal sternites spinose; armature most prominent on head and thorax and usually more pronounced in ♂♂. Mesothorax short and at best 2x (♀♀) or 3x (♂♂) longer than pronotum; in ♀♀ ± strongly widening towards posterior. Mesonotum with 2–6 posterior-medial spines on a common base between the bases of the tegmina; no distinctly enlarged posterior mesonotals. Metanotum broader than long. Median segment slightly longer than metanotum, and distinctly separated. Abdomen of ♀♀ broadened medially with segment IV widest; all segments wider than long. Abdominal tergites II–VI of ♀♀ often ± expanded laterally and with dentate/spinose leaf-like lateral appendages. Abdomen of ♂♂ slender, segments II–VII roughly parallel-sided or gently narrowing; occasionally with dentate lateral appendages on II–IX. Abdominal sternites II–IV with ± distinct, impressed excavations laterally. ♀♀ with a prominent praeopercular organ on sternum VII, usually formed by two spine-like projections. Cerci of both sexes small and compressed basally. ♀♀ with a gonangulum and gonoplacs; gonapophyses VIII considerably longer than gonapophyses IX. Epiproct and subgenital plate of ♀♀ elongated and forming a beak-like ovipositor; apex of epiproct often specialized; subgenital plate acutely keeled longitudinally. Anal segment of ♂♂ with paired thorn-pads at posterior margin. Vomer well developed and sclerotized, roughly triangular in shape and with a single, acute apical hook. Poculum large and strongly convex. Femora rectangular in cross-section, all carinae ± spinose or dentate. Profemora straight, at best slightly flattened interobasally. No medioventral carina on femora, ventral surface either smooth or with a longitudinal row of minute spines. Outer ventral carinae of metatibiae armed with prominent triangular or hook-like spines. Area apicalis with a distinct spine medio-apically. Tarsi short, basitarsi hardly longer than 2nd tarsomere. Eggs (Figs. 412–447): Large to very large (capsule length 5.5–10.2 mm), barrel or lemon-shaped; polar-area round to ± conical. Capsule ± circular in cross-section, 3x wider than long. . . . . . . . . . . . . . . . . . . glaber More slender insects (Figs. 121–122); mesothorax only slightly widening towards posterior and with posterior only 1.6x wider


HENNEMANN ET AL.

5. 6. 7. 8. 9. 10. 11. -

12. 13. 14. -

than anterior portion; abdominal tergites III–VI < 3x wider than long and only IV with broad leaf-like lateral expansions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .gintingi n. sp. Abdominal tergites II–VI at best gently rounded laterally; pre-median pronotals and posteromedial mesonotals simple spines (latter may be composite) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Lateral margins of abdominal tergites II–VI greatly deflexed and lobed (Figs. 161–162); pre-median pronotals very long and dichotomic; posteromedial mesonotals strongly composite (Fig. 164); Central Sumatra . . . . . . . . . . . . . . . . . . . kerincia n. sp. Mesothorax with 2–4 posteromedial mesonotals between bases of tegmina.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Mesonotum with six prominent posteromedial mesonotals on a strongly raised common base (Fig. 190); . . . . . . . . . . muelleri Apex of subgenital plate angular and ± notched medially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Apex of subgenital plate obtusely rounded (Fig. 117); Peninsular Malaysia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . erringtoniae Large (body length > 10 cm); pre-median mesonotals not considerably enlarged (Fig. 181); tegmina reaching to abdominal segment III; Singapore & S-Peninsular Malaysia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .macroptera n. sp. Smaller (body length < 9 cm); pre-median mesonotals strongly enlarged (Fig. 209); tegmina reaching to abdominal segment II; Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rosenbergii Epiproct long and projecting over apex of subgenital plate; subgenital plate with apex roundly angular or notched and base rounded.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. grayii species-group Epiproct short and not reaching apex of subgenital plate; subgenital plate with apex ± pointed and base with a distinct angular median projection; only Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. echinata species-group Epiproct with lateral teeth; Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Epiproct without lateral teeth and apex bifid; not Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Large (body length > 10 cm); mesonotum 1.5x longer than pronotum; anterior-mesal mesonotals small and with four posteromedial mesonotals between bases of tegmina (Figs. 257–258) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . grayii Smaller (body length < 10 cm); mesonotum hardly longer than pronotum; anterior-mesal mesonotals prominent and about equal in size to anterior mesonotals, only with two posteromedial mesonotals between bases of tegmina (Figs. 228–229). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . dehaanii Very small (body length < 6 cm); apex of subgenital plate obtusely rounded (Fig. 266); Sumatra . . . . . . . . . . . . . . . . . . . parva Larger (body length > 8 cm); apex of subgenital plate broadened, angular and notched medially (Fig. 240); Vietnam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gorochovi n. sp. Lateral spines of mesonotum unequal in size, antero-lateral mesonotals enlarged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Lateral spines of mesonotum roughly equal in size, antero-lateral mesonotals not enlarged (Fig. 304) . . . . . . . . . . . . saussurei Large (body length > 9 cm); abdominal tergites spinulose; tegmina short and at best reaching posterior margin of abdominal tergum II; lowland form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . echinata Smaller (body length < 8 cm); abdominal tergites rugose dorsally; tegmina long and at least reaching half way along abdominal tergum III; mountainous form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . scabra

* ♀♀ of H. aculeata n. sp. and H. mecheli (Redtenbacher, 1906) are not known

♂♂ 1. 2. 3. 4. 5. 6. 7. 8. -

Ventral surfaces of meso- and metafemora with a longitudinal median row of spines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Ventral surfaces of meso- and metafemora smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2. muelleri species-group Pre-median pronotals and posteromedial mesonotlas simple spines (latter may be composite) . . . . . . . . . . . . . . . . . . . . . . . . . 3 Pre-median pronotals and posteromedial mesonotals very long and dichotomic or antler-like (Fig. 165); Central Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . kerincia n. sp. Metanotum and median segment unarmed; tegmina overlapping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Metanotum and median segment with a posteromedian pair of spines; tegmina and alae small and slender, not overlapping; Simeulue Island . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . jacobsoni Macropterous, alae considerably longer than tegmina and at least reaching half way along abdomen . . . . . . . . . . . . . . . . . . . . 5 Micropterous, alae shorter or at beast equal in length to tegmina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Alae long, at least reaching to abdominal segment VII; two posteromedial mesonotals between bases of tegmina; no lateral lobe on abdominal tergum VIII. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Alae shorter, only reaching to abdominal tergum VI (Fig. 98); four posteromedial mesonotals between bases of tegmina (Figs. 103–104); abdominal tergum VIII with a distinct lateral lobe; West Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . . aculeata n. sp. Anterior coronals simple spines; mesonotum short with disc < 2.5x longer than wide; anterior mesonotals and pre-median mesonotals distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Anterior coronals multi-spinose and crenulate (Fig. 146); mesonotum slender with disc > 4x longer than wide (Fig. 145); anterior mesonotals and pre-median mesonotals obsolete (Figs. 145–146); Sumatra. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .glaber Post-median mesonotals obsolete or lacking; epiproct very small and rounded (Fig. 401); cerci with an apical tooth dorsally (Fig. 178); Singapore & S-Peninsular Malaysia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .macroptera n. sp. With a pair of post-median mesonotals; epiproct elongated and projecting over posterior margin of anal segment (Fig. 185); cerci club-like (Fig. 400); Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .mecheli Anterior coronals simple spines; mesonotum with distinct spines on disc . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Anterior coronals multi-spinose and crenulate (Fig. 126); no distinct spines on disc of mesonotum (Fig. 127); Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .gintingi n. sp.



31

9. 10. 11. 12. 13. 14. 15. 16. -

With 2–4 posteromedial mesonotals between bases of tegmina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Mesonotum with six prominent posteromedial mesonotals on a strongly raised common base (Fig. 188); Sumatra . . . muelleri Large (body length > 7 cm); tegmina plain with anterior margin bright green (Figs. 107–108); epiproct elongated and projecting over posterior margin of anal segment (Fig. 119); Peninsular Malaysia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . erringtoniae Smaller (body length < 7 cm); tegmina with a whitish lateral marking in apical half (Figs. 200–202); epiproct very small and rounded (Fig. 398); Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rosenbergii Abdominal tergites II–IV with a prominent central pair of spines; lateral margins of anal segment simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. grayii species-group No enlarged central pair of spines on abdominal tergites II–IV; lateral margins of anal segment with a tooth sub-basally; only Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. echinata species-group Body length > 5,5 cm; not Sumatra. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Very small (body length < 4 cm); Sumatra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . parva Body length < 7,5 cm, stocky insects ; mesonotum short and just slightly longer than pronotum, with two posteromedian mesonotals between bases of tegmina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Larger (body length > 7,5 cm) and more slender insects; mesonotum 1.5x longer than pronotum with four large posteromedial mesonotals between bases of tegmina (Fig. 251–252); Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . grayii Mesonotum with large anterior-mesonotals and a pair of pre-median mesonotals (Figs. 226–227); lateral spines of abdominal tergites II–VI minute; Borneo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . dehaanii Anterior mesonotals indistinct and pre-median mesonotals lacking (Fig. 237); lateral spines of abdominal tergites II–VI prominent; Vietnam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gorochovi n. sp. Lateral spines of mesonotum unequal in size and antero-lateral mesonotals ± enlarged; epiproct small and rounded, wider than long; abdominal tergites II–VI tapering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Lateral spines of mesonotum equal in size, antero-lateral mesonotals not enlarged (Fig. 299–300); epiproct elongated, slender and longer than wide (Fig. 297); abdominal tergites II–VI almost parallel-sided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . saussurei Large (body length > 6,5 cm); abdominal tergites II–VI spinulose dorsally; antero-lateral mesonotals strongly enlarged (Fig. 282); lowland form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . echinata Small (body length < 6 cm); abdominal tergites II–VI minutely rugulose dorsally; antero-lateral mesonotals just slightly larger than other lateral spines of mesonotum (Fig. 314); mountainous form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . scabra

4.1.1 The muelleri species-group The ten species of the muelleri species-group are the sister-group of the remainder of Haaniella. They are more closely related to Heteropteryx Gray, 1835 than the two other groups, which is supported by the smooth medioventral surface of the meso- and metafemora, long apically bifid epiproct of ♀♀, increasingly elongated antennae and non setose eggs. All these characters are shared with Heteropteryx. Three species have well developed but unequally sized tegmina and alae in the ♂♂, thus resembling Heteropteryx whose ♂♂ have fully developed wings that cover the entire abdomen. In contrast, ♂♂ of all species of the grayii-group and echinatagroup are exclusively brachypterous. The species contained in this group are characterized by the smooth ventral surface of the meso- and metafemora and more or less lemon-shaped eggs, which have the polar-area and operculum ± convex and roundly conical. The eggs have the surface of the capsule and operculum irregularly granulose or tuberculose and destitute of setae. Abdominal tergites II–IV of ♂♂ never bear an enlarged median pair of spines but may be spinose posteriorly in certain taxa. Females may have single abdominal tergites laterally deflexed and dilated into lobes or rounded expansions and ♂♂ of certain species may have abdominal tergites VII–IX with a ± distinct lateral lobe. The lateral invaginations of abdominal sternites II–VI, that are typical and often very prominent in members of the other two species-groups, are very indistinct or lacking in members of the muelleri-group. The epiproct of ♀♀ is always bifid apically, in some species increasingly fused with the anal segment and always projects over the apex of the subgenital plate. The antennae are increasingly elongated when compared to the grayii-group or echinatagroup, those in ♂♂ usually almost equal in length to the body and in ♀♀ at least reaching to abdominal segment V. In the two other species-groups the antennae are considerably shorter than the body in ♂♂ and at beast reach abdominal segment III. There are 24 antennomeres in ♀♀ and 22 in ♂♂. The head has the vertex usually more strongly convex than in the grayii-group and echinata-group and occasionally armed with crenulate lobes. The ventral spines of the metatibiae are never conspicuosly incurving as in members of the grayii-group and echinatagroup but more or less straight or sometimes decidedly broadened, triangular in shape and tooth-like. The coxal trochanters and intersegmental membranes between abdominal sternites I–III are never brightly coloured as in the grayii-group and echinata-group.


HENNEMANN ET AL.

The majority of species occurs throughout Sumatra with one representative on Simeulue Island some 150 km off the west coast of Sumatra, one species in Singapore and two in Peninsular Malaysia. None occurs in Borneo. For a comparison and distinction from the other two species-groups of Haaniella see Table 1.

Haaniella aculeata Hennemann, Conle, Brock & Seow-Choen n. sp. (Figs. 98–104, 361, 366, 403, 489) [Acutely spined Haaniella] HT, ♂: Balun VI.1914; Pad. Bovenl. (Sumatra) E. Jacobson; E. Jacobson, Pad. Bovenl. Sum. Balun VI.1914; Heteropteryx ? dilatata Park. ? K. Günther det. [RMNH].

Diagnosis: ♂♂ of this distinctive new species are characterised by the prominent armature of the legs, which includes a large dagger-like medio-apical dorsal spine on the femora. The elongated wings are shared with H. glaber (Redtenbacher, 1906), H. macroptera n. sp. and H. mecheli (Redtenbacher, 1906), but are comparatively shorter than in these three latter species with the alae merely projecting somewhat over the posterior margin of abdominal segment V. While the colouration resembles H. erringtoniae (Redtenbacher, 1906) from Peninsular Malaysia, whose ♂♂ do not have elongated wings, it differs from the macropterous ♂♂ of the three aforementioned species by the more pronounced body armature, having four instead of two posteromedian mesonotals between the bases of the tegmina and having the lateral margins of abdominal tergites V–VI expanded into a spinose posterior lobe (Fig. 98). Without knowledge of the ♀♀, the exact systematic position of H. aculeata n. sp. is difficult to decide, but the most closely related species appear to be H. erringtoniae (Redtenbacher, 1906), H. macroptera n. sp. and H. muelleri (Haan, 1842). Etymology: The name (aculeatus lat. = acutely spinose) refers to the strongly developed armature of the legs of this peculiar new species. Description: ♂♂ (Figs. 98–99). Medium-sized (body length 70.5 mm) fairly slender for the genus with elongated wings and very prominently and acutely armed extremities. Head and most of thorax dull green, the abdomen mid brown with a greenish hue. Mesonotum with the lateral portions ochre and the lateral margin marked by a fine black line. Lower portion of mesopleurae and most of metapleurae very dark brown to black. Mesosternum dull ochre with the lateral portions blackish brown. Cephalic and thoracic spines green with the tips black, spines of abdomen and legs black. Tegmina dull greenish greenish brown with the anterior portion broadly bright apple green and a few well defined pale green markings interior of the radial vein. Costal region of alae greenish mid brown (fading towards base), the anal fan transparent greyish pink with dark brown veins. All femora dark greenish brown, the tibiae reddish pale to mid brown. Head globose, hardly longer than wide with the vertex roundly convex and armed with ten long and slender spines (Fig. 103). Vertex with four prominent and long coronals which roughly form the four corners of a square; the anterior pair somewhat larger than the posterior pair. Anteriorly with a further pair of slightly smaller but very pronounced, slender supra-occipitals (Fig. 104). Posterior margin with two distinct lateral coronals and behind the eyes with one fairly long supra-orbital spine. Antennae at least reaching to posterior margin of abdominal segment II (broken in the holotype). Pronotum slightly longer than wide with lateral margins distinctly concave and posterior margin wider than anterior margin; antero-lateral pronotals and postero-lateral pronotals distinct and just before transverse median sulcus with a strong pair of pre-median pronotals as well as two somewhat shorter interposterior pronotals near posterior margin (Fig. 103). Mesothorax slender, elongate, some 2.5x longer than pronotum with posterior portion distinctly widened and the mid portion very gently constricted in dorsal view (Fig. 104). Mesonotum gently narrowed medially and about 4x longer than wide and disc armed with six prominent spines; a pair of moderately long anterior mesonotals, a very long and slender pair of pre-median mesonotals and a somewhat shorter pair of post-median mesonotals (Fig. 103). Between the bases of the tegmina with four very long and slender posteromedials on a common base; the anterior two directed anteriad, the posterior two directed posteriad (Fig. 103). One or two short spines are present on lateral margins. Tegminal well pronounced but shorter than all other mesothoracic spines. Mesopleurae with a long and slender supra-coxal and five long and slender, unequally sized laterals along lower margin; the most posterior spine much shorter than all previous. Metapleurae strongly dilated in the median portion and with five long and slender spines; a further spine close to metacoxae and the supra-coxal very prominent and slender. Sensory area of profurcasternum moderately developed and profurcasternum armed with a pair of short, black spines. Mesosternum smooth except for three short but robust, REVISION OF HETEROPTERYGINAE


33

black lateral spines, which increase in size towards the posterior; also a pair of minute posterior mesosternals between coxae. Metasternum with four stout black spines on each side of the median line, the outermost longest. Tegmina elongate-ovate, slender, roughly 2/3 the length of alae and reaching about half way along abdominal tergum III (Fig. 98); radial vein somewhat raised in basal portion of tegmen. Alae slightly projecting over posterior margin of abdominal tergum V; near base with a well developed tympanal area. Abdominal segments gently narrowing towards posterior, III–VII slightly narrowing from III towards VII and averaging 1.3x longer than wide. Lateral margins of III somewhat deflexed posteriorly, in IV–VI much expanded and forming a strongly projecting, spinose lobe posterolaterally; the lobe with three long and very slender spines and a further but much smaller spine shortly behind mid of segment. VII with the posterolateral angles semicircularly deflexed and this portion with three very weakly developed teeth only (Fig. 101). VI–IX with a small pair of posteromedian spines close to median line (most pronounced on VII and VIII). Sternites II–VI with eight black spines; one pair pre-medially, one pair posteriorly and two spines near each lateral margin. Sternum VII only with two short, black spines near the posterior margin. Anal segment somewhat constricted medially with the posterolateral portions widened and broadly rounded; the posterior margin indented medially (Fig. 101). Epiproct elongate, much longer than wide, slightly narrowed towards the posterior with the apex gently notched; decidedly projecting over posterior margin of anal segment (Fig. 403). Cerci strongly compresseed basally with the apical portion club-like. Vomer triangular with the outer margins strongly swollen and the median portion distinctly impressed, the apical hook almost straight; dull ochre (Fig. 403). Poculum with the posterior margin broadly rounded. Legs long and moderately slender with all carinae (including dorsal carinae of tibiae) very prominently armed with large and acute, triangular black teeth; metafemora almost reaching to posterior margin of abdominal segment VI and metatibiae projecting considerably beyond apex of abdomen. Femora dorsally with a very characteristic and prominent, backward directed, dagger-shaped medio-apical spine (Fig. 99). Metatibiae armed with six very prominent and broadly triangular, black teeth and a few intercalated teeth on two outer ventral carinae and four distinct, black, triangular teeth on each dorsal carina (Fig. 366). Ventral surface of all femora smooth (Fig. 361). Comments: Since this new species is only known from the single ♂ holotype, nothing is known about its variability or host-plants. Females and eggs are unknown. Distribution (Fig. 489): So far only known from the type-locality Balun, a small village in the South Solok Regency of the province Sumatera Barat in western Sumatra. The abbreviation “Pad. Bovenl.” on the original labels by the collector E. Jacobson means “Padangsche Bovenlanden” and refers to the Padang Highlands. Specimens examined: 1 TABLE 2. Measurements [mm] of Haaniella aculeata n. sp. ♂ HT [RMNH] Body

70.5

Head

5.3

Pronotum

5.2

Mesonotum

13.2

Metanotum

4.0

Median Segment

4.8

Tegmina

19.4

Alae

30.0

Profemora

18.8

Protibiae

22.0

Mesofemora

14.7

Mesotibiae

18.9

Metafemora

22.3

Metatibia

26.3

Antennae

> 38.0


HENNEMANN ET AL.

Haaniella erringtoniae (Redtenbacher, 1906) rev. stat. (Figs. 94, 105–120, 360, 369–370, 397, 412–413, 428, 450–452, 490) [Errington’s Haaniella] Heteropteryx erringtoniae Redtenbacher, 1906: 169. HT, ♀: Muséum Paris, Pr. Malacca, Selangor, Errington, et Chape 1900, Heteropteryx Erringtonae Redtenb. type!; Redtenbacher Det. 1901 [MNHN—in alcohol]. Haaniella erringtoniae, Günther, 1932: 313. Dräger, 2011: 46, figs. 10: 4a–c (♀ abdomen), 11: 1 (♂ thorax), 12 (♀), 13 (♂), 14–15 (nymphs), 37:2a–b (egg). Haaniella mülleri erringtoniae, Günther, 1944: 72. Haaniella muelleri, Brock, 1995: 95. Brock, 1999: 141 & 183, figs. 95 a–c (♂), b (alae), c (abdomen), d–e (♀), e abdomen), f (egg), pl. 54 (♀) & 55 (♂). Seow-Choen, 2000a: 151, figs. 7 (♂), 8 (♀). Seow-Choen, 2000b: 40, pl. 103 (♂), 104 a–b (♀), c (egg). Zompro, 2004: 197. Otte & Brock, 2005: 149 (in part). Haaniella mülleri, Schulten, 1995: 98.

Material examined (58 ♂♂, 57 ♀♀, 5 nymphs, eggs): PENINSULAR MALAYSIA: 1 ♀: Selangor or FMS, BKT or Langrove, FR, Runba, 200ft, 5.1921, H.C. Abraham [RMNH]; 1 ♀ (nymph): Selangor, FMS, at light, 3.9.1934, H. Klante det „mecheli Redtb.“ [RMNH]; 1 ♀ (nymph): Selangor, FMS, Kuala Lumpur, Gombak Valley, 19. Oct. 1921, H.M. Pendlebury, det. Klante “mecheli Redtb.” [RMNH]; 1 ♂ (nymph): Malaysia, Selangor, Bukit Kutu, 3500ft, 7.9.1929, H.M. Pendlebury, det. Klante “mecheli Redtb.” [RMNH]; 1 ♂, 1 ♀: bred by Potvin, 1994 [RBINS]; 1 nymph (n1): Malaysia, Kuala Lumpur, 13.2.1987 / Leg. Ruhberg [ZMUH]; 1 ♀: Malay Peninsula, Genting Tea Estate, 8th June 1992, Coll. C.L. Chan, S.W. Lee & Tan Man Wah [NHMUK]; 1 ♂: Malay Peninsular, Cameron Highlands (?), 1979, Coll. Natives [NHMUK]; 1 ♂: 1500–1600 m, 22.VI.196 Seow-Choen & Dolly, Genting Highlands [coll. FSC]; 1 ♂, 1 ♀: 1500–1600 m, VIII.2009, Seow-Choen, Genting Highlands [coll. FSC]; 1 ♂: 1500–1600 m, VIII.2012, Seow-Choen, Genting Highlands [coll. FSC]; 2 ♀♀: 1500– 1600 m, VI.2013, Seow-Choen, Genting Highlands [coll. FSC]; 1 ♀: 1500–1600 m, VIII.1994, Seow-Choen, Genting Highlands [coll. FSC]; 1 ♀: ex Zucht U. Ziegler (Berlin), XI.1992, urspr.: W-Malaysia, Selangor, Templer Park [coll. FH, No. 0385-1]; 3 ♂♂, 2 ♀♀: ex Zucht F. Hennemann, 1995, urspr.: W-Malaysia, Selangor, Templer Park [coll. FH, No.’s 0385-2 to 6]; 4 ♂♂, 3 ♀♀, eggs: ex Zucht F. Hennemann, 1996–1998, urspr.: W-Malaysia, Selangor, Templer Park [coll. FH, No’s 0385-7 to 13, E & ED]; 3 ♂♂, 4 ♀♀: ex Zucht H. Dräger 2011–2013, Herkunft: W-Malaysia, Selangor, Templer Park [coll. FH, No’s 0385-14 to 20]; 1 ♂: West-Malaysia, Pahang, Genting Highlands, 1500–1600 m, leg. Seow-Choen & Dolly 22.VI.1996 [coll. FH, No. 0385-21]; 1 ♀: WestMalaysia, Pahang, Genting Highlands, 1500–1600 m, leg. F. Seow-Choen VIII.2009 [coll. FH, No. 0385-22]; 1 ♀: ex Zucht O. Conle, 1997, Herkunft: West-Malaysia, Selangor, Templer Park (PSG No. 112) [coll. OC]; 15 ♂♂, 13 ♀♀, 1 ♂ (nmyph): ex Zucht O. Conle, 2003, Herkunft: West-Malaysia, Selangor, Templer Park (PSG 112) [coll. OC]; 26 ♂♂, 25 ♀♀: ex Zucht O. Conle, 2006, Herkunft: West-Malaysia, Selangor, Templer Park (PSG 112) [coll. OC]. Diagnosis: Very close and perhaps the adelphotaxon of H. macroptera n. sp. from Singapore and southern Peninsular Malaysia. Females however differ by the averaging shorter tegmina, smaller lateral spines of abdominal tergites II–VI, more rounded outer angles of the posterior margin of the anal segment (Figs. 115–116), obtusely rounded apex of the subgenital plate (Fig. 117) and shorter and less curved ventral spines of the metatibiae (Fig. 369). Males readily differ by being micropterous and more slender, with the mesothorax relatively more elongate, elongated and slender epiproct (Fig. 119) and sinple cerci, which lack the dorso-apical tooth seen in H. macroptera n. sp.. The eggs are smaller, more globose and have the central elevation of the operculum acutely conical (flattened apically in H. macroptera n. sp.). Etymology: Named in honour of Mdm. Errington, who discovered this species and collected the female holotype (Redtenbacher, 1906: 169). Description: ♀♀ (Figs. 105–106). Fairly large (body length 89.5–114.1 mm) and slender with long and slender legs. Colour variable, mostly pale to dark brown with pretty pale drab or straw mottling and speckles. More rarely dull greenish specimens occur or ones which have a dark brown median long stripe along the dorsal body surface. Often with a bold pale V-shaped marking on mesonotum. Ventral surface of body pale reddish brown and



35

sternites with a whitish patch posterolaterally. Tegmina ranging from plain pale to dark brown and sometimes with pale mottling; anterior margin occasionally green or whitish and often with a distinct white spot near the base. Femora as well as pro- and mesotibia with a distinct pale transverse band just after middle, metatibia with two to three more or less distinct pale bands. Body armature usually pale red with black points but may also be green. Head slightly longer than wide, globose with vertex roundly convex and armed with ten spines (Fig. 110). Vertex with four distinct coronals which together roughly form the four corners of a square; the anterior pair larger than the posterior pair. Anteriorly with a pair of much smaller supra-occipitals (Fig. 109). Posterior margin with two fairly distinct lateral coronals and behind the eyes with one fairly long supra-orbital. Antennae ± reaching posterior margin of abdominal tergum IV; with 24 antennomeres. Pronotum slightly longer than wide with lateral margins distinctly concave and posterior margin wider than anterior margin; antero-lateral pronotals and posterolateral pronotals distinct and just before transverse median sulcus with a strong pair of pre-median pronotals as well as two small but more distant inter-posterior pronotals near posterior margin (Fig. 109). Mesothorax gently and gradually widening towards the posterior with posterior portion some 1.8x wider than anterior margin. Mesonotum roughly parallel-sided, almost twice as long as wide and about 2x the length of pronotum. Lateral margins with 3– 4 fairly small and slender spines, disc with a pair of ± distinct anterior-mesonotals, a strong pair of pre-median mesonotals close to median line and a further pair of smaller and more distant post-median mesonotals (Fig. 110). Between bases of tegmina with two (sometimes three or rarely four) posteromedial mesonotals on a common base; if present the anterior ones are decidedly smaller than the posterior ones (Fig. 109). Tegminal fairly distinct and slender. Mesopleurae with a long and slender supra-coxal spine and four almost equally sized laterals at lower margin. Metapleurae with lateral margin rounded and armed with six long and slender laterals, the most posterior one decidedly smaller than previous; supra-coxal distinct. Sensory area of profurcasternum indistinct. Mesosternum with a lateral row of 3–4 small spines (Fig. 111). Metasternum with 8–10 small spines. Tegmina elongate-oval and ± reaching half way along abdominal tergum III. Alae translucent grey with black veins. All abdominal tergites smooth dorsally. Lateral margins of II–VI moderately deflexed, rounded and with three (rarely four) fairly small spines; very indistinct on VII. Sternites II–VI with four spines on each side of the median line; these most prominent on sternum II and decreasing in size towards VI. Praeopercular organ on sternum VII formed by two fairly distant, small spines and a small convex median swelling inbetween (Fig. 117). Posterior margin of anal segment gently notched. Epiproct tectiform longitudinally, slightly downcurving with the apex deeply notched and forming two acute points (Figs. 115–117). Subgenital plate with dorsal margins roundly excavated medially (Fig. 115) and the apex obtusely rounded; not reaching apex of epiproct (Fig. 117). All legs long and slender, metafemora reaching to abdominal segment VII and metatibiae ± projecting beyond apex of abdomen. All carinae sparsely dentate and metatibiae with medium-sized, fairly slender triangular teeth on the two outer ventral carinae (Fig. 369). Ventral surface of all femora smooth (Fig. 360). ♂♂ (Figs. 107–108). Medium sized (body length 72.0–77.8 mm) and slender insects with long and slender legs. General colour dull green to greyish brown with a ± distinct pale ochre stripe along lateral margins of mesonotum. Ventral surface of body dull ochre, the mesosternum pale ochre to straw. Tegmina plain mid to dark brown or very dull green with the anterior margin bright pale green. Femora mid to dark brown (often with a slight greenish hue), the tibiae dull ochre to dark reddish brown. Armature of legs mostly black. Shape and armature of head as in ♀♀, but armature more prominent with all spines comparatively longer and more slender (Fig. 112). Antennae very long and almost reaching to apex of abdomen; with 22 antennomeres. Pronotum almost twice as long as wide, about equal in length to head, with the lateral margins gently concave; spines as in ♀♀ but decidedly longer. Mesothorax slender, elongate, some 2.3x longer than pronotum with anterior and posterior portions roughly equal in width and the mid portion very gently constricted in dorsal view (Fig. 113). Mesonotum gently narrowed medially and about 4.2x longer than wide; disc with six spines arranged as in ♀♀, but all much longer and more elongate (Fig. 112). Also the posteromedian mesonotals between the bases of tegmina distinctly longer than in ♀♀ and often with an additional much smaller lateral spine at the base (Fig. 112). Mesopleurae with a long and slender supra-coxal spine and four fine spines along lower margin; occasionally a few smaller spines near the lower margin. Armature of metapleurae as in ♀♀. Sensory area of profurcasternum weakly developed. Mesosternum smooth except for 3–4 small, black lateral spines. Abdominal sternum I with a pair of minute pair of spines between coxae (Fig. 114). Metasternum as in ♀♀. Tegmina ovate with the apex fairly acute and reaching ± half way along abdominal tergum II. Abdominal segments II–VII averaging 1.5x longer than wide, IV–VII gradually decreasing in length and II widening towards anterior. Tergites II–VII smooth dorsally and with a ± decided fine longitudinal median carina; IV–VII with the lateral margins moderately rounded and deflexed in


HENNEMANN ET AL.

posterior half. II–VII with 3–4 fairly long but very slender lateral spines, which are restricted to the deflexed posterior portion in IV–VII (Fig. 107). Sternites II–VI with four slender spines, VII unarmed or with a small pair of posterior spines. Anal segment somewhat narrowed towards posterior with posterior margin triangularly notched medially (Fig. 119). Epiproct elongated, longer than wide, notched apically and projecting over apex of abdomen (Fig. 119). Cerci small, club-like and compressed dorsoventrally. Vomer triangular and much longer than wide with the apical hook somewhat curved dextrally; pale ochre (Fig. 397). Poculum with posterior margin broadly rounded and notched medially (Fig. 397). Legs all very long and slender, mesofemora almost as long as pro- and mesonotum combined, metafemora reaching to abdominal segment VI and metatibiae projecting distinctly beyond apex of abdomen. All carinae more or less distinctly dentate, the teeth fairly slender and pointed; two outer ventral carinae of metatibia with 5–7 slender, triangular and backward directed black teeth (Fig. 370). Ventral surface of all femora smooth. Nymphs: These have the body and leg armature much more obvious than the adult insects and also the lateral margins of the abdominal tergites or more dilated and deflexed. Often prettily mottled. Variability: This species is fairly constant in the armature of the body and legs, with variability mainly restricted to the relative size of the cephalad and thoracic spines and teeth of the extremities. Females usually have two posteromedial mesonotals between the bases of the tegmina but rarely there may be a further smaller pair of spines anterior to the main pair. While the colouration is quite constant in ♂♂, ♀♀ show a wide range of different colour patterns (Figs. 450–452), specimens ranging from almost plain pale to mid brown to almost dull green. There are several intermediate steps with specimens having only certain parts of the body with a greenish hue, furnished with green speckles or having all of the thoracal spines green. Some specimens are prettily mottled with dark and pale brown or black with green spines, others may have a dull brown longitudinal median stripe along the entire dorsal surface of the body (less defined on abdomen; Fig. 451). Often there is a very large but weakly defined V-shaped marking on the mesonotum, the femora have pale brown to ochre median tranverse band and the tibiae are to a variable degree mottled with pale ochre. Abdominal sternites II–VII usually bear a pale ochre to white posterolateral spot, which is lacking in plain brown specimens. The tegmina range from plain mid to dark brown or may be speckled with darker/paler brown or green and often possess a small semi-circular white subbasal marking at the anterior margin. The anterior margin is often greenish or more rarely whitish. The only colourvariation seen in ♂♂ concerns to the width of the bright green stripe along the anterior margin of the tegmina. Several hundred eggs from cultured material have been examined and show the anterior expansions of the micropylar plate are somewhat variable in shape. They are either more or less club-like and widened at the apex or slender and almost straight to distinctly incurved. Eggs (Figs. 412–413, 428): Large, almost circular in cross-section and distinctly lemon-shaped with capsule strongly narrowed in anterior portion and polar-area roundly conical; about 1.5x longer than wide. Surface of capsule very minutely granulose, slightly glossy and irregularly set with minute but acutely pointed, slender black tubercles. Micropylar plate principally X-shaped, with all four extension fairly slender; anterior extensions incurving in apical half, posterior extensions almost straight and somewhat broadened and rounded apically. Interior portion of micropylar plate gently raised, the outer margin also raised and entire plate surrounded by a blunt bulge; surface as in capsule. Micropylar cup a small black knop, at both sides accompanied by a short carina. Operculum almost circular, marginated and conically raised medially. General colouration dull greyish brown, the anterior margin of the capsule, outer margin and bulge surrounding the plate very dark grey to black. Measurements [mm]: length incl. operculum 8.7–9.1, length 7.8–8.0, width 5.1–5.2, height 5.6–5.8. Comments: The identity and status of H. erringtoniae (Redtenbacher, 1906) had been in dispute, mostly because the ♀ holotype from “Malacca” (= Peninsular Malaysia) had not been traced in MNHN until recently. The specimen was finally located in the alcohol collection by Emmanuel Delfosse (MNHN) in 2012, who kindly provided photos of the specimen, thus clarifying the taxonomic status of this species. Recent authors (e.g. Brock, 1995; Brock, 1999; Otte & Brock, 2005) have treated H. erringtoniae as a synonym of the Sumatran H. muelleri (Haan, 1842), but examination and comparison of the holotype of H. erringtoniae and other Malayan specimens with Haan’s type specimens of H. muelleri in RMNH have clearly shown these to represent two distinct species. Consequently, H. erringtoniae is here re-established as a valid species (rev. stat.) and appears to be restricted to the Malay Peninsula, while H. muelleri is endemic in Sumatra. Günther (1944: 72) treated H. erringtoniae as a subspecies of H. muelleri, which was not accepted by Brock (1995: 95; 1999: 183) who listed it as a synonym of the latter species and stated that there was no justification for the subspecies status suggested by Günther (1944).



37

H. erringtoniae has been cultured in Europe since the late 1980’s from stock originating from Peninsular Malaysia (Selangor: Templer Park) and was included on the Phasmid Study Group (PSG) culture-list as culture No. 112 “Haaniella muelleri”. Hence, all references to the captive rearing, life cycle or biology of “H.muelleri” in fact relate to H. erringtoniae. This species is fairly easy to culture in humid but somewhat ventilated conditions, using e.g. bramble (Rubus fruticosus, Rosaceae), oak (Quercus spp., Fagaceae) and guava (Psidium guayava, Myrtaceae) as alternative foodplants. Eggs take approximately 10–12 months to hatch, nymphs being slow growing and adults quite long-lived. Although not very common in Peninsular Malaysia, H. erringtoniae is very common locally along a service road leading up to the casino in the Genting Highlands (Pahang). Seow-Choen (2000b: 40) provided a list of native foodplants in Peninsular Malaysia (→ 7.6). Distribution (Fig. 490): Peninsular Malaysia: Selangor [MNHN—type locality]; Selangor, Runba 200 ft. [RMNH]; Selangor [RMNH]; Selangor, Batu Caves [Brock, 1999: 144]; Selangor, Kuala Lumpur, Gombak Valley [RMNH]; Selangor, Kuala Lumpur [ZMUH]; Selangor, Templer Park [coll. FH, coll. OC]; Selangor, Ulu Langat [Brock, 1999: 144]; Selangor, Bukit Kutu 3500 ft. [RMNH]; Pahang, Genting Highlands 1500–1600 m [coll. FH, coll. FSC]; Pahang, Genting Tea Estate [NHMUK]; Negeri Sembilan, Pasoh [Brock, 1999: 144]. Specimens examined: 120 TABLE 3. Measurements [mm] of Haaniella erringtoniae (Redtenbacher, 1906) ♀ HT [MNHN]

♂ [Genting Highlands, coll. FH]

♀ [Genting Highlands, coll. FH]

♂♂ (coll. FH)*

♀♀ (coll. FH)*

Body

114.1

76.0

99.0

72.0–77.8

89.5–110.0

Head

11.9

6.0

8.9

5.9–6.5

8.5–9.7

Pronotum

9.1

6.2

8.2

5.8–6.4

7.2–8.6

Mesonotum

18.5

14.6

17.3

12.0–15.0

15.1–18.3

Metanotum + Median Segment

12.0

9.0

11.5

8.2–9.3

11.8–14.3

Tegmina

24.8

12.0

23.0

10.8–14.4

21.5–25.5

Profemora

22.8

20.6

22.2

19.5–20.5

22.4–25.4

Protibiae

26.4

23.2

22.5

20.0–25.2

24.7–28.6

Mesofemora

21.0

15.8

18.0

20.0–22.5

18.0–20.2

Mesotibiae

22.9

14.0

21.0

17.6–18.2

20.0–24.5

Metafemora

30.5

24.0

28.0

24.5–26.0

28.5–32.0

Metatibia

28.4

27.8

34.5

26.0–29.0

32.0–39.3

Antennae

-

> 49.0

65.0

66.0–68.0

62.0–67.0

* captive reared

Haaniella gintingi Hennemann, Conle, Brock & Seow-Choen n. sp. (Figs. 121–137, 359, 373–374, 399, 416–417, 458–459, 490) [Ginting’s Haaniella] Haaniella sp. “Sibayak”, Dräger & Kneubühler, 2013: 52, figs (♂, ♀, nymphs, egg). HT, ♀: Museum Leiden, Indon. Sumatra, Ketambe, Atjeh, Tenggara 350m, 1972/1974 H.D. Rijksen, Haaniella muelleri simplex det. P.E. Bragg, Feb. 1993 [RMNH]. PT, 1 ♂, 3 ♀♀: 27.II.2014, Roadside leading to Danau Toba, Prov. Sumtra Utara, approx. 1400 m [coll. FSC]. PT, 1 ♀: IV.2013, edge of Sibolangit Botanic Gardens, 1220 m, Brestagi, Prov. Sumatra Utara [coll. FSC].


HENNEMANN ET AL.

PT, 4 ♀♀: ex Zucht H. Dräger, 2014, Herkunft: NW-Sumatra, Prov. Sumatera Utara, Mt. Sibayak 1400–1600 m [coll. FH, No’s 0753-1 to 4]. PT, 3 ♂♂, 3 ♀♀, 12 eggs: ex Zucht H. Dräger, 2014, Herkunft: NW-Sumatra, Prov. Sumatera Utara, Mt. Sibayak 1400–1600 m [coll. FH, No’s 0753-5 to 10, E]. PT, 1 ♂♂, 1 ♀: ex Zucht F. Hennemann, 2016, Herkunft: NW-Sumatra, prov. Sumatera Utara, Mt. Sibayak 1400-1600 m, [coll. FH, No's 0753-11 & 12]. PT, 3 ♂♂, 1 ♀, : NW-Sumatra, Prov. Sumatera Utara, Berastagi, 1500 m, 2011, leg, Jimmy Gideon Ginting [coll. OC]. PT, 2 ♂♂, 2 ♀♀: NW-Sumatra, Prov. Sumatera Utara, Mt. Sibayak, 1550 m, 2011, leg, Jimmy Gideon Ginting [coll. OC]. PT, 3 ♂♂, 1 ♀: NW-Sumatra, Prov. Sumatera Utara, Jaranguda, 1450 m, 2011, leg, Jimmy Gideon Ginting [coll. OC]. PT, 1 ♂: Ex Zucht B. Kneubühler 2012 (F1), Herkunft: NW-Sumatra, Prov. Sumatera Utara, Berastagi, leg, Jimmy Gideon Ginting [coll. OC]. PT, 1 ♀: Ex Zucht B. Kneubühler 2013 (F2), Herkunft: NW-Sumatra, Prov. Sumatera Utara, Berastagi & Mt. Sibayak, leg, Jimmy Gideon Ginting 2011 [coll. OC]. PT, 11 ♂♂, 9 ♀♀: Ex Zucht O. Conle 2012–2014 (F1), Herkunft: NW-Sumatra, Prov. Sumatera Utara, Berastagi & Mt. Sibayak, leg, Jimmy Gideon Ginting 2011 [coll. OC]. PT, 1 ♂, 3 ♀♀: Indonesia, North Sumatra, Mount Sibayak, Ex breeding R. Krijns, 2014 [RBINS].

Diagnosis: This very distinctive new species differs from all other members of the muelleri species-group by numerous characters. The most closely related species and possible adelphotaxon seems to be H. glaber (Redtenbacher, 1906) n. comb., with which it shares the obsolete mesothoracic armature, large, crenulate and multi-dentate anterior coronals, presence of an enlarged triangular sub-apical tooth on the posterodorsal carina of the femora and broadened, black triangular teeth on the two ventral carinae of the metatibiae of both sexes. Females however differ by the much more slender body and relatively more elongate body segments, less trapezoidal mesothorax, averaging smaller tegmina, strongly laterally deflexed abdominal tergum IV (Figs. 121– 122), different structure of the praeopercular organ (Fig. 134) and obtusely rounded apex of the subgenital plate (Fig. 134). Males readily differ from those of H. glaber by being micropterous (Figs. 123, 125), more slender and by completely lacking the anterior and pre-median mesonotals (Figs. 129–130). The eggs are characteristic for the fairly elongate capsule and very short anterior and posterior extensions of the micropylar plate (Figs. 416–417). Etymology: Dedicated to Jimmy Gideon Ginting (Berastagi, North Sumatra), who first collected this species on Mount Sibayak near Berastagi and provided a culture-stock, that is now being successfully reared in Europe. Description: ♀♀ (Figs. 121–122, 124). Medium sized (body length 90.0–103.0 mm), moderately slender species with much reduced body spination but large crenulate anterior coronals and a strongly widened abdominal tergum IV. Colour variable (→ see comments on variability below) and ranging from ochre to dark brown, sometimes with a greenish hue, with defined dark markings or a longitudinal median stripe on body. Usually with a fine dull green stripe along lateral margins of mesonotum. Body armature pale to dull orange or red, the larger spines with black tips. Anterior margin of tegmina bright apple green. Head a little longer than wide, globose with vertex conically elevated and prominently armed (Fig. 126). Anterior coronals strongly enlarged and forming composite or crest-like swellings with the apex laterally compressed and usually tri-dentate, posterior coronals much smaller and represented by simple but strong spines (Figs. 126–127). Anteriorly with a pair of very small supra-occipitals (sometimes obsolete). Posterior margin with two fairly distinct but stout lateral coronals, which are slightly smaller than the posterior coronals. Behind the eyes with a supra-orbital spine about equal in size to the lateral coronals. Antennae ± reaching half way along abdominal tergum IV; with 24 antennomeres. Pronotum longer than head and a little longer than wide, widening towards posterior and with lateral margins distinctly concave. Antero-lateral pronotals very small, the posterior pronotals somewhat more pronounced and the pair of inter-posterior pronotals very indistinct and considerably smaller than the outer posterior pronotals. Just before transverse median sulcus with a pair of very low and blunt pre-median pronotals (Fig. 126). Mesothorax gently and gradually widening towards the posterior with posterior portion about 1.6x wider than anterior margin. Mesonotum very slightly narrowing towards the posterior, almost 2x longer than wide and roughly 1.8x the length of pronotum. Lateral margins with 5–7 blunt spiniform tubercles, disc with 8–10 minute blunt, paired spinules or spiniform tubercles (Fig. 127). Between bases of tegmina with two fairly prominent, stout posteromedian mesonotals on a somewhat raised common base (Fig. 126). Tegminal spine very small. Mesopleurae with a fairly slender and moderately distinct supra-coxal spine and 5–7 unequally sized, stout laterals along lower margin (Fig. 128). Metapleurae with lateral margin strongly rounded and armed with 5–6 fairly prominent, slender spines, the posterior three spines in the deflexed portion of pleuron decidedly longer than the anterior ones; supra-coxal small. Sensory area of profurcasternum distinct, strongly convex and triangular in REVISION OF HETEROPTERYGINAE


39

outline. Meso- and metasternum with a variable number of minute spines or spiniform tubercles. Tegmina oval in shape and ± reaching 2/3 along abdominal tergum II. Alae translucent dark grey with black veins. All abdominal tergites smooth dorsally but III–V sometimes with a pair of minute posteromedian tubercles. Abdominal tergites II–III widening, IV with the lateral margins strongly and almost semicircularly deflexed (Figs. 121–122), V and VII narrowing and with lateral margins just slightly deflexed and gently rounded; tergum IV by far the widest segment and almost 2x the width of VII. Lateral margins of II–VI armed with 4–5 small, slender spines; these very indistinct to almost obsolete on VI. Sternites II–VI with six or eight small and pair-wise arranged spines; these most prominent on II and decreasing in size towards VI. Praeopercular organ on sternum VII formed by two short and blunt spiniform processes and a small, black, rounded median swelling just in front of these (Fig. 134). Posterior margin of anal segment with a roundly triangular indention and the outer portions roughly semicircular. Epiproct tectiform longitudinally, gradually narrowing towards a slightly downcurving and triangularly incised, acutely bifid apex (Figs. 132–134). Apex of subgenital plate bluntly rounded and hardly reaching half way along epiproct (Fig. 134). All legs moderately long and slender, mesofemora somewhat longer than mesothorax, metafemora slightly projecting over posterior margin of abdominal segment V and metatibiae just not reaching apex of abdomen. All carinae acutely but irregularly dentate; the femora with an enlarged and prominent triangular pre-apical tooth on posterodorsal carina. Two outer ventral carinae of metatibiae armed with 6–8 prominent and strong, triangular and slightly backward directed black teeth, some of these in the basal portion melted with each other (Fig. 373). Ventral surface of all femora smooth (Fig. 359). ♂♂ (Figs. 123, 125). Fairly small (body length 61.0–65.5 mm) and slender for the genus with reduced body spination but prominent, crenulate anterior coronals and moderately long, slender legs. Colour dull ochraceous brown with a slight greenish hue, the mesonotum and mesosternum contrasting ochre. Mesonotum with a dull green to bluish green line along lateral margins and mesopleure dull green to bluish green. Body armature dark green, the larger spines with black tips. Tegmina plain dark brown with the anterior margin bright apple green. Femora dull ochracheous brown with slight greenish hue, the tibiae dull ochre. Armature of legs mostly black. Shape and armature of head as in ♀♀, but armature more prominent with all spines much longer and more slender (Fig. 129). Antennae reaching to abdominal segment VIII; with 22 antennomeres. Pronotum about 1.3x longer than wide, somewhat longer than head with the lateral margins gently concave; spines as in ♀♀ but the premedian pronotals much larger and decidedly longer than all other spines (Fig. 129). Mesothorax very slender and fairly elongate, some 2.5x longer than pronotum with posterior portion widened and the mid portion gently constricted in dorsal view (Fig. 130). Mesonotum almost parallel-sided and averaging 4.4x longer than wide; disc only with a few paired tubercles, but the pair of posteromedial mesonotals between the bases of the tegmina decidedly longer and more slender than in ♀♀. Tegminal very indistinct. Mesopleurae with a moderately long and slender supra-coxal spine and 4–5 unequally sized laterals along lower margin, the anterior ones very small and rather resembling spiniform tubercles, the posterior 2–3 spines larger and well defined. Armature of metapleurae as in ♀♀. Sensory area of profurcasternum distinct and generally as in ♀♀. Mesosternum smooth except for about eight paired spines in posterior half (Fig. 131), the four posterior ones being more pronounced than the anterior ones; a pair of minute posterior mesosternals between coxae. Metasternum with five low and stout spines on each side of the median line. Tegmina fairly small, ovate in outline and not reaching posterior margin of median segment. Abdominal segments II–V very slightly increasing in length, VI and VII decreasing. II–VI rectangular and of uniform width, VI some 1.4x longer than wide. Tergum VII with lateral margins roundly deflexed and somewhat lamellate (Fig. 136). Tergites II–VI entirely smooth and lateral margins only with a single spine near posterior angle; sometimes a single very small spine in median portion. VII with a pair of pre-median tubercles and a transverse row of four blunt tubercles near posterior margin. Sternites II–V with eight paired spines, two near lateral margins, one pair pre-medially and one pair near posterior margin. VI lacking the pre-medial pair of spines, VII only with a few small paired tubercles; both slightly tectiform longitudinally. Anal segment gently narrowing towards posterior, the posterior margin widely triangularly excavated (Fig. 136). Epiproct very short. Cerci small, club-like and compressed laterally in basal portion. Vomer slender with the apical portion strongly narrowed and gradually tapered towards a fairly long, straight apical hook; mid brown with the apex black (Fig. 399). Poculum with posterior margin broad and angular, the basal portion with a prominent blunt median projection (Fig. 135). Legs all fairly long and slender, mesofemora reaching half way along pronotum, metafemora ± reaching half way along abdominal segment VI and metatibiae projecting somewhat beyond apex of abdomen. All carinae of femora acutely armed with pointed and fairly slender, spine-like teeth and usually with an enlarged pre-apical tooth on


HENNEMANN ET AL.

posterodorsal carina. Ventral carinae of pro- and mesotibiae with about four fairly small teeth. Dorsal carinae of all tibiae entirely smooth. Two outer ventral carinae of metatibiae armed with 7–8 strong, triangular and slightly backward directed black teeth (Fig. 374). Ventral surface of all femora smooth. Variability: Both sexes show variability in the armature of the head, thorax as well as the number, arrangement and size of the ventral teeth of the metatibiae. The two enlarged anterior coronals of the vertex vary considerably in size and general shape. Females show considerable variability concerning to the colouration. While mostly ranging from mostly plain ochre to dark brown (Figs. 458) some specimens have a slight greenish hue and occasionally specimens occur which have a more or less defined dark longitudinal median stripe along the entire dorsal body surface (Fig. 459). A ♀ collected by the fourth author in Berastagi had most of the body pale ochre with only the head, meso- and metapleurae as well as the legs darker brown and the tegmina with the anterior 2/3 dull brown (Fig. 459). No such variability in colour is seen in ♂♂. Eggs (Figs. 416–417): Of average size, barrel-shaped and fairly elongate, capsule circular in cross-section and 1.7x longer than wide. Anterior portion somewhat constricted, the polar-area slightly convex and roundly conical. Entire capsule surface irregularly pitted, the pits all with a rounded granule in centre; these granules becoming more decidedly raised and tubercle-like towards anterior margin of capsule and on operculum. Portion between micropylar plate and polar-area with a faint longitudinal bulge. Micropylar plate large principally X-shaped in outline with all four extension very short and broad; posterior extensions slightly directed posteriad, the anterior extensions very gently incurving and hardly longer than wide. Surface of micropylar plate like capsule. Entire plate surrounded by a broad and roundly raised bulge. Micropylar cup represented by a rounded impression marginated by a C-shaped carina ventrally; above micropylar cup a gently raised oval area. Operculum circular, moderately convex and roundly conical; surface as in capsule. General colouration dull grey, the outer margin of the operculum and micropylar plate black. Measurements [mm]: length incl. Operculum 7.0–8.4, length 6.2–7.7, width 3.8–4.5, height 4.1–5.0. Comments: This species is not rare in North Sumatra, being frequently found in the region of Berastagi, on Mount Sibayak and along the road leading from Berastagi to Lake Toba. It has been found on a native Ficus-tree as well as on Rubus moluccanus (Rosaceae) but feeding was not observed in the wild. Culture stock has been exported to Europe by Jimmy Gideon Ginting (Sumatra), who found several specimens on Mount Sibayak near Berastagi (Sumatera Utara Province) at an altitude of 1400–1600 m in November 2010. It was first successfully reared by Bruno Kneubühler (Luzern, Switzerland) in 2011 and since distributed as “Haaniella sp. Sibayak”. It accepts bramble (Rubus fruticosus, Rosaceae), rose (Rosa spp., Rosaceae), hazel (Corylus avellana, Betulaceae) and oak (Quercus spp., Fagaceae) as alternative foodplants. Breeding has proven fairly easy in slightly ventilated but humid conditions. At average temperatures of 25°C eggs hatch after an incubation of 8–12 months and nymphs reach maturity after 5–6 months. Females produce some 15 eggs per month. Distribution (Fig. 490): Sumatra, endemic. N-Sumatra: Prov. Aceh, Gunung Leuser NP, Ketambe, Atjeh, Tenggara 350 m [RMNH]; NW-Sumatra: Prov. Sumatera Utara, Mount Sibayak 1400–1600 m [coll. FH, coll. OC]; NW-Sumatra: Prov. Sumatera Utara, Berastagi 1500 m [coll. OC]; NW-Sumatra: Prov. Sumatera Utara, Jaranguda 1450 m [coll. OC]; NW-Sumatra: Prov. Sumatera Utara, road between Berastagi and Danau Toba, 1200–1400 m [coll. FSC]; NW-Sumatra; Prov. Sumatera Utara, edge of Sibolangit Botanic Gardens, 1220 m [coll. FSC]. Specimens examined: 56 TABLE 4. Measurements [mm] of Haaniella gintingi n. sp. ♀ HT [RMNH]

♂, PT [coll. FSC]

♀♀, PT [coll. FSC]

♂♂, PT [coll. FH]*

♀♀, PT [coll. FH]*

Body

98.4

65.0

90.0–100.0

61.0–65.5

84.0–103.0

Head

9.0

5.0

7.0–9.0

4.8–5.0

6.8–8.9

Pronotum

7.3

5.0

7.0–8.0

4.8–5.0

7.2–9.0

Mesonotum

15.2

12.0

15.0–17.0

11.9–12.3

14.2–16.5

......continued on the next page



41

TABLE 4. (Continued) ♀ HT [RMNH]

♂, PT [coll. FSC]

♀♀, PT [coll. FSC]

♂♂, PT [coll. FH]*

♀♀, PT [coll. FH]*

Metanotum

-

5.0–5.5

3.1–3.3

–

Median Segment

-

6.0–6.5

3.3–3.4

–

Tegmina

21.7

9.0

17.0–20.0

6.2–7.8

15.5–19.0

Profemora

19.3

17.0

20.0–25.0

15.0–15.5

18.5–22.0

Protibiae

22.3

18.0

22.0–24.0

15.2–16.3

20.0–23.0

Mesofemora

17.1

15.0

17.0–18.0

11.4–13.0

16.3–18.0

Mesotibiae

20.0

15.0

19.0–20.0

12.5–13.4

17.4–20.0

Metafemora

25.1

23.0

27.0–29.0

17.8–19.3

22.6–27.5

Metatibia

30.0

25.0

30.0–31.0

19.6–20.0

26.0–30.0

Antennae

-

53.0

60.0–63.0

53.0–55.0

55.0–62.0

*captive reared

Haaniella glaber (Redtenbacher, 1906) n. comb., rev. stat. (Figs. 79, 134–152, 358, 375–376, 402, 491) [Smooth-bodied Haaniella] Leocrates glaber Redtenbacher 1906: 167. HT, ♀: Lakat Sumatra, acheté des M. Schneider; Leocrates glaber Redt.; Heteropteryx glaber Redtb. 126. [MHNG]. [Synonymised in error with Haaniella muelleri (Haan, 1842) by Brock, 1995: 96] rev. stat. Dohrn, 1910: 401. Heteropteryx glaber, Karny, 1923 : 234 Haaniella mülleri simplex Günther, 1944: 72, fig. 1 (♀). HT, ♀: Mus. Leiden, F.J. Weynman, Tebing-.Tinggi, Haaniella mülleri n. ssp. Typus K. Günther Stett. Ztg. 1944, Abb. 1, gehört Mus. Leiden, Haaniella mülleri simplex Günther, 1944 Typus, Typus [RMNH]. n. syn. Brock, 1995: 96. [Erroneously listed as a synonym of Haaniella muelleri (de Haan, 1842)] Bragg, 1996: 112.

Material examined (2 ♂♂, 1 ♀, 2 nymphs): SUMATRA: 1 ♀: S. Sumatra, G. Tanggamoes, Giesting, 750m, VII.1934, L.J. Toxopeus, Heteropteryx glabra Redt. ♀ (= Haaniella mülleri simplex Gthr. 1944) H. Klante det. [RMNH]; 1 ♂ (nymph): Müller, Sumatra [RMNH]; 1 ♂: A.L. v. Hasselt. Riv. Papanoeli, Sumatra Aug. 1894; Heteropteryx mecheli Redt. K. Günther det. [RMNH]; 1 ♂, 1 ♀ (penultimate instar): C. J. Louwerens, Sumatra, Benkoelen, Lebong Tanday, 1927 [MNHN]. Diagnosis: This fairly distinctive species appears to be most closely related to H. gintingi n. sp. with which it shares the obsolete mesothoracic armature of ♀♀, weakly armed mesonotum of ♂♂, large, crenulate and multidentate anterior coronals, large triangular sub-apical lobe on the posterodorsal carina of the femora and conspicuously broadened, black triangular teeth on the ventral carinae of the metatibiae of both sexes. It however differs from H. gintingi n. sp. by the much more stocky appearance (♀♀ in particular) and broader, relatively shorter legs of both sexes. Females furthermore differ by the strongly widening, trapezoidal mesothorax, laterally expanded abdominal tergites II–VI (only IV distinctly widened in gintingi), larger tegmina, different structure of the praeopercular organ and angular to slightly notched apex of the subgenital plate (Fig. 149). Males are readily distinguished from those of H. gintingi n. sp. by the plain brown colouration and being macropterous with the alae almost covering the entire abdomen (Figs. 141–143). The long alae and elongated tegmina are shared with ♂♂ of H. aculeata n. sp., H. macroptera n. sp. and H. mecheli (Redtenbacher, 1906). The more elongate and considerably more slender mesothorax, very minute spines of the mesonotum and characteristic crenulate anterior coronals of the vertex however readily distinguish ♂♂ of H. glaber from these three species. The convex vertex of both sexes,


HENNEMANN ET AL.

massive body, strongly widening, trapezoidal mesothorax and laterally dilated abdominal tergites II–VI of ♀♀ resemble the genus Heteropteryx Gray, 1835 and emphasize the close relation between the muelleri species-group of Haaniella and that genus. Etymology: The name (glaber lat. = smooth) refers to the unarmed mesothorax and abdominal tergites of ♀♀ this species. Description: Since no live specimens were available for examination, the colouration is described from preserved specimens only. The previously unknown ♂ is here described for the first time. ♀♀ (Figs. 138–140). Of average size (body length 86.0–88.4 mm), very broad and massive for the genus, with much reduced body armature, strongly laterally dilated abdominal tergites III–V and prominent cephalic projections. General colour plain ochraceous or greyish mid to dark brown, sometimes with a slight greenish hue. Tegmina of same colour as body, sometimes with a fine green line along basal half of anterior margin or with two black basal markings on both sides of the radial vein. Head a little longer than wide, globose with vertex conically elevated and prominently armed (Fig. 144). Anterior coronals strongly enlarged and forming laterally compressed, bi-dentate projections; posterior coronals much smaller and represented by simple but strong spines (Fig. 144). Anteriorly only with a pair of very minute supra-occipitals (sometimes indistinct). Posterior margin with two small but strong lateral coronals, which are considerably smaller than the posterior coronals. Behind the eyes with a short supra-orbital spine about equal in size to the lateral coronals. Antennae ± reaching to posterior margin of abdominal tergum IV; with 24 antennomeres. Pronotum longer than head, about 1.2x longer than wide, widening towards posterior and decidedly narrowed medially. Antero-lateral pronotals very small and blunt, the postero-laterals somewhat more pronounced and the pair of inter-posterior pronotals very minute and considerably smaller than the outer posterior pronotals. Some distance before transverse median sulcus with a pair of low and very small pre-median pronotals (Fig. 144). Mesothorax strongly, gradually widening towards the posterior with posterior portion some 2x wider than anterior margin. Mesonotum very slightly narrowing in the posterior half, almost 2x longer than wide and roughly 2x the length of pronotum. Lateral margins with 3–4 very minute spiniform tubercles, the disc merely set with with 6–8 small pair-wise arranged spiniform tubercles (Fig. 144). Between bases of tegmina with two fairly prominent, stout and rather blunt posteromedial mesonotals. Tegminal spine short but acutely pointed (Fig. 144). Mesopleurae with a fairly distinct but short supra-coxal spine and 5–6 unequally sized, stout laterals along lower margin (Fig. 144). Metapleurae with lateral margin strongly rounded and deflexed medially and armed with five moderately prominent spines, the posterior three spines in the deflexed portion decidedly longer than the anterior ones; supracoxal obsolete. Sensory area of profurcasternum very large, roughly triangular in outline and covering most of the segments surface; with longitudinal median furrow in posterior portion (Fig. 79). Meso- and metasternum set with a variable number of blunt spiniform tubercles. Tegmina fairly large, oval in shape and slightly projecting over posterior margin of abdominal tergum II. Alae translucent dark grey with black veins. All abdominal tergites smooth dorsally. II–III widening, IV widest and almost 3.5x wider than long, V–X strongly narrowing. Lateral margins of III–VI strongly deflexed and weakly rounded but most distinct in IV; in II–V armed with 5–6 very small spines, the remaining with lateral margins unarmed. Sternites II–VI with four pairs of spines, these most prominent on II and decidedly decreasing in size towards VI. Praeopercular organ on sternum VII formed by two distinct and acutely pointed spines near posterior margin of segment (Fig. 149). Posterior margin of anal segment rounded (Fig. 148). Epiproct tectiform longitudinally, gradually narrowing towards a slightly downcurving and triangularly incised, acutely bifid apex (Figs. 147–149). Apex of subgenital plate broad, angular, with a shallow median indention and roughly reaching half way along epiproct (Fig. 149). All legs moderately long and fairly stocky, mesofemora somewhat longer than mesothorax, metafemora slightly projecting over posterior margin of abdominal segment V and metatibiae ± reaching apex of abdomen. All carinae except anterior carinae of pro- and mesotibiae dentate; the femora with an enlarged and prominent triangular pre-apical tooth on posterodorsal carina. Two outer ventral carinae of metatibiae armed with 6–8 prominent and strong, triangular teeth, some of these in the basal portion paired and the two apical ones smaller than the median teeth (Fig. 375). Ventral surface of all femora smooth (Fig. 358). ♂♂ (Figs. 141–143). Medium sized (body length 68.0–69.8 mm) and slender for the genus with rather reduced thoracic spination but very prominent head armature, long tegmina and fully developed alae. Colour dull ochraceous brown, the mesonotum and mesosternum dull drab. Mesopleurae and lateral margins of mesonotum dark brown. Body armature dull reddish brown with black tips. Tegmina plain mid to dark brown with most of the



43

anterior margin creamish white to pale yellow. Femora ochracheous brown, the tibiae dull ochre. Armature of legs with black tips, the ventral teeth of the metatibiae contrasting black. Shape and armature of head as in ♀♀, but armature much more prominent with all spines much longer and more slender. The anterior coronals very large and forming laterally compressed, crenulate and lobe-like lobes (Fig. 146). Antennae broken in all examined specimens. Pronotum about 1.2x longer than wide, somewhat longer than head with the lateral margins very gently concave; armature much more prominent than in ♀♀. Antero-lateral pronotals short and fairly robust, the postero-lateral pronotals decidedly longer and more slender; the interposterior pronotals roughly equal in size to the outer postero-lateral pronotals but somewhat more slender. Before transverse median sulcus with a pair of prominent and strong pre-median pronotals (Fig. 146). Mesothorax very slender and elongate, almost 2x longer than pronotum with posterior portion widened and the mid portion gently constricted in dorsal view (Fig. 145). Mesonotum almost parallel-sided and averaging 4x longer than wide; disc smooth except for a small pair of anterior mesonotals, as well as a pair of minute pre-median and post-median mesonotals. Between the bases of the tegmina with a pair of fairly prominent, slender and upright posteromedial mesonotals (Fig. 146). Tegminal fairly distinct and somewhat anteriad directed. Mesopleurae only with 3–4 spines along lower margin which gradually increase in size towards the posterior; supra-coxal prominent and acutely pointed. Metapleurae strongly and almost semicircularly deflexed in the median portion, armature as in ♀♀ but spines comparatively longer and more slender; supra-coxal spine of moderate size and smaller than that of the mesopleurae. Sensory area of profurcasternum distinct, ovate in shape and longitudinally divided by a median furrow. Mesosternum smooth except for 2–3 pairs of short and stout spines in the posterior half and fairly distinct pair of posterior mesosternals between the coxae. Metasternum with ten short and stout paired spines. Abdominal sternum I with a pair of short spines near posterior margin. Tegmina elongate, slender, some 3.5x longer than wide, slightly narrowing towards the apex and reaching about half way along abdominal tergum IV; some 3/5 the length of alae. Sub-basally with a fairly prominent roundly conical and anteriad directed hump. Alae fully developed and reaching to abdominal tergum VIII or IX, the anal fan transparent grey with a slight orange wash and the veinature brown. Abdominal segments II–IV increasing and, VI–VII decreasing in length, IV longest and a little more than 2x longer than wide. II slightly trapezoidal in shape, III–VI roughly parallel-sided. Tergites smooth, except for a pair of small posteromedian humps on VII–IX. Lateral margins smooth except for a single, slender and backward directed spine near the posterior angle. Tergum VII somewhat widenen in the posterior half with the lateral margins slightly deflexed and gently rounded. Sternites with four pairs of spines, these increasing in size from II towards VI. Anal segment almost rectangular in dorsal aspect, gently narrowed medially and the posterior margin with a broadly triangular indention; outer lateral angles expanded and broadly rounded (Fig. 151). Epiproct small, shieldshaped, almost semicircular and with a fine longitudinal median carina (Fig. 402). Cerci small, club-like, slightly curved and compressed laterally in basal portion. Vomer with the apical portion strongly narrowed and gradually tapered towards a gently curved apical hook; pale reddish brown with the apex black (Fig. 402). Poculum large, reaching about half way along anal segment, tri-carinate in the posterior portion and the posterior margin roundly angular (Fig. 152). Legs all moderately long and slender, mesofemora reaching more than half way along pronotum, metafemora ± reaching half way along abdominal segment VI and metatibiae projecting considerably beyond apex of abdomen. Protibiae entire smooth, dorsal carinae of mesotibiae smooth and dorsal carinae of metatibiae only with one or two very minute teeth sub-basally. All other carinae dentate and the posterodorsal carina of mesofemora with an enlarged triangular, sub-apical tooth. Metatibiae with armature as in ♀♀, but teeth relatively more prominent and acutely pointed (Fig. 376). Ventral surface of all femora smooth. Nymphs: As in other species of the genus nymphs of this species have the body and leg armature much stronger developed and the legs relatively shorter and broader than adults. Variability: Since only a very few specimens are known, the range of variability in this species cannot be fully defined. The armature of the body and legs of the examined specimens does not show any considerable variation. While the ♀ type specimens of H. glaber and H. muelleri simplex merely have a small and slender white sub-basal marking at the anterior margin of the tegmina, the ♀ from Mount Tanggamus in RMNH has a large but irregularly shaped blackish latero-basal marking and a further but much smaller black basal marking interior of the radial vein. Furthermore, the latter specimen has the posterior half of the tegmina darker brown than the basal half, but this may be due to the insect was freshly moulted when captured. Comments: There has been considerable confusion about the identity of this species in the past. Günther (1944: 69) omitted the Sumatran H. glaber (Redtenbacher, 1906) from his arrangement of the genus Haaniella


HENNEMANN ET AL.

Kirby, 1904, which shows he regarded it as a member of Heteropteryx Gray, 1835, as did Redtenbacher (1906: 167) who originally described this species in the genus Leocrates Stål, 1875, a synonym of Heteropteryx. Brock (1995: 96) erroneously synonymised H. glaber species with Haaniella muelleri (Haan, 1842). However, examination of the ♀ HT in MHNG has clearly shown H glaber is a distinct species, clearly differing by e.g. the more convex vertex, which bears a pair of conspicuous crenulate extensions, strongly broadening trapezoidal mesothorax, very broad abdomen and more prominent triangular ventral teeth of the metatibiae. Consequently, H. glaber is here reestablished (rev. stat.). Günther (1944: 72) described Haaniella muelleri simplex based on a single ♀ from Tebing Tinggi, NE-Sumatra in RMNH, which was erroneously listed as a synonym of H. muelleri by Brock (1995: 96). Examination of Günther’s specimen in RMNH has shown this subspecies to be conspecific with H. glaber (n. syn.). Although widely distributed throughout Sumatra, this species has been very rarely encountered and only seven specimens have so far been recorded. Nothing is known about the natural habits and host plants of H. glaber. Eggs unknown. Distribution (Fig. 491): Sumatra, endemic. E-Sumatra: Prov. Riau, Lakat [MHNG—type-locality]; SSumatra: Prov. Lampung, Giesting, Mount Tanggamus 750 m [RMNH]; SW-Sumatra: Prov. Benkulu, Lebong Tandai [MNHN]; NE-Sumatra: Prov. Sumatera Utara, Deli Serdang, Tebing Tinggi [RMNH]; W-Sumatra: Prov. Sumatera Utara, Tapanuli district, Kapanoek River “Papanoeli” [RMNH]. Specimens examined: 7 TABLE 5. Measurements [mm] of Haaniella glaber (Redtenbacher, 1906) ♀ HT [MHNG]

♀ HT of muelleri simplex [RMNH]

♀ [RMNH]

♂ [RMNH]

♂ [MNHN]

Body

86.0

88.4

86.5

68.0

69.8

Head

-

6.2

6.8

4.7

5.5

Pronotum

7.6

7.4

6.8

5.3

6.0

Mesonotum

13.8

14.2

14.2

12.3

12.3

Metanotum

5.5

-

-

4.0

-

Median Segment

4.0

-

-

3.9

-

Tegmina

21.0

19.9

18.1

22.8

22.8

Alae

18.0

16.0

-

36.0

40.0

Profemora

17.2

17.8

17.8

17.3

18.0

Protibiae

18.6

20.2

20.2

18.7

18.7

Mesofemora

14.9

15.3

15.3

14.2

15.5

Mesotibiae

17.0

16.9

16.9

15.3

16.9

Metafemora

23.1

22.5

22.5

20.0

21.2

Metatibia

26.5

25.2

25.2

22.2

23.1

Antennae

> 50.0*

-

58.0

> 17.0*

> 34.0*

* apex broken

Haaniella jacobsoni Günther, 1944 (Figs. 153–160, 492) [Jacobson’s Haaniella] Haaniella jacobsoni Günther, 1944: 73, figs. 3 & 4. HT, ♂: E. Jacobson, Sinabang (Simalur) Sumatra III.1913; Haaniella jacobsoni ♂ Typus K. Günther 1944; Typus, RMNH Leiden Holotype [RMNH]; AT, ♀: S.A.J. Voorthuis, Sinabang Simaluee 28. April 1918; Haaniella jacobsoni n. sp. ♀, Allotypus, K. Günther 1944; Typus Allotyp, Paratype [RMNH];



45

PT, ♂: S.P. Buitendyk, Poeloe Babi, Simalur, II.1908; Haaniella jacobsoni Gtr., 1944, Paratypus [SMTD]. Bragg, 1996: 111. Zompro, 2004: 198. Otte & Brock, 2005: 149.

Diagnosis: This characteristic species is easily distinguished from all other members of the muelleri species-group by the prominent posteromedian pair of spines on the metanotum and somewhat less distinct median pair of spines on the median segment, which are positioned between the tegmina and alae. The wings are smaller than in all other species, slender, not overlapping but leave a free space inbetween them and do not reach the posterior margin of the median segment. Further characteristic features of H. jacobsoni are the prominent leg-armature and the prominent, rounded lateral expansions of abdominal tergites IV–VI of ♀♀, as well as the dorsally spinose abdominal tergites and rounded lateral lobes of tergites VII–VIII of ♂♂. The latter two features are only shared with H. kerincia n. sp., which appears to be the most closely related species. Also in H. kerincia n. sp. the wings are not overlapping, but they are comparatively larger than in H. jacobsoni and do not expose a considerable portion of the metanotum and median segment inbetween them. Furthermore, this species differs by the very long and dichotomic posteromedian pronotals and posteromedian mesonotals as well as the much smaller size. Etymology: Named after the collector E. Jacobson (Günther, 1944: 77). Description: Since this species is so far only known from the three original type specimens, the colouration is described from preserved specimens only. That of the ♀♀ is based on the allotype and that of the ♂♂ on the holotype in RMNH. The paratype ♂ in SMTD is obviously discoloured due to provisional preservation in ethanol, hence cannot serve for describing the natural colouration. ♀♀ (Figs. 153–154). Medium sized species (body length 90.4 mm) with prominent body aramture, strongly laterally expanded abdominal tergites IV–VI and very small, slender and not medially overlapping tegmina and alae. General colouration reddish mid brown and all over with dark brown mottling and speckles. All femora with an irregularly shaped dull ochre median transverse band, the tibiae with two or three irregular transverse annulations. Tegmina with the anterior margin dull cream. Head slightly longer than wide, globose with the vertex gently convex and armed with ten spines. Vertex with four strong coronals which roughly form the four corners of a square; the anterior coronals more prominent than the posterior ones. Anteriorly with a pair of smaller, very slender supra-occipitals. Posterior margin with two fairly distinct lateral coronals, which are roughly equal in size to the posterior coronals. Behind the eyes with a fairly distinct supra-orbital spine, about equal in size to the pair of lateral coronals. Antennae at least reaching to abdominal tergum II (tips broken in the unique specimen available). Pronotum longer than head, some 1.3x longer than wide with lateral margins distinctly concave and posterior margin somewhat wider than anterior margin. Antero-lateral pronotals distinct and accompanied by a much smaller spine posteriorly, postero-lateral pronotals distinct and somewhat larger than the antero-lateral pronotals, near posterior margin with a somewhat smaller pair of inter-posterior pronotals and just before transverse median sulcus with a very strong pair of pre-median pronotals. Mesothorax gently and gradually widening towards the posterior with posterior portion about 1.7x wider than anterior margin. Mesonotum roughly parallel-sided, almost twice as long as wide and about 2x the length of pronotum. Lateral margins set with about five fairly small and slender regularly spaced spines, the disc with a pair of prominent outward directed anterior-mesonotals, a very strong pair of pre-median mesonotals close to median line and a further pair of slightly smaller and more distant post-median mesonotals. Four distinct and fairly strong posteromedial mesonotals on a swollen common base between bases of tegmina; the anterior two spines somewhat smaller than the posterior pair. Tegminal of moderate size. Mesopleurae with a strong and very prominent supracoxal spine and about eight prominent but unequally sized spines along lower margin, the two median ones of which are very prominent, gently back-curving and much larger than the remaining (Fig. 154). Metapleurae with lateral margin deflexed, semicircularly rounded and armed with five spines, the three median ones of which are considerably larger than the other two; supra-coxal prominent but smaller than that of the mesopleurae. Metanotum with a fairly distinct slightly posteriorly directed pair of posteromedian spines between the tegmina. Sensory area of profurcasternum well developed and roundly triangular in outline, this segment furthermore with a very distant pair of minute posterolateral spines. Mesosternum with a lateral row of four well developed spines, which distinctly increase in size towards the posterior. Metasternum with 3–4 spines on each side of the median line, the outermost spine noticeably larger than the remaining. Tegmina small, slender, elongate-oval, reaching about 2/3 along median segment and not overlapping, leaving a free space between each other. Alae somewhat shorter than


HENNEMANN ET AL.

tegmina. Median segment with a small pair of central spines and a transverse row of four minute spines near posterior margin. Abdominal tergites II–VII with a very small pair of latero-anterior spines, a very minute to obsolete median pair of spinules and a transverse row of six minute spines close to posterior margin. II–IV strongly widening, following narrowing and IV widest segment; all transverse. IV–VI with lateral margins strongly deflexed into an almost semicircular, dentate lobe; II–VI each with four short spines along lateral margins, which are most prominent on IV–VI. Tergum VII only with three and VIII with two very minute lateral spines. Sternites II–VI with four spines on each side of the median line; these most prominent on II and decreasing in size towards VI. Praeopercular organ on sternum VII formed by two blunt spine-like posteromedian swellings. Posterior margin of anal segment gently concave with the outer angles fairly acute (Fig. 158). Epiproct tectiform longitudinally, slightly downcurving, narrowed in the apical half with the apex notched and forming two short but acute points (Fig. 158). Subgenital plate strongly convex with the apex roundly angular and only reaching about 1/3 along epiproct (Fig. 157). All legs long and slender, metafemora reaching to anterior margin of abdominal segment VII and metatibiae distinctly projecting beyond apex of abdomen. Protibiae entirely unarmed. All other carinae, except for the dorsal carinae of the mesotibiae, acutely dentate. The apicalmost tooth on the posterodorsal carina of the femora somewhat enlarged. Metatibiae with the dorsal carinae very minutely and sparsely dentate and the ventral carinae with a variable number of medium-sized triangular teeth, which decrease in size towards the base of tibia. Ventral surface of all femora smooth. ♂♂ (Figs. 155–156). Medium sized (body length 67.0–68.8 mm) and moderately slender with prominent body armature, dorsally spinose abdominal tergites and small, very slender and not overlapping tegmina. General colour dull drab to ochre and sometimes with a slight greenish hue. Dorsal portion of mesonotum straw and thoracic armature dull green basally with the apical portion dark brown. Mesosternum pale ochre, median portion of abdominal sternites II–VI reddish mid brown with an orange wash. Tegmina with a weakly defined pale basal or sub-basal lateral marking and a faint, washed and roughly oval blackish median lateral marking; the anterior margin may be dull green in the basal half. Femora with an irregular, straw to ochre median transverse band, the tibiae irregularly marbled with straw and dark brown. Shape and armature of head as in ♀♀, but armature much more prominent with all spines considerably longer and more slender. Antennae broken in all specimens available. Pronotum about 1.3x longer than wide, a little longer than head with the lateral margins gently concave; spines as in ♀♀ but more slender and the inter-posterior pronotals considerably larger. Mesothorax slender and fairly elongate, some 2.5x longer than pronotum with posterior portion gently widened and the mid portion very slightly constricted in dorsal view (Fig. 155). Mesonotum almost parallel-sided and a little more than 4x longer than wide; disc with six spines arranged as in ♀♀, but all more elongate and slender and the antero-lateral mesonotals somewhat more distinct. The spines along the lateral margins small and much less developed than in ♀♀. Posteromedian swelling between bases of tegmina more prominent than in ♀♀ with the four spines considerably longer and more slender. Mesopleurae with a very long and slender supra-coxal and about six strong laterals along lower margin; two spines in the posterior half much enlarged and noticeably longer than the remaining. Armature of metapleurae as in ♀♀, the supra-coxal very prominent and long. Profurcasternum as in ♀♀. Metanotum with a distinct posteromedian pair of spines between the tegmina. Mesosternum smooth except for a longitudinal lateral row of three spines, which increase in size towards the posterior, and a pair of small, blunt posterior mesosternals between the coxae. Metasternum as in ♀♀, but spines more developed and the outermost spine in particular very prominent. Tegmina very small, elongateoval and reaching less than half way along median segment; not overlapping and leaving a free space between each other. Alae slightly projecting under tegmina medio-posteriorly. Median segment armed with a distinct median pair of laterally directed spines, a minute median spine on each side and a transverse row of four minute spines close to posterior margin, the inner ones of which are noticeably larger than the outer two spines. Abdominal segments II– IV slightly increasing, V–VII decreasing in length; IV longest segemnt and about 2.3x longer than wide. II gently narrowing towards posterior, III–VI roughly uniform in width. Tergites II–VII with a minute pair of latero-anterior spines, a small pair of median spines and a transverse row of six spines close the posterior margin; the four inner spines very prominent, long and slender. Lateral margins of II–VI with three small, slender spines in the posterior half, this portion gently deflexed and rounded in V and VI. Tergum VII with the lateral margins very strongly deflexed to form a large, roughly semicircular to angular lobe, which laterally extends by more than half of the segments width (Fig. 160). VIII with a similarly shaped but broader lateral lobe and also lateral margins of IX slightly deflexed (Fig. 160). Anal segment gently tectiform, narrowed medially and the posterior margin with a distinct, roundly triangular median excavation; the outer lateral angles expanded, slightly protuded ventrally and REVISION OF HETEROPTERYGINAE


47

roundly triangular. Cerci small, laterally compressed and with the apex broadly rounded. Vomer triangular. Sternites II–VI with four spines on each side of the raised median longitudinal carina; these most prominent on II and decreasing in size towards VI. Poculum strongly convex with posterior margin angular, distinctly labiate, downcurving and notched medially (Fig. 159). Legs all long and moderately slender, mesofemora somewhat longer than mesonotum, metafemora reaching to abdominal segment VI and metatibiae projecting distinctly beyond apex of abdomen. Armature generally as in ♀♀ but more pronounced, the apical tooth on the posterodorsal carina of the femore more decidedly enlarged. Eggs: The ♀ allotype in RMNH has an egg in its ovipositor, which is however impossible to remove without dissecting the specimen. From an external view it shows to be of the typical lemon-like shape seen in other members of the muelleri species-group, having the polar-area distinctly conical. It is plain dark grey in colour and has a distinctly pitted capsule surface. The micropylar plate is surrounded by a very dark grey bulge and the posterior extensions of the plate have the apex strongly expanded dorsally to form a rounded hook-like shape. Comments: Nothing is known about the natural habitat and host-plants of this very characteristic species, which is the only representative of the genus known to occur on the Mentawai islands. It is hoped that future collections on these islands but Simeulue in particular will reveal fresh material. The three types in RMNH and SMTD are still the only known specimens of H. jacobsoni and there have been no records since its description in 1944. Distribution (Fig. 492): So far only known from Simeulue Island (Prov. Sumatera Barat), 150 km off the west coast of Sumatra (Sinabang [RMNH] and Poeloe Babi [SMTD]). Endemic. Specimens examined: 3 TABLE 6. Measurements [mm] of Haaniella jacobsoni Günther, 1944 ♂, HT [RMNH]

♂, PT [SMTD]

♀, AT [RMNH]

Body

68.8

67.0

90.4

Head

5.5

5.2

7.0

Pronotum

5.8

5.6

7.2

Mesonotum

14.7

14.5

16.8

Metanotum

4.0

3.9

5.6

Median Segment

4.9

4.8

5.8

Tegmina

8.0

7.0

13.2

Profemora

17.0

18.5

19.8

Protibiae

18.9

20.0

22.7

Mesofemora

14.7

15.3

17.1

Mesotibiae

15.0

16.2

19.8

Metafemora

20.3

21.0

25.0

Metatibia

22.0

23.0

28.2

Antennae

>20.0

>60.0

>47.0

Haaniella kerincia Hennemann, Conle, Brock, Seow-Choen & Gorochov n. sp. (Figs. 161–170, 448–449, 489) [Spiny Mt. Kerinci Stick-insect] HT, ♂: Sumatra: Prov. Jambi, 35km N. of Sungaipenuh, env. of Nat. park Kerinci-Seblat, Mt. Kerinci, 1500–2000 m., 18– 22.xi.1999 (A.V. Gorochov) [ZIN]. PT, 1 ♂, 2 ♀♀: Sumatra: Prov. Jambi, 35km N. of Sungaipenuh, env. of Nat. park Kerinci-Seblat, Mt. Kerinci, 1500–2000 m., 18–22.xi.1999 (A.V. Gorochov) [ZIN].


HENNEMANN ET AL.

Diagnosis: This remarkable new species is the smallest member of the muelleri species-group and second smallest member of the genus. The most closely related species appears to be H. jacobsoni Günther, 1944 from Simeulue Island, with which it shares the strongly developed body and leg armature, laterally deflexed abdominal tergites II– VI of ♀♀ and broad lateral lobe of abdominal tergites VII–IX of ♂♂. The very large, dichotomic and antler-like pre-median pronotals and posteromedial spines of the mesonotum however are unique in the entire Heteropteryginae and readily distinguishe H. kerincia n. sp. from all other species of Haaniella. Etymology: Named after the type locality Mount Kerinci in the Jambi province of central Sumatra. Description: ♀♀ (Figs. 161–162, 164). Small (body length 56.0–57.0 mm) and heavily spinose with very long and dichotomic to antler-like pre-median pronotals, the posteromedian mesonotals represented by a multispinose swelling and abdominal tergites II–VI strongly laterally deflexed. Colour pale to dark brown, either almost plain or all over with straw to pale ochre markings and speckles. Mottled specimens usually with a bold V-shaped marking on mesonotum and a pale post-median transverse band on femora. Plain specimens may have black markings on most of the thoracal segments. Rarely specimens possess a slight greenish hue. Tegmina plain mid to dark brown. Larger spines of thorax black at the base with the apical half dull ochre. Head a little longer than wide, globose with vertex roundly convex and prominently armed (Fig. 170). Anterior coronals more prominent than all other cephalic spines and often bifid, with the anterior point shorter than the posterior one; posterior coronals somewhat smaller and represented by simple but strong spines (Fig. 170). Anteriorly with a pair of small and stout supra-occipitals. Posterior margin with two fairly distinct but stout lateral coronals, which are slightly smaller than the posterior coronals. Behind the eyes with a fairly indistinct supraorbital spine. Antennae ± reaching to abdominal tergum III; with 24 antennomeres. Pronotum somewhat longer than head and about as wide as long with the posterior portion wider than the anterior and the lateral margins slightly concave. Antero-lateral pronotals distinct and slender, the posterior pronotals about equal in size and the pair of inter-posterior pronotals very indistinct or obsolete. Behind transverse median sulcus with two pairs of small post-median pronotals represented as acute tubercles. Just before transverse median sulcus with a pair of very prominent, long and slender bifid pre-median pronotals (Fig. 170). Mesothorax gently and gradually widening towards the posterior with posterior portion about 1.8x wider than anterior margin. Mesonotum very slightly narrowing towards the posterior, some 1.3x longer than wide and roughly 1.6x the length of pronotum. Lateral margins with a variable number of very short and stout, rather blunt spines. Disc with strong and long anterior mesonotals, a small pair of anterior-mesal mesonotals and two pairs of pre-median mesonotals; the anterior pair very prominent, long and slender, the posterior pair much smaller and about equal in size to the anterior-mesal mesonotals (Fig. 170). Between bases of tegmina with a prominent rounded swelling, which is armed with a cluster of about 10–12 unequally sized, but fairly short spines (Fig. 164). Tegminal spine small. Mesopleurae with a very long and prominent supra-coxal spine and 6–7 unequally sized, pointed laterals along lower margin. Metapleurae with lateral margin moderately rounded and armed with five fairly prominent, slender spines, the median three spines in the deflexed portion decidedly longer than the others; supra-coxal distinct. Sensory area of profurcasternum moderately developed. Mesosternum with a row of three lateral spines, which strongly increase in size towards the posterior. Tegmina oval in shape and projecting distinctly over posterior margin of abdominal tergum II; radial vein strongly raised and forming a rounded keel sub-basally. Alae translucent dark grey with black veins, sometimes reaching slightly beyond apex of tegmina. Wings hardly overlapping. Abdominal tergites II–VI sparsely tuberculose with a transverse row of six more acute tubercles near posterior margin. Lateral margins of II slightly, those of III–VI strongly deflexed and forming a dentate, semicircular lobe (most prominent in IV–VI); all armed with 5–6 short but acute, triangular teeth. Lateral margins of VII deflexed and semicircularly rounded but unarmed. Sternites II–VI with eight fairly prominent spines, which decrease in size from II towards VI. Praeopercular organ on sternum VII formed by two short and blunt spines and a small, rounded median swelling just in front of these. Posterior margin of anal segment broadly rounded with a shallow median indention. Epiproct tectiform longitudinally, narrowing towards a slightly downcurving and triangularly incised, acutely bifid apex; gently constricted in median portion (Fig. 167). Apex of subgenital plate bluntly bi-dentate with a very deep, trinagular median excavation and almost reaching half way along epiproct (Fig. 166). All legs moderately long and slender, mesofemora considerably longer than mesothorax, metafemora slightly projecting over posterior margin of abdominal segment VI and metatibiae distinctly projecting beyond apex of abdomen. All carinae acutely but irregularly dentate. Two outer ventral carinae of metatibiae armed with several enlarged, triangular black teeth. Ventral surface of all femora smooth.



49

♂♂ (Figs. 163, 165). Small (body length 40.0–43.0 mm) and slender for the genus with very long and dichotomic or antler-like spines on head and thorax and long, slender, acutely dentate legs. Colour variable, ranging from dull ochraceous brown (holotype) to dark brown (paratype), the holotype with most of the mesonotum, median portion of abdominal tergites VIII–IX drab and a large triangular drab marking which extends over abdominal segments II and III. All larger spines of head, thorax and legs dull greenish ochre. Abdominal tergum VIII with a straw anterolateral marking in the paratype. Tegmina often with a washed dark marking in the central area. Legs dull ochracheous brown to dark brown, the femora with the basal half paler in colour and often with a faint pale post-median transverse band. Two of the three live ♂♂ examined from photographs taken on Gunung Tujuh, Kerinci-Seblat NP by Jeremy Holden (Vietnam) have a pale straw or whitish anterolateral marking on abdominal tergum VII and the entire anterior margin of the tegmina pale ochre or pale yellow. One of these has the entire body armature contrasting pale yellow, while the armature is mostly dull orange in the third specimen. Head about as wide as long, armature as in ♀♀, but armature more prominent with all spines much longer and more slender. Supra-orbital spine distinctly longer than in ♀♀. Antennae reaching to abdominal segment V; with 22 antennomeres. Pronotum about 1.3x longer than wide, somewhat longer than head with the lateral margins gently concave; spines as in ♀♀ but the bifid pre-median pronotals much larger, decidedly longer than all other spines and occasionally with a further much shorter spine towards anterior. Mesothorax some 1.6–1.7x longer than pronotum with posterior portion widened. Mesonotum gently narrowing towards the posterior and averaging 2.3x longer than wide; disc with a pair of very long and slender anterior mesonotals and one or two pairs of extremely long and slender pre-median mesonotals, otherwise rugulose. Lateral margins with a row of blunt spiniform tubercles and occasionally with two fairly long and slender spines in anterior half. Posteriomedial mesonotals between the bases of the tegmina extremely long, dichotomic and antler-like, slightly directed posteriad, at least bifid and often accompanied by one or two further but smaller anterior spines (Fig. 165). Tegminal small. Mesopleurae with a very prominent, long and slender supra-coxal spine and two strong spines in posterior half of lower margin; this margin otherwise with a few spiniform tubercles. Armature of metapleurae as in ♀♀ but supracoxal spine very prominent, extremely long, slender and almost extending by half of body width; also the other spine along lower margin much longer than in ♀♀. Sensory area of profurcasternum fairly distinct. Mesosternum rugose with three a lateral of three fairly distinct but blunt spines, which decidedly increase in size towards posterior; the pair of posterior sternals well pronounced. Metasternum with a variable number of minute spines and a distinct lateral spine just before excavation of metacoxae. Tegmina ovoid in outline with the apical portion gradually narrowing and not overlapping, leaving a small space inbetween them; radial vein conspicuously raised and forming a rounded keel sub-basally. Tegmina ± reaching to posterior margin of median segment; alae slightly projecting beyond apex of tegmina and reaching almost half way along abdominal tergum II. Abdominal segment II slightly narrowing, II–VI roughyl equal in width, parallel-sided and averaging 1.3x longer than wide. Tergites all sparsely and irregularly rugulose and tuberculose, lateral margins of II–VI with 3–4 very short and indistinct spines or tubercles in posterior portion. Tergum VII with lateral margin moderately deflexed and rounded, VIII with lateral margins strongly widened and almost semicircular, IX with a roundly angular lateral lobe in posterior portion (Fig. 169). Sternites II–VI armed with two pairs of distinct elongate spines, which are most prominent on II and decrease in size towards VI; posterior pair considerably larger than anterior pair. Sternum VII only with paired tubercles. Anal segment somewhat narrowing towards posterior, the posterior margin with a shallow median indention and the outer angles broadly rounded (Fig. 169). Epiproct very short. Vomer fairly small, triangular, somewhat longer than wide with the medio-basal portion broadly impressed and with an almost straight, black terminal hook; dull ochre. Poculum with posterior margin roundly rectangular, labiate and downcurving (Fig. 168). Legs all long and slender, mesofemora almost as long as pro- and mesonotum combined, metafemora ± reaching half way along abdominal segment VII and metatibiae projecting considerably beyond apex of abdomen. All carinae of acutely armed with pointed and fairly slender, spine-like teeth, although only very minute teeth on dorsal carinae of tibiae. Two outer ventral carinae of metatibiae armed with 7–8 strong and rather triangular teeth. Ventral surface of all femora smooth. Variability: Although so far only known from a total of seven specimens, three of them examined from photographs of live insects only, considerable variability is seen in the colouration as well the number and size of the thoracic spines. The latter concerns to ♂♂ in particular. A specimen photographed on Gunung Tujuh in the Kerinci Seblat National Park has the entire body armature contrasting pale yellow and considerably more strongly developed than all the ♂♂ males examined (Fig. 449).


HENNEMANN ET AL.

Comments: Eggs unknown. Distribution (Fig. 489): Central Sumatra, so far only known from Mount Kerinci and Mount Tujuh in the Kerinci-Seblat National Park in the Jambi province, an area well known for its rich fauna and flora and harbouring endemic species. Obviously peculiar to higher altitudes, since all records are from highly mountainous, mossy forests between 1500 and 2000 m. Specimens examined: 4 (7). TABLE 7. Measurements [mm] of Haaniella kerincia n. sp. ♂, HT [ZIN]

♂, PT [ZIN]

♀♀, PT [ZIN]

Body

43.0

40.0

56.0–57.0

Head

4.0

3.0

5.0

Antennae

20.0*

29.0

27.0–28.0*

Pronotum

4.0

4.0

5.5

Mesonotum

7.0

6.5

7.0

Metanotum and median segment combined

6.0

6.0

6.0

Tegmina

7.0

6.5

12.0–13.0

Alae

5.5

5.5

10.0–11.0

Profemora

12.0

12.0

12.0

Protibiae

13.5

13.0

13.0

Mesofemora

10.0

10.0

11.0

Mesotibiae

11.0

11.0

12.0

Metafemora

14.0

13.5

16.5

Metatibia

16.0

16.0

18.5

* tips broken

Haaniella macroptera Hennemann, Conle, Brock & Seow-Choen n. sp. (Figs. 80, 171–181, 367–368, 401, 414–415, 453–455, 489) [Long-winged Haaniella] Haaniella erringtoniae, Dräger, 2011: 41, fig. 16 (♂). HT, ♂: Singapore, Nee Soon swamp forest, leg. F. Seow-Choen, XII.2013[LKCNHM]. PT, ♂: Singapore, Nee Soon swamp forest, leg. F. Seow-Choen & Chua EK, X.2013[LKCNHM]. PT, ♀: Singapore, Nee Soon swamp forest, leg. F. Seow-Choen V.2013[LKCNHM]. PT, ♀: Singapore, Freshwater Forest, X.2013, F. Seow-Choen [NHMUK]. PT, ♂: Singapore, Nee Soon swamp forest, leg. Seow-Choen, Choo ME & CHUA EK, XI.2014 [coll. FSC]. PT, ♀: Singapore, Nee Soon swamp forest, leg. F. Seow-Choen & EK Chua, XII.2014 [coll. FSC]. PT, ♀: Singapore, Nee Soon swamp forest, leg. F. Seow-Choen & EK Chua, Choo ME, VIII.2014 [coll. FSC]. PT, ♀: Singapore, Nee Soon swamp forest, leg. EK Chua & Mohamed Azlin Sani, VII.2013 [coll. FSC]. PT, 3 ♂♂: Singapore, bred by F. Seow-Choen, from eggs of female, Nee Soon Swamp Forest, VIII.2015 [coll. FSC]. PT, ♀: Singapore, Nee Soon swamp forest, leg. F. Seow-Choen V.2013 [coll. FH, No’s 0869-1]. PT, ♂: Singapore, Nee Soon swamp forest, leg. F. Seow-Choen VII.2013 [coll. FH, No. 0869-2]. PT, 3 eggs: Singapore, Nee Soon swamp forest, leg. F. Seow-Choen V.2013 [coll. FH, No’s 0869-E].

Diagnosis: Closely related and possibly the sister-taxon of H. erringtoniae (Redtenbacher, 1906). Males are very similar to H. mecheli (Redtenbacher, 1906), but differ by the larger size, somewhat shorter wings (Fig. 173), more pronounced body aramture, different spination of the mesothorax and metatibiae, short epiproct, narrowed and roundly angular posterior margin of the poculum, shape of the vomer (Fig. 401) and dorso-apical tooth of the cerci



51

(Fig. 178). From H. erringtoniae ♂♂ of this new species readily differ by the elongated tegmina and fully developed alae, which reach as far back as to abdominal segment VII, the short epiproct and the distinct subapical dorsal tooth of the cerci. Females can be distinguished from those of H. erringtoniae by the averaging larger size, having the tegmina and alae averaging longer and roughly reaching to the posterior margin of abdominal tergum III (at best slightly projecting over tergum II in erringtoniae), larger lateral spines of abdominal tergites II–VI, more angular outer angles of the posterior margin of the anal segment (Fig. 176), notched apex of the subgenital plate (Fig. 177) and longer more decidedly curved ventral spines of the metatibiae (Fig. 367). The distinctive eggs differ from those of H. erringtoniae by the averaging larger dimensions, more distinctly granulose capsule, differently shaped micropylar plate and flattened, medially impressed apex of the operculum. The latter feature is unique within Heteropteryginae. Etymology: The name refers to the well-developed alae and elongate tegmina of ♂♂ of this new species. Description: ♀♀ (Figs. 171–172). Large (body length 107.0–126.0 mm) and fairly slender with moderately long and slender legs; largest member of the muelleri species-group. Colour variable, ranging from pale to mid brown and sometimes reddish brown, usually irregularly marbled and speckled with dark brown. Cephalic and thoracic spines green at the base, then pale red and with the tips black. Ventral surface of body usually of paler colour than dorsal surface. Tegmina ranging from plain pale to dark brown and sometimes with sparse pale mottling; anterior margin often with a distinct white spot near the base. Occasionally specimens may have a broad whitish median transverse band across the tegmina, a large basal white marking (Fig. 455) and/or have the anterior margin bright white. Head slightly longer than wide, globose with vertex roundly convex and armed with ten spines (Fig. 181). Vertex with four distinct coronals which roughly form a square; the anterior coronals larger than the two posterior ones. Anteriorly with a further pair of small supra-occipitals (Fig. 181). Posterior margin with two fairly distinct lateral coronals. Behind the eyes with a fairly distinct supra-orbital spine, about equal in size to the lateral coronals. Antennae ± reaching posterior margin of abdominal tergum IV; with 24 antennomeres. Pronotum longer than wide with lateral margins distinctly concave and posterior margin wider than anterior margin; antero-lateral pronotals and postero-lateral pronotals distinct and just before transverse median sulcus with a strong pair of pre-median pronotals as well as two minute inter-posterior pronotals near posterior margin (Fig. 181). Mesothorax gently and gradually widening towards the posterior with posterior portion some 1.8x wider than anterior margin. Mesonotum slightly but gradually narrowing towards posterior about 1.8x longer than wide and about 1.9x the length of pronotum. Lateral margins with 3–4 fairly distinct and slender spines as well as a few much smaller intercalated spines. Disc with a pair of ± distinct anterior-mesonotals and a strong pair of pre-median mesonotals close to median line; between these a variable number of smaller spines. Usually a small pair of fairly distant, small postmedian mesontals present and otherwise with a few minute irregularly dispersed spiniform tubercles (Fig. 181). Between bases of tegmina with two strong posteromedial mesonotals on a common base. Tegminal spine fairly distinct and slender. Mesopleurae with a long and slender supra-coxal spine and 4–5 almost equally sized, slender laterals along lower margin; often with a few much smaller intercalated spines. Metapleurae with lateral margin gently rounded and armed with five long and slender spines in the anterior 2/3 and a much smaller spine shortly before metacoxae; supra-coxal prominent. Sensory area of profurcasternum moderately distinct and roughly triangular in outline (Fig. 80). Mesosternum with a lateral row of three stout spines, which decrease in size towards the posterior; otherwise with a few irregularly dispersed spiniform tubercles in the lateral portions. Metasternum with a variable number of minute spinules and a fairly pronounced lateral spine. Tegmina large, elongate-oval and ± reaching half way along abdominal tergum III. Alae translucent grey with black veins. All abdominal tergites smooth dorsally. Lateral margins of II–VI very gently deflexed and with 3–4 fairly small spines; lateral margins of VII smooth. Sternites II–VI with four spines on each side of the median line; these most prominent on sternum II and decreasing in size towards VI. Praeopercular organ on sternum VII formed by a small black median swelling some distance off the posterior margin, which is accompanied by two small, back-curving spines (Fig. 177). Posterior margin of anal segment roundly rectangular and very indistinctly indented medially (Fig. 176). Epiproct tectiform longitudinally, gradually narrowing and gently downcurving with the apex deeply notched and forming two acute points (Figs. 176–177). Subgenital plate with dorsal margins roundly excavated medially (Fig. 175); the apex roundly angular, weakly notched and not reaching apex of epiproct (Fig. 177). All legs long and slender, mesofemora about equal in length to mesonotum, metafemora reaching about half way along abdominal segment VI and metatibiae rougly reaching to apex of abdomen. All carinae sparsely dentate and metatibiae with prominent,


HENNEMANN ET AL.

but slender and slightly curved teeth on the two outer ventral carinae, which increase in size towards the apex of tibia (Fig. 367). Ventral surface of all femora smooth. ♂♂ (Figs. 173–174). Large (body length 83.0–84.0 mm) and fairly slender insects with long and moderately slender legs, elongated tegmina and fully developed alae. General colour dull ochre to mid brown and often with a greenish hue. Head, pronotum and mesonotum dark green, the latter with a pale ochre stripe along lateral margins. Tegmina plain mid to dark brown and to a variable degree furnished with pale washed pale markings and spots; the basal 3/4 of the anterior margin either bright apple green (Figs. 174, 453) or white over entire length (Fig. 454). Femora mid to dark brown (often with a slight greenish hue), the tibiae ranging from very pale green to dull ochre. Armature of legs mostly black; teeth on ventral carinae of metatibiae in particular. Shape and armature of head as in ♀♀, but armature more prominent with all spines comparatively much longer and more slender. Antennae very long and almost reaching to apex of abdomen; with 22 antennomeres. Pronotum about 1.7x longer than wide, somewhat longer than head, with the lateral margins gently concave; spines as in ♀♀ but decidedly longer, the pre-medials in particular very prominent and long. Mesothorax moderately slender and elongate, some 1.7x longer than pronotum with anterior and posterior portions roughly equal in width and the mid portion very gently constricted in dorsal view. Mesonotum gently narrowed medially and almost 3x longer than wide; disc unarmed except for a prominent pair of antero-lateral mesonotals and a very strong pair of pre-median mesonotals; sometimes with a minute pair of post-median spines and with two or three fairly stout spines on lateral margins. The two posteromedial mesonotals between the bases of tegmina very long, slender and somewhat larger than the pre-medials. Tegminal moderately distinct. Mesopleurae with a very long and slender supra-coxal spine and 3–4 strong spines along lower margin; also with a variable number of much smaller intercalated spines close to lower margin. Armature of metapleurae as in ♀♀ but all spines comparatively longer. Sensory area of profurcasternum as in ♀♀ and noticeably raised. Mesosternum smooth except for three stout black lateral spines, which increase in size towards the posterior; a pair of minute posterior mesosternals between coxae. Metasternum with three short, black spines and a much more pronounced lateral spine on each side. Tegmina elongate, slender, about 3/5 the length of alae and ± reaching half way along abdominal segment IV. Alae reaching to abdominal segment VII, the anal fan pale and transulcent pink with dark grey veins. Abdominal segments II–VII averaging 1.8x longer than wide, IV–VII gradually decreasing in length and II somewhat widening towards anterior. Tergites II–VII smooth dorsally; IV–VII with the lateral margins moderately rounded and deflexed in posterior half. II–VI with 3–4 fairly long but very slender, black lateral spines, which are restricted to the deflexed posterior portion in VI. VII only with three very small spines in posterior portion (Fig. 180). Sternites II–VI with eight fairly slender, black spines, VII unarmed. Anal segment somewhat narrowed towards posterior with posterior margin deeply notched medially and the outer angles broadly rounded (Fig. 179). Epiproct very short and with a shallow median notch apically. Cerci club-like and compressed dorsoventerally with the upper apical portion expanded to form a roundly triangular projection (Fig. 178). Vomer broadly triangular with the apical hook somewhat curved dextrally; dull reddish brown (Fig. 401). Poculum with posterior margin broadly angular and labiate (Fig. 180). Legs all very moderately long and slender, metafemora reaching almost half way along abdominal segment VI and metatibiae projecting noticeably beyond apex of abdomen. All carinae more or less distinctly dentate, the teeth fairly slender and pointed Metatibiae with the two dorsal carinae set with numerous small, black teeth, the two outer ventral carinae with about five strong triangular black teeth and several further, much smaller teeth particulatly in the basal portion (Fig. 368). Ventral surface of all femora smooth. Variability: Both sexes show variability in the colouration and slight variation concerning to the size and position of the mesothoracic spines. The most significant variability is, that ♂♂ usually have the anterior margin of the tegmina bright apple green (Fig. 453) but it may be distinctly white in some specimens (Fig. 454). No intermediate forms are known and such a situation is unusual in Haaniella since it has not yet been observed in any other species. Variability in the colouration of the tegmina is also seen in ♀♀, some specimens having a broad whitish transverse median band or basal marking (Fig. 455) or a white stripe along anterior margin. Eggs (Figs. 414–415): Large, sub-circular in cross-section and distinctly lemon-shaped with capsule strongly narrowed in anterior portion and polar-area strongly protuded and roundly conical; about 1.5x longer than wide. Surface of capsule granulose and irregularly set with numerous minute but acutely pointed, slender black tubercles. Micropylar plate principally X-shaped, with all four extension moderately slender; anterior extensions gently broadened in apical half, posterior extensions strongly upcurving and broadly rounded apically. Interior portion of micropylar plate gently raised, the outer margin also raised and entire plate surrounded by a blunt bulge; surface as



53

in capsule. Micropylar cup a small and flat black knop, at both sides accompanied by a short and rounded carina. Operculum almost circular, marginated and conically raised medially; the median extension with the top flattened and a central impression. General colouration greyish mid brown, the anterior margin of the capsule, operculum as well as the outer margin and bulge surrounding the plate very dark grey to black. Measurements [mm]: length incl. operculum 9.1–10.4, length 7.8–8.9, width 4.9–5.2, height 5.6–6.1. Comments: All records of H. macroptera n. sp. restrict to swamp forests or low-land forests. Natural foodplants include Rubus moluccanus (Rosaceae) and Dillenia suffruticosa (Dilleniaceae). Captive rearing in Europe is currently being attempted and bramble (Rubus fruticosus, Rosaceae) appears to be accepted as alternative foodplants (personal communication with Ian Abercrombie, Kent, U.K.). Distribution (Fig. 489): Singapore: Nee Soon swamp forest [LKCNHM, coll. FH, coll. FSC]; Freshwater Forest [NHMUK]; South Peninsular Malaysia: Johor, Gunung Kulai [photos by the fourth author]; Johor, Endau Rompin [photos by the fourth author]. Specimens examined: 13 TABLE 8. Measurements [mm] of Haaniella macroptera n. sp. ♂, HT [LKCNHM]

♂♂, PT

♀♀, PT

Body

83.0

83.0–84.0

107.0–126.0

Head

6.0

6.0–7.0

10.0–11.0

Pronotum

7.0

7.0–7.5

9.0–10.0

Mesonotum

11.0

12.0

17.0–20.0

Metanotum

5.0

5.0

6.0–7.5

Median Segment

4.5

4.0

7.0–8.0

Tegmina

29.5

29.0–31.0

29.0–34.0

Alae

47.0

45.0–46.0

25.0–26.0

Profemora

20.0

19.0–20.0

23.0–25.0

Protibiae

22.0

20.0–23.0

22.0–27.0

Mesofemora

17.0

17.0–18.0

19.0–22.0

Mesotibiae

17.0

17.0–18.0

20.0–23.0

Metafemora

24.5

25.0–26.0

29.0–33.0

Metatibia

27.0

25.0–27.0

33.0–40.0

Antennae

71.0

72.0–74.0

75.0–80.0

Haaniella mecheli (Redtenbacher, 1906) n. comb., rev. stat. (Figs. 182–185, 400, 490) [Mechel’s Haaniella] Leocrates mecheli Redtenbacher, 1906: 167. HT, ♂: Sumatra, Indragiri, leg. Mechel 1899 (VI.D.46), [34] [NHMB]. Hennemann & Conle, 1999: 11. [Erroneously listed as a synonym of Haaniella muelleri (Haan, 1842)] Haaniella muelleri, Otte & Brock, 2005: 149 (in part).

Diagnosis: Very similar and very closely to H. macroptera n. sp. from Singapore and southern Peninsular Malaysia, but ♂♂ (the only sex known) differ by the smaller size, brown general colouration, somewhat longer wings (Fig. 182), less pronounced body aramture, spination of the mesothorax and metatibiae, elongated epiproct, posteriorly rounded poculum and shape of the vomer (Fig. 400). The true relation to H. macroptera n. sp. can however not be defined without knowledge of the still unknown ♀♀. Also similar to the macropterous ♂♂ of H. glaber (Redtenbacher, 1906) but differing by the somwhat larger size and more stocky shape, broader and less


HENNEMANN ET AL.

elongated mesothorax, more prominent spines of the mesonotum, simple anterior coronals, shorter poculum, longer and more slender epiproct (Fig. 185), dorsally dentate tibiae and different armature of the metatibiae. Etymology: Named after Anton v. Mechel, who collected the holotype of this species amongst several other Phasmatodea described by Redtenbacher (1906–1908). Description ♂♂ (Fig. 182). The unique holotype lacks most of both antennae and has both protibiae broken. Due to the old age of the specimen the colour and in particular that of the anal fan of the alae is badly faded. Medium-sized (body length 74.1 mm) with long wings. General colour dull ochre brown, the lateral margins of the mesonotum with a fine dull ochre line. Armature of head and body dark orange with black tips, that of the legs mostly black. Tegmina greyish mid brown with the anterior margin pale cream. Anal fan of the alae very pale transparent orange with reddish brown longitudinal and dark brown transverse veins. Head globose, hardly longer than wide with the vertex roundly convex and armed with ten fairly long and slender spines. Vertex with four prominent coronals which roughly form the four corners of a square; the anterior coronals somewhat larger than the two posterior ones. Anteriorly with a further pair of small supra-occipitals. Posterior margin with two distinct lateral coronals and behind the eyes with a fairly long supra-orbital spine. Pronotum slightly longer than head and about 1.3x longer than wide, narrowed medially with lateral margins distinctly concave and the posterior margin wider than anterior margin. Antero-lateral pronotals and postero-lateral pronotals small and just before transverse median sulcus with a prominent pair of pre-median pronotals; a small pair of inter-posterior pronotals present. Mesothorax slender, elongate, some 1.6x longer than pronotum with posterior portion distinctly widened and the mid portion very gently constricted in dorsal view. Mesonotum gently narrowed medially and about 3.4x longer than wide. Disc armed with six spines; a pair of short anterior mesonotals, a moderately distinct pair of pre-median mesonotals and a decidedly shorter pair of post-median mesonotals. Between the bases of the tegmina with two stout and fairly short posteromedial mesonotals on a common base. Lateral margins with three stout spines. Tegminal spine small. Mesopleurae with a long and strong supra-coxal spine and about five fairly short but distinct, unequally sized laterals along lower margin; the most posterior spine much shorter than all previous. Metapleurae widened in the posterior portion and with five strong, somewhat unequally sized spines; the supra-coxal very prominent Mesosternum smooth except for three short but robust, black lateral spines, which increase in size towards the posterior. Metasternum set with a few small spines and a fairly prominent and strong, black spine near outer lateral margin. Tegmina very elongate, slender, roughly 2/ 3 the length of alae and reaching about half way along abdominal tergum V (Fig. 182); radial vein strongly raised in the basal portion of tegmen. Alae fully developed and almost reaching to posterior margin of abdominal tergum IX; near base with a large tympanal area. Abdominal segments III–VII very gently narrowing towards posterior and averaging 1.2x longer than wide; V–VII decreasing in length. Lateral margins of V–VII roundly deflexed in posterior half; in II–VI with four short but slender spines, VII only with two small spines in the deflexed posterior portion. Sternites II–VI with eight black spines; one pair pre-medially, one pair posteriorly and two much smaller spines near each lateral margin. Sternum VII unarmed. Anal segment with basal two thirds narrowing, then widening, the posterior margin distinctly indented medially and the outer angles broadly rounded (Fig. 184). Epiproct elongate, considerably longer than wide, slightly narrowed towards the posterior and with the apex notched medially; projecting distinctly beyond posterior margin of anal segment (Fig. 185). Cerci very short, compresseed basally and with the apical portion distinctly club-like (Fig. 400). Vomer roughly triangular with the outer margins strongly swollen and the median portion distinctly impressed, slightly curved dextrally; colour dull drab (Fig. 400). Posterior margin of poculum broadly rounded and with a very shallow median indention; distinctly projecting over posterior margin of tergum IX (Fig. 183). Legs moderately long and slender with all carinae (except the dorsal carinae of the pro- and mesotibiae) acutely dentate; metafemora reaching about half way along abdominal segment VI and metatibiae projecting considerably beyond apex of abdomen. Two outer ventral carinae of metatibiae armed with about five prominent, elongate and straight triangular teeth as well as a few much smaller intercalated teeth. Ventral surface of all femora smooth. Comments: This species is so far only known from the unique holotype in NHMB, hence no information is available on the natural habits, host-plants or variability. Females and eggs unknown. Hennemann & Conle (1999: 11) have mistaken H. mecheli as a macropterous variety of H. muelleri (Haan, 1842) and hence synonymised it with this species. Subsequent examination and careful comparison with H. muelleri and the other species of the genus that have macropterous ♂♂, namely H. aculeata n. sp., H. glaber (Redtenbacher, 1906) and H. macroptera n. sp., has shown H. mecheli to represent a distinct and valid species. Consequently, it is here re-established (rev. stat.). REVISION OF HETEROPTERYGINAE


55

Distribution (Fig. 490): So far only known from the type-locality. E-Sumatra: Prov. Riau, Indragiri Hulu, Indragiri River [NHMB]. Specimens examined: 1 TABLE 9. Measurements [mm] of Haaniella mecheli (Redtenbacher, 1906) ♂ HT [NHMB] Body

74.1

Head

7.2

Pronotum

6.9

Mesonotum

12.5

Metanotum

-

Median Segment

-

Tegmina

30.7

Alae

45.8

Profemora

18.7

Protibiae

> 14.9*

Mesofemora

14.9

Mesotibiae

14.0

Metafemora

21.8

Metatibia

25.6

Antennae

-

* apex broken

Haaniella muelleri (Haan, 1842) (Figs. 186–196, 492) [Müller’s Haaniella] Phasma (Heteropteryx) mülleri Haan, 1842: 108, pl. 11: 4 (♂) & 5 (♀). LT, ♂: Müller Padang; Haaniella mulleri (deHaan) Det. S. Nakata 1965; Lectotype des. Brock 1995: 95; RMNH Leiden Lectotype Heteropteryx muelleri de Haan 1842 [RMNH]; PLT, 1 ♂, 1 ♀, 3 nymphs: RMNH Leiden Paralectotype Heteropteryx muelleri de Haan 1842 [RMNH]. [Not: PLT, 2 ♀♀: H. muelleri var. b., RMNH Leiden Paralectotype Heteropteryx muelleri de Haan 1842 [RMNH]. These are H. rosenbergii (Kaup, 1871) → see below). Heteropteryx Mülleri, Westwood, 1859: 82. Heteropteryx mülleri, Redtenbacher, 1906: 172. Dohrn, 1910: 402. Haaniella mülleri, Kirby 1904b: 397. Günther, 1932: 313. Brock, 1995: 95 (in part). [Designation of lectotype] Haaniella muelleri, Zompro, 2004: 197, figs. 112a & b (♂, ♀). [♂ only, ♀ is H. rosenbergii (Kaup, 1871)] Otte & Brock, 2005: 149 (in part). Haaniella mülleri mülleri, Günther, 1944: 72. Bragg, 1996: 112. [Not: Haaniella mülleri, Schulten, 1995: 98; misidentification relating to H. erringtoniae (Redtenbacher, 1906)] [Not: Haaniella muelleri, Brock, 1999: 141 & 183; misidentification relating to H. erringtoniae (Redtenbacher, 1906)] [Not: Haaniella muelleri, Seow-Choen, 2000a: 151; misidentification relating to H. erringtoniae (Redtenbacher, 1906)] [Not: Haaniella muelleri, Seow-Choen, 2000b: 40; misidentification relating to H. erringtoniae (Redtenbacher, 1906)] [Not: Haaniella muelleri ssp., Zompro, 2000b: 64; misidentification relating to H. rosenbergii (Kaup, 1871)] [Not: Haaniella (Heteropteryx) erringtoniae novaeguineae Günther, 1930: 737, figs. 4 (♂) & 5 (♀); erroneous synonym by Otte & Brock, 2005. 149] [Not: Leocrates glaber Redtenbacher 1906: 167; erroneous synonym by Brock, 1995: 96]


HENNEMANN ET AL.

[Not: Leocrates mecheli Redtenbacher, 1906: 167; erroneous synonym by Hennemann & Conle, 1999: 11].

Material examined (8 ♂♂, 3 ♀♀, 11 nymphs): SUMATRA: 1 ♂: Sumatra, Mt. Singalang, Luglio 1887, O. Beccari [MCSN]; 1 ♀: C. J. Louwerens, Sumatra, Benkoelen, Lebong Tanday, 1927 [MNHN]; 1 ♂, 1 ♀ (nymph): Banka, N.W. vd. Wileck, det. K. Günther [RMNH]; 1 ♀ (nymph): E. Jacobson, Félémoun, Sumatra, V. Jan. 1917, det. Bragg [RMNH]; 2 ♂♂, 2 ♀♀ (nymphs): Indrapoera, Sumatra, Weyers [RBINS]; 1 ♂: Sumatra, Lebong Tandai, 9.–14.XI.1922, C.J. Brooks [NHMUK, 1936-681]; 1 ♀: Sumatra, Montes Battak, ex coll. Fruhstorfer; H. Fruhstorfer vend. 25.V.1904.; K. Günther determ. Vers. Nr. 131, 1932; Heteropteryx mülleri de Haan 1842, K. Günther det. [ZMUH]; 1 ♂ (penultimate instar): Sumatra, Deli, ex coll. Fruhstorfer; K. Günther determ. Vers. Nr. 131, 1932; Haaniella mülleri rosenbergi Redt., K. Günther det. [ZMUH]; 1 ♀, 2 ♀♀ (nymphs): Mus. Caes. Vind. Padang, Consul Schild; det. Holdhaus, Heteropteryx mülleri [NHMW, No. 287]; 1 ♂ (penultimate instar nymph): Badang, 96, Fcons. Schild, Mus. Caes. Vindobon, det. Redtenb. Heteropteryx mülleri, Heteropteryx mülleri de Haan larva [NHMW, No. 287]. NO DATA: 1 ♂: coll Karny [NHMW, No. 287]. Diagnosis: Closely related and most certainly the sister-taxon of H. rosenbergii (Kaup, 1871). Both sexes can be separated from H. rosenbergii by the more slender appearance and relatively longer legs, having six posteromedial mesonotals on a strongly raised and swollen common base between the bases of the tegmina (only 2–4 in rosenbergii), and having a distinct pair of supra-antennals. Males readily differ from those of H. rosenbergii by the colouration of the tegmina, which have the anterior margin broadly green (with a conspicuous white lateral sub-apical marking in rosenbergii), longer and more slender mesonotum, which is 3x longer than the pronotum (only 2.5x longer in rosenbergii) and obsolete anterior mesonotals. Females differ by having the leg armature more prominent with the dorsal carinae of the tibiae armed with several acute teeth and having the lateral margins of abdominal tergites III–VI more distinctly expanded with the spines considerably longer. Males strongly resemble those of H. erringtoniae (Redtenbacher, 1906) from Peninsular Malaysia, with which they share the broadly green anterior margin of the tegmina and conspicuously elongated epiproct. They however clearly differ from this species by the smaller size, obsolete anterior mesonotals, lack of post-median mesonotals, six prominent posteromedial mesonotals and relatively longer, more slender mesothorax. Etymology: Named after its collector Salomon Müller (Heidelberg, Germany), who collected numerous insects and other animals throughout Indonesia now held in the collections of RMNH. Description: Haan (1842, pl. 11: 4 & 5) provided beautiful coloured figures of both sexes. There have been numerous descriptions of this species but all relate to H. erringtoniae from Peninsular Malaysia. The colouration is described from preserved specimens only, since no live specimens or photos have been available. ♀♀ (Figs. 186–187). Fairly large (body length 84.0–96.5 mm) with long and slender legs and between the bases of the tegmina with six posteromedial mesonotals on a swollen common base (Fig. 190). Colour pale to dark brown or dull drab. Body armature dull orange to red with black points. Tegmina plain mid to dark brown with the anterior margin occasionally greenish in basal portion. Alae translucent greyish brown with black veins. Head slightly longer than wide, globose with vertex roundly convex and armed with twelve spines (Fig. 191). Vertex with four strong coronals which roughly form the four corners of a square; the anterior coronals a little longer than the two posterior ones. Anteriorly with a further pair of somewhat smaller supra-occipitals (Fig. 190). Supra-antennals distinct. Posterior margin with two fairly distinct lateral coronals. Behind the eyes with a fairly long supra-orbital spine, about equal in size to the posterior pair of coronals. Antennae ± reaching posterior margin of abdominal tergum IV. Pronotum slightly longer than wide with lateral margins distinctly concave and posterior margin somewhat wider than anterior margin; antero-lateral pronotals very minute, postero-lateral pronotals distinct and just before transverse median sulcus with a strong pair of pre-median pronotals as well as two small, distant inter-posterior pronotals near posterior margin (Fig. 191). Mesothorax gently and gradually widening towards the posterior with posterior portion almost 2x wider than anterior margin. Mesonotum roughly parallelsided, about twice as long as wide and about 2x the length of pronotum. Lateral margins with 2–3 fairly small and short spines, disc with a pair of fairly small anterior-mesonotals, a very strong pair of pre-median mesonotals close to median line and a further pair of smaller and more distant post-median mesonotals (Fig. 191). Between bases of tegmina with a very prominent and characteristic swelling, that bears six strong but fairly short posteromedial



57

mesonotals (Fig. 190). Tegminal small. Mesopleurae with a long and slender supra-coxal and 4–5 prominent but unequally sized laterals along lower margin (Fig. 191). Metapleurae with lateral margin strongly rounded and armed with five prominent spines, the three median ones decidedly larger than the other two; supra-coxal distinct. Sensory area of profurcasternum weakly developed. Mesosternum with a lateral row of three small spines and often some further small spines in the median portion. Metasternum with 8–10 small spines, the outermost spine more pronounced than the others. Tegmina oval and ± reaching half way along abdominal tergum III. All abdominal tergites smooth dorsally except for a pair of minute spines just behind the middle. Lateral margins of II–III and VI– VII slightly, those of IV and V strongly deflexed and almost semicircularly rounded, all armed with four fairly strong spines; these very small on VII. Sternites II–VI with four spines on each side of the median line; these most prominent on sternum II and decreasing in size towards VI. Praeopercular organ on sternum VII formed by two blunt spine-like swellings (Fig. 194). Posterior margin of anal segment gently concave. Epiproct tectiform longitudinally, slightly downcurving with the apex deeply incised and forming two elongate, slender and acute points (Figs. 193–194). Subgenital plate with dorsal margins gently concave medially (Fig. 192) and the apex angular with a ± decided median notch; not reaching apex of epiproct (Fig. 194). All legs long and slender, metafemora reaching to posterior margin of abdominal segment VI and metatibiae distinctly projecting beyond apex of abdomen. All carinae prominently and acutely dentate; metatibiae with distinct triangular teeth on the two outer ventral carinae. Ventral surface of all femora smooth. ♂♂ (Fig. 188). Medium sized (body length 60.0–73.0 mm) and slender for the genus with very long and slender legs and six posteromedial mesonotals on a swollen common base between the bases of the tegmina. General colour dull drab with a greenish hue on entire body except abdomen. Mesonotum with a ± distinct pale ochre stripe along lateral margins, mesopleurae dark brown. Ventral surface of body ochre, the mesosternum pale ochre to straw. Tegmina plain mid to dark brown with the anterior margin bright apple green. Alae translucent greyish brown with black veins. Femora dull green, tibiae ochre. Armature of legs mostly black. Shape and armature of head as in ♀♀, but armature more prominent with all spines much longer and more slender (Fig. 189). Pronotum about 1.4x longer than wide, about equal in length to head, with the lateral margins gently concave; spines as in ♀♀ but decidedly longer (posterior pronotals in particular). Mesothorax very slender and elongate, some 3x longer than pronotum with posterior portion widened and the mid portion very gently constricted in dorsal view (Fig. 189). Mesonotum almost parallel-sided and almost 5x longer than wide; disc with six spines arranged as in ♀♀, but the anterior mesonotals and post-medial mesonotals obsolete (Fig. 189). Posteromedian swelling between bases of tegmina more prominent than in ♀♀ with the six posteromedial mesonotals comparatively longer and more slender. Tegminal small. Mesopleurae with a long and slender supracoxal and 4–5 strong laterals along lower margin. Armature of metapleurae as in ♀♀. Sensory area of profurcasternum roughly triangular in outline, the segment furthermore with a minute spine near each posterolateral corner. Mesosternum smooth except for three small, black lateral spines and a pair of very minute posterior mesosternals between coxae. Metasternum as in ♀♀, but spines less developed. Tegmina ovate and slightly projecting over posterior margin of median segment. Abdominal segments II–VII averaging 1.6x longer than wide, II relatively shorter and widening towards anterior. Tergites II–VII smooth dorsally; IV–VII with the lateral margins moderately rounded and deflexed in posterior half. II–VI with four small and slender, unequally sized lateral spines, which are restricted to the deflexed posterior portion in IV–VI. Sternites II–VI with four slender and distinct black spines, which decrease in size from II towards VI; VII only with a single pair of minute posterior spines. Anal segment somewhat constricted post-medially with the posterior margin deeply excavated triangularly (Fig. 195). Epiproct elongated, longer than wide, notched apically and projecting over apex of abdomen (Fig. 195). Cerci small, club-like and compressed dorsolaterally. Vomer broadly triangular and longer than wide with the apical hook somewhat curved dextrally; dull reddish brown. Poculum with posterior margin broadly angular, labiate and very gently excavated medially (Fig. 196). Legs all very long and slender, mesofemora longer than mesonotum, metafemora reaching to abdominal segment VII and metatibiae projecting strongly beyond apex of abdomen. All carinae acutely dentate, the teeth fairly slender and pointed; two outer ventral carinae of metatibia with 6–7 prominent, triangular black teeth. Ventral surface of all femora smooth. Variability: Some variability is seen in the degree of body armature in both sexes. The ♀ in MNHN (Fig. 187) has the entire body and leg armature more pronounced than the paralectotype in RMNH or ♀ in ZMUH, with the abdominal tergites III–VI being considerably more deflexed. The latter ♀ has the anterior mesonotals and postmedian mesonotals very weakly developed (Figs. 190–191).


HENNEMANN ET AL.

Comments: Haan (1842: 108) originally described this species from several examples taken on Mount Singgalang (Padang District, W-Sumatra) and provided beautiful colour illustrations of both sexes. Examination of the two additional specimens from an undefined locality described by Haan as “Heteropteryx muelleri var. b.” have proven to represent fairly small and weakly armed specimens of H. rosenbergii (Kaup, 1871). There has been considerable confusion concerning the status of H. muelleri and related species, which has resulted in numerous erroneous synonyms. Several of these synonyms were caused by Günther’s attempt of an intrageneric arrangement of Haaniella Kirby, 1904 (Günther, 1944: 69, 72), which listed several of the described taxa as subspecies of H. muelleri. The following taxa have erroneously been listed as synonyms of H. muelleri (Brock, 1995; Brock, 1999; Hennemann & Conle, 1999; Otte & Brock, 2005) and are here re-established or the synonymies corrected: Heteropteryx glaber Redtenbacher, 1906 rev. stat., Haaniella erringtoniae (Redtenbacher, 1906) rev. stat., Haaniella erringtoniae novaeguineae Günther, 1930 (= Haaniella rosenbergii (Kaup, 1871) n. syn.), Haaniella muelleri simplex Günther, 1944 (= Heteropteryx glaber Redtenbacher, 1906 n. syn.), Leocrates mecheli Redtenbacher, 1906 rev. stat. and Haaniella rosenbergii (Kaup, 1871) rev. stat.. Several papers have dealt with the captive rearing of stock originating from Peninsular Malaysia (e.g. Schulten, 1995) and numerous authors recorded H. muelleri from Peninsular Malaysia (e.g. Brock, 1999; SeowChoen, 2000a; Seow-Choen, 2000b), but all these records are based on misidentifications and relate to H. erringtoniae (Redtenbacher, 1906). In fact, nothing is known about the natural habits and host-plants of H. muelleri. Haan (1842: 108) stated this species to live under dead and dehydrated leaves in the jungle, information most certainly forwarded by the collector of the type specimens. Eggs unknown. Distribution (Fig. 492): Sumatra, endemic. W-Sumatra: Prov. Sumatera Barat, Padang district, Batang Singgalang [RMNH—type locality]; W-Sumatra: Prov. Sumatera Barat, Padang district, Mount Singgalang [MCSN]; W-Sumatra: Prov. Sumatera Barat, Indrapura [RBINS]; W-Sumatra: Prov. Sumatera Utara, Tapanuli district, Kapanoek River “Papanoeli” [RMNH]; SW-Sumatra: Prov. Benkulu, Lebong Tandai [MNHN, NHMUK]; NE-Sumatra, Prov. Sumatera Utara, Medan (= Deli) [ZMUH]; N-Sumatra: Prov. Sumatera Utara, Toba Highlands (= Batak Mountains) [ZMUH]; NW-Pulau Bangka, Prov. Bangka-Belitung [RMNH]. Specimens examined: 22 TABLE 10. Measurements [mm] of Haaniella muelleri (Haan, 1842) ♂ LT [RMNH]

♂ PLT [RMNH]

♀ PLT [RMNH]

♂ [MCSN]

♀ [ZMUH]

♀ [NHMW]

Body

60.0

60.5

84.0

73.0

96.5

90.0

Head

4.5

4.5

5.9

5.8

7.2

7.0

Pronotum

4.9

5.0

7.2

6.4

8.3

7.5

Mesonotum

13.1

12.7

15.8

14.0

16.2

16.0

Metanotum

3.5

3.2

4.9

-

-

-

Median Segment

3.4

3.8

6.2

-

-

-

Tegmina

10.0

9.2

19.2

12.7

18.4

22.0

Alae

6.8

6.9

15.1

-

-

-

Profemora

18.1

17.9

19.4

21.5

21.0

19.0

Protibiae

20.2

19.8

22.2

22.3

22.0

20.0

Mesofemora

15.1

13.9

16.4

16.0

17.5

15.0

Mesotibiae

15.9

16.6

18.3

16.5

18.8

13.0

Metafemora

21.5

-

25.1

25.5

25.6

25.0

Metatibia

24.2

-

29.6

27.0

31.0

29.0

Antennae

> 32.0

>35.0

> 36.0

75.0

> 19.0

> 23.0



59

Haaniella rosenbergii (Kaup, 1871) rev. stat. (Figs. 82, 197–213, 371–372, 398, 418–419, 456–457, 491) [Rosenberg’s Haaniella] Heteropteryx Rosenbergii Kaup, 1871a: 35, pl. 2: 2. HT, ♀: Java (?), durch Hermann v. Rosenberg [HLMD]. Kaup, 1871b: 20, pl. 1: 12. [Description of egg] Dohrn, 1910: 404. Haaniella Rosenbergi, Kirby, 1904b: 397. Günther, 1932: 313. Haaniella mülleri rosenbergi, Günther, 1944: 72. Haaniella (Heteropteryx) erringtoniae novae guineae Günther, 1930: 737, figs. 4 (♂) & 5 (♀). HT, ♀: Sattelberg, Deutsch Neu Guinea H. Rolle, Berlin W.; H. Rolle vend. 1.VII.1908; Typus; Haaniella Kirby (= Heteropteryx aut) K. Günther det.; H. e. novaeguineae n. subsp. K. Günther det.; Haaniella erringtoniae Redt. K. Günther det. [ZMUH]; PT, ♂: Sattelberg, Deutsch Neu Guinea H. Rolle, Berlin W.; H. Rolle vend. 1.VII.1908; Typus; H. e. novaeguineae n. subsp. K. Günther det.; Haaniella erringtoniae Redt. K. Günther det. [ZMUH]. (Erroneously synonymised with H. erringtoniae Redtenbacher, 1906 by Günther, 1935: 124 and with H. muelleri var. b de Haan, 1842: 1908 by Günther, 1936: 124—erroneous data) n. syn. Haaniella erringtoniae, Günther, 1935: 124. Phasma (Heteropteryx) mülleri var. b, Haan, 1842: 108. PLT (of muelleri), 2 ♀♀: H. muelleri var. b., RMNH Leiden Paralectotype Heteropteryx muelleri de Haan 1842 [RMNH]. n. syn. Haaniella mülleri var. b., Günther, 1936: 124. Günther, 1944: 72. Haaniella muelleri ssp., Zompro, 2000b: 64. [Not: Haaniella muelleri, Brock, 1995: 95; erroneous synonym] [Not: Haaniella muelleri, Brock, 1999: 95; erroneous synonym] [Not: Haaniella muelleri, Otte & Brock, 2005: 149; erroneous synonym]

Material examined (15 ♂♂, 25 ♀♀, 3 nymphs): SUMATRA: 2 ♀♀, 2 ♂♂: Sumatra, Excell. V. Studt. G. [MNHU]; 1 ♀ (nymph): Padang-Pandjang, West-Sumatra, H. Rolle, Berlin W., det. Ebner Heteropteryx dehaani ?, coll. R. Ebner [NHMW, No. 287]; 1 ♂, 1 ♀: Padang-Pandjang, West-Sumatra, H. Rolle, Berlin W. [NHMW, No. 281]; 1 ♀: Sumatra, Montes Battak, ex coll. Fruhstorfer [MHNG]; 1 ♀: Niederländisch Indien” [HNMM]; 1 ♂: Sumatra, Montes Battak, ex coll. Fruhstorfer; Collectio Br. v. W.; det. Redtenb. Heteropteryx mülleri; 25.304; Heteropteryx mülleri de Haan [NHMW, No. 287]; 1 ♂: Sumatra, Padang, 80. [ZMUH]; 1 ♂: Sumatra, 79–55 [NHMUK]; 3 ♀♀, 1 egg (ex ovipositor): W-Sumatra, Padang, II.1995 [coll. FH, No’s 0113-1 to 3 & E]; 1 ♂, 1 ♀: NW-Sumatra, Prov. Aceh, Subulussalam, leg. F. Seow-Choen IV.2013 [coll. FH, No’s 0113-4 to 5]; 2 ♂♂, 3 ♀♀: NW-Sumatra, Prov. Aceh, Subulussalam, leg. F. Seow-Choen IV.2013 [coll. FSC]; 1 ♂: Sumatra, ex. Mus. Berlin (MNHU) [coll. OC, No. 0032-1]; 2 ♀♀: WSumatra, Prov. Sumatera Barat, Padang Distr., Mt. Singgalang, 600 m, Annai Valley N.R., 8.2009, leg. St. Lakl [coll. OC, No's 0032-2 & 3]; 2 ♂♂: W-Sumatra, Prov. Sumatera Barat, Padang Distr., Annai Valley, 900–1000 m, SE slopes of Mt. Singgalang, 6.2007, leg. St. Jakl [coll. OC, No's 0032-5 & 5]; 1 ♂: W-Sumatra, Prov. Sumatera Barat, Padang Distr., Mt. Tandikat, 600–900 m, 8.2009, leg. St. Jakl [coll. OC, No 0032-6]. MENTAWAI ISLANDS (SIBERUT ID.): 2 ♂♂, 4 ♀♀, 1 ♂ (penultimate instar), 1 ♀ (penultimate instar): Mentawai Islands, Siberut Island prope Sumatra, VII.2007 [coll. OC, No's 0032-7 to 14]. PHILIPPINES [in error]: 1 ♀: Mt. Banahao, ex Coll. UMB [coll. OZ, No. 31-7]; 1 ♀: Mt. Banahao, O. Schütz [UMB]; 1 ♀: Philippinen [UMB]; 1 ♀ (nymph n4): Philippinen, O. Schütz [UMB]. Diagnosis: Closely related and very similar to H. muelleri (Haan, 1842), which is most certainly the sistertaxon. Both sexes of H. rosenbergii (Kaup, 1871) may however be distinguished by the somewhat stockier appearance and relatively shorter legs, distinct anterior mesonotals, having only 2–4 posteromedial mesonotal spines between the bases of the tegmina (six on a strongly swollen common base in muelleri) and lacking the distinct pair of supra-antennals seen in muelleri. Males readily differ by the colouration of the tegmina, which bear a conspicuous white lateral sub-apical marking (anterior margin broadly green in muelleri) and considerably


HENNEMANN ET AL.

shorter and broader mesonotum, which is only about 2.5x longer than the pronotum (3x longer in muelleri). Females have the leg armature less pronounced with the dorsal carinae of the tibiae only very minutely dentate and the lateral margins of abdominal tergites III–VI less expanded with the spines considerably smaller. Etymology: Dedicated to Hermann v. Rosenberg, who donated the holotype to HLMD (Kaup, 1871: 35). Description: ♀♀ (Figs. 197–199). Medium sized (body length 70.0–89.0 mm) species with moderately long but weakly dentate legs. Colour ranging from dark brown to almost black, either ± plain or with ochre markings and mottling on body and legs. Occasionally with a large but weakly defined V-shaped ochre marking on mesonotum. Body armature dull orange or brown. Tegmina plain dark brown to black, sometimes with some dull ochre speckles and rarely with a sub-basal black spot just over radial vein. Head longer than wide, globose with vertex roundly convex and armed with ten spines (Figs. 209–210). Vertex with four strong but fairly stout coronals which roughly form a square; the anterior coronals somewhat more pronounced than the two posterior ones. Anteriorly with a further pair of small supra-occipitals (Fig. 209). Posterior margin with two fairly distinct but stout lateral coronals. Behind the eyes with a supra-orbital spine, about equal in size to the posterior coronals. Antennae ± reaching to posterior margin of abdominal tergum IV; with 24 antennomeres. Pronotum indistinctly longer than wide, widening towards posterior and with the lateral margins gently concave. Antero-lateral pronotals distinct and often with a pair of minute anterior mesal pronotals, posterolateral pronotals distinct and usually with a somewhat smaller pair of inter-posterior pronotals ± equal in size to the anterior-mesal pronotals. Just before transverse median sulcus with a very strong pair of long pre-median pronotals (Fig. 209). Mesothorax gently and gradually widening towards the posterior with posterior portion about 1.7x wider than anterior margin. Mesonotum very slightly narrowing towards posterior, about 1.8x longer than wide and roughly 2x the length of pronotum. Lateral margins with 2–3 fairly distinct and slender spines as well as a similar number of much smaller spines inbetween. Disc with a pair of very distinct anterior-mesonotals, a very strong pair of long pre-median mesonotals close to median line and a further pair of somewhat smaller and more distant pair of post-median mesonotals (Fig. 210); usually some further minute scattered spines are present near the lateral margins. Between bases of tegmina with two (rarely four) fairly prominent, posteriorly directed posteromedial mesonotals on a common base (Figs. 209–210). Tegminal very small and stout. Mesopleurae with a very long and slender supra-coxal and 6–7 unequally sized spines at lateral margin; the median two decidedly larger than the remaining (Fig. 210). Metapleurae with lateral margin strongly rounded and armed with five prominent, slender spines, the anterior one decidedly smaller than the other four spines in the deflexed portion; supra-coxal very prominent and elongate. Sensory area of profurcasternum expanding almost over entire width of segment (Fig. 82). Mesosternum with a lateral row of three short spines. Metasternum with 8–10 short spines, the outermost spine more pronounced than the others. Tegmina oval and ± reaching to posterior margin ofabdominal tergum II; the apex fairly acute. Alae translucent grey with black veins. All abdominal tergites smooth dorsally but III–V sometimes with some very minute spiniform tubercles near posterior margin. Lateral margins of III–VI very gently deflexed, II–VI armed with four small, slender spines; these very indistinct on VI. Sternites II–VI with four spines on each side of the median line; these most prominent on sternum II and decreasing in size towards VI. Praeopercular organ on sternum VII formed by two short and stout spines and a shallow rounded swelling near posterior margin (Fig. 205). Posterior margin of anal segment with a shallow rounded indention. Epiproct tectiform longitudinally, slightly downcurving with the apex triangularly incised and forming two acutely triangular points (Figs. 203–205). Apex of subgenital plate angular with a ± decided median notch; not reaching apex of epiproct (Fig. 205). All legs moderately long and slender, metafemora ± reaching one third along abdominal segment VI and metatibiae slightly projecting beyond apex of abdomen. All carinae acutely dentate; metatibiae with about eight fairly short but pointed and backward directed teeth on the two outer ventral carinae (Fig. 371). Ventral surface of all femora smooth. ♂♂ (Figs. 200–202). Fairly small to medium sized (body length 51.0–67.5 mm) and slender with long and moderately slender legs. Colour very variable (→ see comment on variability below), but usually mid to dark brown with the dorsal surface of the pro- and mersonotum dull green. Ventral surface of thorax ochre, the mesopleurae sometimes blackish along lower margin. Tegmina plain mid to dark brown with a variable white marking in apical half (→ see comments on variability below). Armature of body orange to dull ochre. Femora greenish mid brown to dull green, tibiae ochre to dark reddish brown. Armature of legs mostly black. Shape and armature of head as in ♀♀, but all spines much longer and more slender (Fig. 211). Antennae with 22 antennomeres and reaching to abdominal segment VII. Pronotum about 1.3x longer than wide, somewhat longer



61

than head with the lateral margins gently concave; spines as in ♀♀ but decidedly longer (postero-lateral pronotals in particular). Mesothorax very slender and fairly elongate, some 2.3–2.5x longer than pronotum with posterior portion widened and the mid portion gently constricted in dorsal view (Figs. 212–213). Mesonotum almost parallel-sided and 3.6–4.2x longer than wide; disc with six prominent spines arranged as in ♀♀, but the postmedian mesonotals relatively smaller (Fig. 211). Between the bases of the tegmina with two (sometimes four) very long and slender, posteriad directed posteromedial mesonotals on a common base (Figs. 211–213). Mesopleurae with a very long and slender supra-coxal spine and 4–5 variably sized laterals along lower margin, the posterior 2– 3 spines larger than the anterior ones (Figs. 212–213); occasionally a few smaller spines near the lower margin. Armature of metapleurae as in ♀♀ but spines considerably longer and more slender. Sensory area of profurcasternum indistinct. Mesosternum smooth except for a few small, lateral spines and a pair of minute posterior mesosternals between coxae. Metasternum as in ♀♀, but spines less developed and the outermost one relatively more developed. Tegmina ovate and ± reachingt to posterior margin of median segment. Abdominal segments II–IV slightly increasing, V–VII decreasing in length, II widening towards anterior, IV–V averaging 1.2– 1.3x longer than wide. Tergites II–VI with two pairs of minute spines in anterior half and a transverse row of six minute spines near posterior margin; the two median spines larger than the others and fairly pronounced on II–IV. Lateral margins of II–V with 4–5 small but acute spines, the lateral margins of VI and VII gently deflexed and rounded posteriorly. Sternites II–VI with four spines in anterior 2/3 and four spines near posterior margin, the two median ones of which are larger than the lateral ones; spines gradually decreasing in size from II towards VI. Sternum VII tectiform longitudinally and only with a pair of minute pre-median spines. Anal segment gradually narrowing towards posterior, the posterior margin with a shallow and wide triangular excavation (Fig. 207). Epiproct very short (Fig. 398). Cerci small, club-like and compressed dorsolaterally in basal portion. Vomer triangular, somewhat longer than wide with a distinct and deep median furrow and the apical hook almost straight; dull reddish brown (Fig. 398). Poculum with posterior margin broadly rounded and with a shallow median indention (Fig. 398). Legs all long and fairly slender, mesofemora almost as long as pro- and mesonotum combined, metafemora ± reaching to posterior margin of abdominal segment VI and metatibiae projecting considerably beyond apex of abdomen. All carinae acutely armed with pointed and fairly slender, spine-like teeth; teeth on two outer ventral carinae of metatibia black with orange tips and distinctly backward directed (Fig. 372). Ventral surface of all femora smooth. Variability: This species shows considerable variability concerning to the size, colouration and degree of the body and leg armature. The ♀ holotype in HLMD and the two type specimens of the synonymous H. erringtoniae novaeguineae in ZMUH have the spines of the body considerably longer and more pronounced than all other examined specimens. Also the number of the posteromedial mesonotals between the bases of the tegmina varies. While there are only two such spines in most of the specimens examined, there are four in the ♀ holotype and even five in the ♂ from Batak Mountains in NHMW. The two ♀♀ paralectotypes of Haan’s “H. muelleri var. b” in RMNH are considerably smaller than all other specimens (body lengths 70.0 and 77.0 mm) and also have the body armature less pronounced, with all spines of the body and legs being comparatively shorter than in the other ♀♀ at hand. This is also the case for the ♂ from Padang in ZMUH, which only measures 51.0 mm (Fig. 202). The ♂ allotype of H. erringtoniae novaeguineae (Fig. 201) and ♂ from “Padang-Pandjang” in NHMW are worth mentioning because they are somewhat more slender than the other examined ♂♂. Furthermore these two specimens have a comparatively longer mesothorax and longer tegmina that have most of the apical half whitish with a black network of veins. The latter feature is also true for the small ♂ from Padang in ZMUH (Fig. 202). While the colouration is fairly constant in ♀♀, almost all at hand being plain mid to very dark brown with also the tegmina entirely plain and without any distinctive colour pattern, ♂♂ show some variability. Only one of the ♀♀ in coll. FSC and one ♀ in coll. FH have some paler brown speckles on the body, a washed ochre marking on the metapleurae, a large faint pale brown marking on the mesonotum and all of the femora with a pale ochre median transverse band (Figs. 197–198). The ♂ from Batak Mountains in NHMW, ♂ from W-Sumatra in ZMUH and ♂♂ from Subulussalam in coll. FSC and coll. FH have the head, thorax and all femora dull green and the tegmina are very dark plain brown with only a small, almost clear white apical marking, the latter being reduced to a short and slender white stripe along the anterior margin in the specimens from Subulussalam (Fig. 200). Other ♂♂ are brown and at best with a slight greenish hue on the head, thorax and femora and the white marking of the tegmina has to a variable degree some of the veins marked with dark brown. Eggs (Figs. 418–419): Medium sized, almost circular in cross-section and lemon-shaped with polar-area


HENNEMANN ET AL.

conical; about 1.4x longer than wide. Capsule surface slightly glossy and very unevenly granulose and rugose. Anterior margin swollen. Micropylar plate generally X-shaped, marginated, the four extensions of moderate length and fairly slender; anterior extensions gently incurving, the posterior extensions much broader and short, almost straight. Interior portion of plate slightly raised and surface like capsule. Entire plate surrounded by a distinctly raised bulge. Micropylar cup a somewhat indented, rounded knob anteriorly marginated by a faint and slightly curved transverse carina. Operculum almost circular and distinctly conical. General colouration dark plain grey to almost black. Measurements [mm]: length incl. operculum 7.2, length 6.6, width 4.7, height 5.1. Comments: Kaup (1871a: 35) gave “Java” as the type-locality, but this is most certainly erroneous. Since all subsequent records of H. rosenbergii are from Sumatra, also the holotype most likely originates from that island. Kaup (1871b: 20, pl. 1: 12) provided a brief description and illustration of the egg, which was extracted from the abdomen of the holotype. Günther (1944: 72) treated H. rosenbergii as a subspecies of H. muelleri (Haan, 1842). Brock (1995: 96) misinterpreted Günther’s treatment and listed H. rosenbergii it as a synonym of H. muelleri as did all subsequent authors. Examination and comparison of Kaup’s ♀ holotype in HLMD with Haan’s type specimens of H. muelleri in RMNH as well as further material, including the previously unknown ♂♂ of H. rosenbergii, has shown the latter to represent a distinct species. Hence, H. rosenbergii is here re-established (rev. stat.). Although very closely related and rather similar, there are several morphological characters that clearly distinguish these two species (→ see differentiations above). The two ♀♀ paralectotypes described as “Phasma (Heteropteryx) mülleri var. b” by Haan (1842: 108) in RMNH are small specimens of H. rosenbergii which have the body armature less developed than typical specimens. Haaniella erringtoniae novae guineae (Günther, 1930) was described based on a ♀ and ♂ from “New Guinea, Sattelberg” in ZMUH. Subsequently, Günther (1935: 124) however recognized the locality was wrong and the specimens mis-labelled, hence synonymised his subspecies with H. erringtoniae (Redtenbacher, 1906). In a later paper however Günther (1936: 124) regarded it as a synonym of “H. muelleri var. b”. Brock (1995: 96; 1999: 183) and Otte & Brock (2005: 149) listed it as a synonym of H. muelleri (Haan, 1842). Examination of the type specimens has shown them to be conspecific with H. rosenbergii, hence Günther’s subspecies is here synonymised (n. syn.). The specimens in UMB recorded from Mount Banahao on the island of Mindanao in the Philippines by Zompro (2000b: 64) as “Haaniella muelleri ssp.” are H. rosenbergii (Kaup, 1871), the locality obviously being erroneous. The fourth author collected specimens near Subulussalam in the Aceh Province in northwestern Sumatra in April 2013. This species is very common locally along the rather deforested low land jungles along the sides of the main trunk road leading into the town of Subulussalam from the direction of Kabanjahe. It was the commonost phasmid found along that road during that particular collecting trip. Distribution (Fig. 491): Sumatra. NW-Sumatra: Prov. Aceh, Subulussalam [coll. FH, coll. FSC]; N-Sumatra: Prov. Sumatera Utara, Toba Highlands (= Batak Mountains) [MHNG, NHMW]; N-Sumatra: Prov. Sumatera Barat, Padang district, Padang [ZMUH, coll. FH]; W-Sumatra: Prov. Sumatera Barat, Mount Singgalang 600 m [coll. OC]; W-Sumatra: Prov. Sumatera Barat, Annai Valley 900–1000 m [coll. OC]; W-Sumatra: Prov. Sumatera Barat, Mount Tandikat 600–900 m [coll. OC]; W-Sumatra: Prov. Sumatera Barat, Padang Panjang [NHMW]; Sumatra [MNHU, NHMUK]. Mentawai Islands. Prov. Sumatera Barat, Siberut Island [coll. OC]. Specimens examined: 48 TABLE 11. Measurements [mm] of Haaniella rosenbergii (Kaup, 1871) ♀ HT [HLMD]

♂♂

♀♀

♂ [ZMUH]*

♀ [ZMUH]*

Body

51.0–67.5

82.0–89.0

67.0

86.5

Head

4.0–6.5

7.0–8.2

4.7

7.5

Pronotum

4.2–5.0

6.6–7.8

5.8

7.3

10.2–13.5

12.0–15.0

13.2

14.2

Mesonotum

......continued on the next page



63

TABLE 11. (Continued) ♀ HT [HLMD]

♂♂

♀♀

♂ [ZMUH]*

♀ [ZMUH]*

5.8–8.8

9.2–10.0

-

-

–

–

-

-

7.6–10.0

15.2–17.3

11.2

16.7

–

–

-

-

Profemora

15.2–19.0

15.6–17.5

17.5

18.0

Protibiae

15.9–20.0

17.8–19.5

18.5

18.5

Mesofemora

11.9–15.3

12.5–15.3

14.6

15.2

Mesotibiae

14.2–17.0

14.2–17.0

15.5

15.8

Metafemora

18.6–22.0

20.1–23.5

25.0

24.3

Metatibia

22.0–26.0

21.8–27.0

27.0

25.7

Antennae

30.0–52.0

33.0–45.0

> 45.0

46.0

Metanotum Median Segment Tegmina Alae

* Type-specimens of H. erringtoniae novaeguineae Günther, 1930. ** Type-specimens (two ♀♀) of H. muelleri var. b. (Haan, 1842) have body-lengths of 70.0 and 77.0 mm.

4.1.2 The grayii species-group The grayii species-group of Haaniella contains four species, two of which are endemic in Borneo, one represented in Sumatra and another species found in southern Vietnam. The disjunct distribution of H. gorochovi n. sp. in Vietnam is remarkable but similar distributional patterns, with species of one genus being represented in Borneo as well as Vietnam and South China but obviously absent in intervening areas such as Peninsular Malaysia, Thailand or Kamputschea are for instance known from genera of the related subfamily Dataminae (e.g. Pylaemenes Stål, 1875). The two Bornean species are closely related and certainly sister-taxa. The grayii-group is sister to the echinata-group and both together are the sister-group of the muelleri speciesgroup of Haaniella. Characters shared with the echinata-group are the longitudinal median row of spines on the ventral surface of the meso- and metafemora, specialized apex of the epiproct of ♀♀ and the setose eggs. Characters that distinguish members from those of the closely related echinata-group are the presence of a prominent central pair of spines on abdominal tergites II–IV and lack of a triangular sub-basal tooth on the lateral margins of the anal segment of ♂♂, as well as the long epiproct and apically angulate or notched subgenital plate of ♀♀. The long epiproct of ♀♀, which clearly projects over the subgenital plate, and roundly angular or notched apex of the subgenital plate are shared with the muelleri species-group. The presence of an enlarged pair of spines on abdominal tergites II–IV of ♂♂ well characterizes this speciesgroup and is not found elsewhere throughout the genus Haaniella. The eggs are always barrel-shaped and setose to a variable degree. The Sumatran representative is remarkable for its small size and is the smallest known species of the entire genus with ♂♂ having a body length of less than 40 mm. In contrast to the other two species-groups the number of antennomeres shows variability, ♀♀ having 24–25 and ♂♂ 22–24 antennomeres. For a comparison and distinction from the other two species-groups of Haaniella see Table 1.

Haaniella dehaanii (Westwood, 1859) (Figs. 214–229, 346, 385–386, 389, 406, 424–425, 430, 460–463, 494) [De Haan’s Haaniella] Heteropteryx DeHaanii Westwood, 1859: 83, pl. 3: 7 (♀). HT, ♀: Borneo, 48–40; BMNH(E) #845208 [NHMUK]. Redtenbacher, 1906: 169. Dohrn, 1910: 403.


HENNEMANN ET AL.

Brock et al., (in press). [Type data] Haaniella Dehaanii, Kirby, 1904b: 398. Haaniella dehaanii, Bragg, 1998: 16, figs. 12–14 (♀ abdomen) & 31–32 (egg). Seow-Choen, 2000a: 151, fig. 1 (♂), 2 (♀). Bragg, 2001: 76, figs. 15 e–f (egg), 16 a–e (abdomen of ♀ and egg), pl. 3: a (♂). Dräger, 2011: 50, figs. 10: 6a–c (♀ abdomen), 11: 2 (♂ thorax), 21 (♂,♀), 22 (♀ defensive reaction), 23 (♀), 24 (♂), 37: 4 (egg). Seow-Choen, 2016: 406, Figs. 1024-1028. Haaniella deHaani, Günther, 1932: 313. Haaniella grayi dehaani, Günther, 1944: 71. Heteropteryx dipsacus Redtenbacher, 1906: 170. LT, ♀: Sarawak, Borneo, Coll. G. Doria, 1865–1866, Batan Lupar 1867, leg. O. Beccari, Best. Verz., No. 85 [MSNG]; PLT, ♂: Sarawak, Borneo, Coll. Doria & Beccari [MSNG]. [Synonymised by Bragg, 1998: 16, designation of LT] Haaniella dipsacus, Günther 1932: 313. Haaniella grayi dipsacus, Günther, 1944: 72.

Material examined (48 ♂♂, 45 ♀♀, 3 nymphs, eggs): SARAWAK: 1 ♂, 1 ♀: bred by Potvin, 1994 [RBINS]; 1 nymph: 4400 Penrissen, det. Klante, 1965 [RMNH]; 1 ♀: ex Zucht F. Hennemann, VIII.1996, Sarawak; ex coll. F. Hennemann [SMNS]; 1 ♂: Borneo, NW-Sarawak, Mount Serapi, 100 m, leg. D´Hulster, 1993 [coll. FH, No. 0115-1]; 1 ♂, 1 ♀: ex Zucht W. Potvin, 1996, Sarawak [coll. FH, No’s 0115-2 & 3]; 1 ♀; ex Zucht P. Heusi (Schweiz), 1998, urspr.: Sarawak [coll. FH, No. 0115-16]; 7 ♂♂, 6 ♀♀, 75 eggs: ex Zucht F. Hennemann, 1998, Herkunft: NW-Sarawak [coll. FH, No’s 0115-4 to 17 & E); 11 ♂♂, 6 ♀♀: ex Zucht F. Hennemann, 2013–2014, Herkunft: NW-Sarawak, Mt. Serapi, PSG 126 [coll. FH, No’s 0015-18 to 34]; 5 ♀♀: Ex Zucht O. Conle, 1998, Herkunft: Borneo, NW-Sarawak, Mt. Serapi (PSG No. 126) [coll. OC]; 2 ♂♂: Ex Zucht O. Conle, 1998–99, Herkunft: Borneo, NW-Sarawak, Mt. Serapi (PSG No. 126) [coll. OC]; 3 ♂♂, 3 ♀♀: Ex Zucht O. Conle 1000, Herkunft: Borneo, NW-Sarawak, Mt. Serapi (PSG No. 126); 18 ♂♂, 13 ♀♀: Ex Zucht O. Conle, 2014: Herkunft: Borneo, NW-Sarawak, Mt. Serapi (PSG No. 126) [coll. OC]; 3 ♂♂, 3 ♀♀: Gunong Gading National Park, Lundu, leg. FSC [coll. FSC]. KALIMANTAN: 1 ♂: Sambas, W-Borneo, v. Mort leg., 3577 [MNHU]; 1 ♀ (nymph): Pontianak, Hoost leg. [MNHU]. BORNEO: 2 ♀♀, 1 ♀ (nymph): Museum Paris, Borneo, R. Oberthur, 1898 [MNHN]; 1 ♂: Mus. Caes. Vind. Borneo, Borneo, det. Redtenb. Heteopteryx grayi var., det. Holdhaus Heteropteryx n. sp.,, Heteropteryx grayi Westw. Var [MNHN]. JAVA (in error): 1 ♀: Coll. Br. V.W., Java, Dr. Candèza; det. Redtenb. Heteropteryx dehaani; 7271 [NHMW, No. 280]; 1 ♂ (nymph): Coll. Br. V. W., Java, Türk; det. Redtenb. Heteropteryx dehaanii; 6843 [NHMW, No. 280]. INDIA (in error): 1 ♀: Museum Paris, Indes anglaises, Balasore, R. Oberthür, 1898 [MNHN]. Diagnosis: Closely related to H. grayii (Westwood, 1859) which is here interpreted as the sister-taxon. The lateral teeth of the epiproct of ♀♀ is a synapomorphy of these two species. Both sexes however readily differ from H. grayii by the averaging smaller size, much stockier habitus and relatively shorter body segments, considerably stronger anttero-lateral mesonotals, very large anterior-mesal mesonotals, which are roughly equal in size to the antero-lateral and anterior mesonotals and together with these form a conspicuous transverse row of six spines, and having only two long posteromedial mesonotals between the bases of the tegmina (four in grayii). Furthermore, both sexes have the coxa-trochanter joints dull greyish blue (bright green in grayii) and the interior surface of the metacoxae black (dull red in grayii). Males may also be distinguished from those of H. grayii by the dull reddish brown or black spines of the thorax (bright green in grayii) and colouration of the tegmina, lacking the white apical portion typical for H. grayii. Eggs differ from those of H. grayii by the smaller dimensions, being somewhat more globose, more regularly arranged hairy structures of the capsule and shape of the micropylar plate. Etymology: Named in honour of Wilhelm de Haan (1801–1855), a well-known Dutch entomologist. Description: ♀♀ (Figs. 214–216). Moderately sized (body length 77.0–99.5 mm) and very stocky species with six very prominent anterior mesonotal spines and two moderately distinct posterior mesonotal spines between



65

the bases of the tegmina. Colour variable and ranging from dull ochre over various shades of brown to almost black (usually darkening with age), either plain or with pale markings. Mesonotum often with a large triangular pale ochre to cream or whitish marking anteriorly, more rarely with a pale ochre marking roughly in centre (Fig. 461). Anterolateral angles and the antero-lateral mesonotals often pale cream to whitish. Dorsal armature of head and thorax dull orange to pale reddish brown with the tips sepia. Tegmina ranging from plain dull ochre over brown to almost black and sometimes with a bold pale spot medio-laterally (Fig. 216). Interior surface of metafemora pale red in the basal half; metacoxae black interiorly (Fig. 346). Coxal trochanters of metacoxae dull greyish blue, the membranes between abdominal sternites I–III pale grey (Fig. 389). Head ovoid and about 1.2x longer than wide with the vertex moderately convex in posterior portion; armed with eight spines (Fig. 229). Vertex with two pairs of distinct and almost equally sized coronals which roughly form the four corners of a square (Fig. 228). Posterior margin with two prominent lateral coronals, which are somewhat stronger than the anterior and posterior coronals. Supra-occipitals minute to obsolete. Behind the eyes with a fairly distinct supra-orbital (Fig. 229). Antennae consisting of 25 segments and ± reaching half way along abdominal tergum III. Pronotum slightly longer but considerably wider than head, a little wider than long and trapezoidal with posterior margin some 1.4x wider than anterior margin; the lateral margins concave, indented medially and set with four very minute spines. Antero-lateral pronotals very prominent, postero-lateral pronotals small and near posterior margin with a pair of indistinct inter-posterior pronotals. Just before transverse median sulcus with a pair of very large and strong pre-median pronotals (Fig. 229). Mesothorax gently and gradually widening towards the posterior with posterior portion some 1.2x wider than anterior margin. Mesonotum slightly narrowing towards the posterior and only about 1.2x longer than pronotum. Lateral margins with 4–6 fairly small spines, which decrease in size towards the posterior. Antero-lateral mesonotals very large and often ± backcurving. Disc with a pair of very large often ± backcurving anterior mesonotals and about equally sized anterior-mesal mesonotals; these together with the antero-lateral mesonotals forming a characteristic transverse row of six almost equally sized spines (Figs. 228–229). Pre-median mesonotals distinct but much smaller than the anterior mesonotals; disc otherwise set with a variable number of minute spinules (Fig. 228). Between bases of tegmina with two distinct, posteriad directed posteromedial mesonotals; often two further minute spines in front (Fig. 229). Tegminal spine fairly small. Mesopleurae with a strong supra-coxal spine and a marginal row of 5–7 unequally sized but fairly distinct, slender laterals. Metapleurae with lateral margin rounded and armed with five spines, the anterior three spines prominent and sometimes gently backcurving, the two posterior spines considerably smaller; supra-coxal very prominent and almost equal in size to the six anterior mesonotal spines. Sensory area of profurcasternum moderately distinct and transversely oval in outline; segment otherwise with two minute spinules laterally. Mesosternum with a lateral row of three short but strong spines, which increase in size towards the posterior. Metasternum with 4–5 small and blunt spines on each side of the median line; the outermost largest. Tegmina ± reaching half way along abdominal tergum II, roundly rhomboidal in shape with the apical portion increasingly narrowed; the radial vein strongly raised a forming a prominent longitudinal keel in basal portion of tegmen. Abdominal tergites II–III widening, IV widest and V–X narrowing; II–VII much wider than long, II and VII somewhat longer than III–VI. Lateral margins of III–VI very gently deflexed and rounded and armed with six slender, unequally sized spines. III–V with a small pair of median spines, II–VII all with a transverse posterior row of 6–8 very minute spinules. Sternites II–V with four short spines on each side of the median line, which roughly form a square; VI only with 4–6 minute spines. Praeopercular organ formed by a rounded swelling and two short diverging carinae on sternum VII (Fig. 222). Tergum VIII with lateral margins smooth. Anal segment with a very blunt longitudinal keel, the posterior margin with a distinct triangular excavation (Fig. 221). Epiproct about equal in length to anal segment and projecting considerably over apex of subgenital plate (Fig. 220); the apical portion with six triangular teeth (Fig. 221). Cerci small and compressed laterally. Subgenital plate with apex incised and not reaching apex of epiproct (Fig. 222). All legs fairly long and comparatively slender with all carinae spinose or dentate, the dorsal carinae of the tibiae only minutely denticuate. Metafemora reaching to abdominal segment VII and metatibiae projecting somewhat beyond apex of abdomen. Ventral surface of femora with a longitudinal median row of small spines (Fig. 362). Two ventral carinae of metatibiae armed with about eight spines, which gradually increase in size and become slightly incurving towards the apex of tibia; the most apical spine of the posteroventral carina largest (Fig. 385). ♂♂ (Figs. 217–219). Medium sized (body length 63.0–75.0 mm) and fairly stocky with six large spines near anterior margin of mesonotum and two prominent posteromedial mesonotals between the bases of the tegmina.


HENNEMANN ET AL.

Colouration ranging from dull ochre to very dark brown (usually darkening with age), rarely with a slight greenish hue. Sometimes with a greenish ochre stripe along lateral margins of mesonotum (Fig. 462). Armature of head and thorax dull green and reddish brown to sepia towards the tips. Tegmina ranging from plain dull ochre over brown to almost black, often with most of the apical portion pale cream to white (Fig. 217), more rarely with a broad transverse median band or only the lateral portion of the apical half whitish or with a pale cream to white diagonal stripe. Interior surface of metafemora dull reddish brown in the basal half; metacoxae black interiorly. Coxal trochanters of metacoxae dull greyish blue and membranes between abdominal sternites I–III dull red; coxal trochanters of pro- and mesocoxae dark green. Shape and armature of head as in ♀♀, but armature more prominent with all spines somewhat longer and more slender (Fig. 227). Antennae long, consisting of 24 segments and reaching to abdominal segment VI. Pronotum generally as in ♀♀ but the pre-median pronotals longer and more slender. Mesothorax short, only some 1.4x longer than pronotum, roughly parallel-sided with the posterior portion somewhat widened. Mesonotum gradually narrowing towards the posterior and about 1.2x longer than width of anterior margin; armature as in ♀♀ with all spines comparatively longer and more slender and the pair of posteromedial mesonotals in particular much larger and very prominent (Fig. 227). Tegminal spine minute. Mesopleurae with armature as in ♀♀ but the spines averaging longer and the supra-coxal very large. Sensory area of profurcasternum as in ♀♀. Armature of mesoand metasternum as in ♀♀ but metasternum with a well decided pair of posteromedian spines close to posterior margin. Tegmina short, ± reaching to posterior margin of median segment, roundly rhomboidal in shape with the apical portion increasingly narrowed; the radial vein strongly raised and forming a prominent longitudinal keel in basal portion of tegmen. Abdominal segments II–V gently and gradually narrowing, VI narrowest segment and parallel-sided, VII and VIII slightly widening. IV–VI somewhat longer than wide, II and VII a little wider than long. Lateral margins of tergites II–VI with about seven small and slender, unequally sized spines; the two posterior ones with a common base and somwhat enlarged. VII only with 3–4 very minute spines on lateral margins. Dorsal surface of II–V armed with a pair of median spines, which are very prominent on II and decrease in size towards V; otherwise with a few minute paired spinules. VIII–X unarmed but VIII. Sternites II–V with eight spines, VI only with four minute spines and VII unarmed (Fig. 225). Lateral invaginations of sternites II–IV very slender. Anal segment with posterior margin almost straight and rectangular (Fig. 224). Epiproct very small and roundly triangular. Cerci small, club-like and distinctly compressed laterally in basal portion. Vomer roundly triangular with the terminal hook fairly long, slender and somewhat curved dextrally; ochre to dull orange with the apical portion brown to black (Fig. 406). Poculum roughly reaching to posterior margin of tergum IX with posterior margin broad and roundly angular (Fig. 225). Legs all fairly long and comparatively slender, mesofemora almost as long as pro- and mesonotum combined, metafemora reaching about half way along abdominal segment VII and metatibiae projecting distinctly beyond apex of abdomen. Armature as in ♀♀ but the ventral spines of the metatibiae smaller, straight and backward directed (Fig. 386). Variability: In addition to a variable size and relative length of the legs, the body armature shows some variability. Occasionally specimens may have the spines of the thorax significantly curved, e.g. the type specimens of the synonymous H. dipsacus (Redtenbacher, 1906). The colouration varies from pale over dark brown to almost black but generally specimens darken with age. Females sometimes have great parts of the mesonotum pale to mid ochre (Fig. 460) and more rarely specimens occur that have a pale ochre central marking on the mesonotum (Fig. 462). The most significant variability however is seen in the colouration of the tegmina. In ♀♀ these are either plain dark brown (Fig. 461), but sometimes bear a bold pale spot medio-laterally (fig. 216). In ♂♂ the tegmina either have the apical portion pale straw to creamish white (Fig. 217, 462), bear a broad transverse median band or only have the lateral portion of the apical half whitish (Fig. 463). Eggs (Figs. 424–425, 430): Of moderate size, barrel-shaped with polar-area rounded; 1.2–1.3x longer than wide. Entire surface densely setose, the setae obscuring most of the capsule and micropylar details, fairly short but becoming longer towards anterior portion of capsule and on operculum; setae of capsule generally directed anteriad. Micropylar plate generally X-shaped; anterior extensions at an angle of about 90° to each other with the apical half slightly widened and incurving; posterior extensions straight, shorter than anterior extensions and with the apex ± strongly broadened and club- to roundly hook-shaped. Micropylar cup represented by an elongate impression, posteriorly accompanied by a rounded tubercle. Operculum almost circular and gently convex. General colouration plain mid brown. Measurements [mm]: length incl. operculum 7.6–10.0, length 7.5–9.7, width 5.6–6.5, height 5.9–7.2.



67

Comments: Westwood (1859: 83) originally described Heteropteryx dehaanii from a single ♀ in NHMUK and provided a quite detailed description and illustration (pl. 3: 7). Redtenbacher (1906: 170) described Heteropteryx dipsacus from a ♂ and ♀ in MCSN, which he distinguished from H. dehaani Westwood by the strikingly curved spines of the thorax. Bragg (1998: 16) synonymised H. dipsacus (Redtenbacher, 1906) with H. dehaani (Westwood), as the curvature of the spines of H. dipsacus and the straight spines of H. dehaani is a variable feature within the population on Mount Serapi (NW-Sarawak). Günther (1944: 71) did not recognize the conspecificity of the two taxa and erroneously treated both H. dehaani and H. dipsacus as subspecies of Haaniella grayii (Westwood, 1859). Subsequent collections and observations have however shown H. grayii and H. dehaanii to be sympatric on Mount Serapi, and no intermediate forms have been found. Followingly they are treated as distinct species (Bragg 1992: 5) which are however very closely related and represent sister-taxa. H. dehaanii has been successfully reared in Europe since 1990 (Phasmid Study Group culture list, culture No. 126) and stock had been re-introduced on several occasions since then. Rearing is easy in very humid conditions and alternative foodplants include bramble (Rubus fruticosusm, Rosaceae), raspberry (Rubus idaeus, Rosaceae), roses (Rosa spp., Rosaceae), oaks (Quercus spp., Fagaceae), pyracantha (Pyracantha coccinea, Rosaceae) and salal (Gaultheria shallon, Ericaceae). Distribution (Fig. 494): Restricted to W-Borneo. NW-Sarawak: Batang Lupar [MSNG]; Kubah National Park, Mount Serapi [coll. FH, coll. OC, coll. PEB]; Penrissen [RMNH]; Gunong Gading National Park, Lundu [coll. FSC]. NW-Kalimanatan: Pontianak [MNHN]; Sambas [MNHU]. Specimens examined: 96 TABLE 12. Measurements [mm] of Haaniella dehaanii (Westwood, 1859) ♂♂

♀♀

63.5–73.0 63.0–75.0

77.0–91.8 89.0–99.5

Head

5.8–7.0

9.5–10.6

Pronotum

7.3–7.9

9.2–10.2

Mesonotum

9.5–10.3

12.5–13.2

Metanotum + median segment

9.5–9.7

10.5–12.3

Tegmina

10.7–12.5

15.0–17.4

Profemora

15.3–17.5

16.7–17.8

Protibiae

18.3–19.8

19.0–20.8

Mesofemora

13.0–14.5

20.0–21.3

Mesotibiae

16.0–16.4

16.0–18.8

Metafemora

21.0–21.7

22.2–24.0

Metatibiae

23.2–25.5

27.0–28.5

Antennae

48.0–53.0

42.0–50.0

Body Body (after Bragg)

Haaniella gorochovi Hennemann, Conle, Brock & Seow-Choen n. sp. (Figs. 230–242, 363, 390, 405, 420–421, 464–465, 493) [Gorochov’s Haaniella] HT, ♂: Vietnam, Prov. Gia Lai, Buôn Luoi (20km N Kannack), 3–11.XI.1993, A. Gorochov [ZIN]. PT, 1 ♀ (+ 8 eggs), 2 ♀♀ (nymphs): Vietnam, Prov. Gia Lai, Buôn Luoi (20km N Kannack), 3–11.XI.1993, A. Gorochov [ZIN]. PT, ♀: Vietnam, Prov. Gia Lai, Buôn Luoi (20km N Kannack), 1–10.V.1995, A. Gorochov [ZIN]. PT, nymph: Vietnam, Prov. Gia Lai, Buôn Luoi (20km N Kannack), 22–31.III.1995, A. Gorochov [ZIN].


HENNEMANN ET AL.

PT, 1 ♂, 2 ♀♀ (nymphs): Vietnam, Prov. Gia Lai, Tram Lap (40 km N of Kannack), XII.1988, A. Gorochov [ZIN]. PT, ♂: Vietnam, Prov. Gia Lai, Tram Lap (40 km N of Kannack), 13.XI.1993, A. Gorochov [ZIN]. PT, 2 ♀♀ (nymphs): Vietnam, Prov. Gia Lai, Distr. Ka Bang, Krong Pa, IX.1997, N. Orlov [ZIN]. PT, 2 ♀♀ (nymphs): Vietnam, Prov. Phykhank Nhachang, forest, leaf litter, 27.06.96, T. K. Sergeeva [ZIN]. PT, ♂: Vietnam, Dak Lak Province, Krong Kmar Distr., Chu Yang Sin Nat. Park, 12°25’26’’N, 108°21’52’’E, H ~ 950m a.s.l.; May 2014, coll. Abramov A.V. (exp. of Russia-Vietnam Tropical Centre), No. F3447.13 [MMUM]. PT, ♀: Vietnam, Dak Lak Province, Krong Kmar Distr., Chu Yang Sin Nat. Park, 12°25’26’’N, 108°21’52’’E, H ~ 950m a.s.l.; May 2014, coll. Abramov A.V. (exp. of Russia-Vietnam Tropical Centre), No. F3447.12 [MMUM]. PT, 2 ♂♂, 4 ♀♀, eggs: Vietnam, Lam Dong Prov., Bidoup-Nui Ba N.P., 12°26'N 108°30'E, 21–25.Vii.2014, night coll., leg. J. Constant & J. Bresseel, GTI Project. I.G.32.779 [RBINS]. PT, 2 males, 2 females, 4 eggs: ex Zucht: F. Hennemann 2016, F1-Generation, Herkunft: Vietnam, Lam Dong Prov., BidoupNui Ba N.P., leg. J. Constant & J. Bresseel [coll. FH, No's 0860-1 to 4 & E1]. PT, 1 ♂, 1 ♀: Vietnam, Gia Lai: An Khe Dist., Tram Lap. 14°26'N 108°33'E 14–30 JUN 1996, B. Hubley DC Currie, J. Swann, ROM961046 [ROM].

Diagnosis: The systematic position of this new species is not fully clarified, but features such as the conspicuous central pair of spines on abdominal tergites II–IV of ♂♂ and long epiproct of ♀♀, which projects considerably over the apex of the subgenital plate, place H. gorochovi n. sp. in the grayii species-group. The most closely related species and possible sister-taxon appears to be the small Sumatran H. parva Günther, 1944, with which it shares the lack of lateral teeth of the epiproct of ♀♀. From H. parva both sexes of this new species however readily differ by the much greater size, considerably more developed and acute body and leg armature, contrasting red cephalic and thoracic spines and having two very prominent instead of four small posteromedial mesonotals between the bases of the tegmina. Females furthermore differ from those of H. parva by the broadened and distinctly notched apex of the subgenital plate. The eggs are much larger than those of H. parva and have the four extensions of the micropylar plate decidedly longer. Etymology: Named in honour of the collector, Dr Andrej Gorochov (ZIN), who has published numerous works on orthopteroid insects and their classification, particularly on the Grylliidae. Description: ♀♀ (Figs. 230–231). Medium sized (body length 90.0–101.0 mm) and stocky species with a short and very broad mesothorax, reduced mesothoracic armature and a bifid epiproct. Colour variable and ranging from dull ochre to dark greyish brown (usually darkening with age), either plain or with various pale markings or darker speckles and occasionally with a faint, broad dull brown longitudinal median stripe along dorsal body surface. Complete armature of head and body bright red with the tips black. Base of the large pre-median pronotals, anterior-mesal mesonotals and posteromedial mesonotals usually black. Tegmina ranging from plain dull ochre to dark brown and sometimes with most of the basal portion pale cream to whitish; these pale areas forming a roughly heart-shaped marking when the tegmina are closed. Interior surface of metafemora and metacoxae pale orange. Coxal trochanters of metacoxae dull greyish, the membranes between abdominal sternites I–III pale orange (Fig. 390). Head ovoid and about 1.2x longer than wide with the vertex moderately convex in posterior portion; armed with ten spines (Figs. 234–236). Vertex with two pairs of distinct and almost equally sized coronals which roughly form a square (Fig. 234). Posterior margin with two prominent lateral coronals, which are somewhat smaller than the anterior and posterior coronals. Behind the eyes with a fairly prominent supra-orbital, which is roughly equal in size to the lateral coronals (Fig. 234). Supra-occipitals distinct and supra-antennals small. Antennae consisting of 26 segments and ± reaching to posterior margin of abdominal tergum III. Pronotum slightly longer and considerably wider than head, a little longer than wide and trapezoidal with posterior margin some 1.4x wider than anterior margin; the lateral margins concave, indented medially and set with 3–4 minute spines. Antero-lateral pronotals distinct but variable in size and sometimes accompanied by a somewhat smaller spine on a common base posteriorly, the postero-lateral pronotals smaller than the antero-lateral pronotals. Near posterior margin with one or two pairs of small to obsolete inter-posterior pronotals and near anterior margin usually with a pair of small anterior-mesal pronotals. Just before transverse median sulcus with a pair of very large and strong pre-median pronotals, which are the largest spines of the entire body (Figs. 235–236). Mesothorax gently and gradually widening towards the posterior with posterior portion some 1.2x wider than anterior margin. Mesonotum distinctly narrowing towards the posterior with the anterior portion laterally overlapping the mesopleurae and almost as wide as posterior portion of mesothorax; only about 1.4x longer than pronotum. Lateral margins of mesonotum with 4–6 fairly small spines, which slightly decrease in size towards the posterior. Antero-lateral mesonotal very large and



69

much more prominent than other lateral spines. Disc with a pair of very small to medium sized anterior mesonotals. The anterior-mesal mesonotals ranging in size from small or almost obsolete (specimens from Bidoup-Nui Ba National Park, → see comments on variability below, Fig. 235) to large and prominent; occasionally even represented by two pais of spines (Fig. 236). Between bases of tegmina with two distinct, upright posteromedial mesonotals (Figs. 234–236). Disc otherwise with a fine longitudinal median carina and 6–8 minute paired spinules in the median portion. Mesopleurae with a strong supra-coxal spine and a marginal row of 5–6 unequally sized laterals, the anterior one of which is considerably larger than the remaining; disc often with a further medium sized spine in the anterior half. Metapleurae with lateral margin rounded and armed with five spines, the anterior three spines prominent with the third largest, the two posterior spines considerably smaller; supra-coxal very prominent and about equal in size to the mesopleural supra-coxal. Sensory area of profurcasternum very large and covering most of anterior half of segment; a small spine near each lateral angle. Mesosternum with a lateral row of three fairly distinct spines and a few further but smaller spines interiorly. Metasternum with six medium sized spines on each side of the median line; the outermost pair largest. Abdominal sternum I with a pair of fairly large, backcurving spines near posterior margin. Tegmina slightly projecting over posterior margin of median segment, roundly rhomboidal in shape with the apical portion increasingly narrowed; the radial vein strongly raised a forming a prominent longitudinal keel in basal portion of tegmen. Tegminal spine moderate. Abdominal tergites II– III widening, IV widest and V–X narrowing; II–VII much wider than long. Lateral margins of III–VI very gently deflexed and rounded and armed with 5–6 slender, unequally sized spines, which are most decided on IV–VI. III– VII with a transverse row of six minute spinules along posterior margin, III with one pair and IV–VII with two to three pairs of small median spines, forming two longitudinal rows and placed and two faint longitudinal carinae. Tergites VIII and IX also with two faint longitudinal, roughly parallel median carinae, each of which are armed with several minute spinules; lateral margins of VIII with 4–6 very minute teeth. Lateral invaginations of sternites II–IV very indistinct. Sternites II–VI with eight paired spines, which are most decided on III and slightly decrease in size towards VI. Praeopercular organ on sternum VII formed two short median spines at posterior margin and a prominent median swelling some distance off the posterior margin, which is apically protuded into a blunt, posteriad directed hook (Figs, 238, 240). Anal segment with a very blunt longitudinal median keel and the posterior margin broadly rounded (Fig. 239). Epiproct about equal in length to anal segment, gently downcurving and distinctly projecting over apex of subgenital plate (Fig. 238); the apex bifid (Figs. 239–240). Cerci small and compressed laterally. Subgenital plate with apex somewhat deflexed and the posterior margin roundly excavated to form two blunt, rounded outer angles (Fig. 240). All legs stocky and of maderate lengthwith all carinae spinose or dentate, but the dorsal carinae of the tibiae only minutely denticulate. Metafemora somewhat projecting over posterior margin of abdominal segment V and metatibiae roughly reaching to apex of abdomen. Ventral surface of femora with a longitudinal median row of 4–5 small spines in the basal half (Fig. 363). Two ventral carinae of metatibiae armed with about six incurving spines, which gradually increase in size towards the apex of tibia; the most apical spine of the anteroventral carina largest (Fig. 230). ♂♂ (Figs. 232–233). Fairly small (body length 59.0–61.0 mm) and stocky insects with a short and broad mesothorax and prominent body armature. Colouration ranging from dull ochre to dark brown (usually darkening with age) with pale and dark mottling and speckles; sometimes with a slight dark longitudinal stripe on pro and mesonotum. Complete armature of head and body bright red with the tips black. Base of the large pre-median pronotals, anterior-mesal mesonotals and posteromedial mesonotals usually black. Most of tegmina whitish to cream with only the apical portion and basal lateral portions brown to black (Fig. 233). Interior surface of metafemora and metacoxae dull orange. Coxal trochanters of metacoxae dull grey and membranes between abdominal sternites I–III dull orange. Shape and armature of head as in ♀♀, but armature considerably more prominent with all spines comparatively longer and more slender (Fig. 237); supra-orbitals in particular much larger than in ♀♀. Antennae fairly long, consisting of 26 segments and reaching to abdominal segment IV or V. Pronotum generally as in ♀♀ but the pre-medials longer and even more prominent. Mesothorax short, only some 1.3x longer than pronotum, roughly parallel-sided with the only posterior portion somewhat widened. Mesonotum gradually narrowing towards the posterior with the anterior portion slightly overlapping mesopleurae and almost as wide as posterior margin of mesothorax; armature generally as in ♀♀ with all spines comparatively stronger and more elongate (Fig. 237). Anterior mesonotals obsolete or lacking and anterior-mesal mesonotals much stronger than in ♀♀. Mesopleurae with armature as in ♀♀ but the spines averaging longer and the supra-coxal very large. Sensory area


HENNEMANN ET AL.

of profurcasternum as in ♀♀. Armature of meso- and metasternum as in ♀♀ but metasternum with a well decided pair of posteromedian spines close to posterior margin. Tegmina short, ± projecting over posterior margin of median segment, roundly rhomboidal in shape with the apical portion increasingly narrowed; the radial vein strongly raised and forming a prominent longitudinal keel in basal portion of tegmen. Tegminal spine fairly distinct. Abdominal segments II–VII gently and gradually narrowing, VII narrowest segment and roughly parallelsided, VIII widening. II–III somewhat longer than wide, IV and V roughly square and VI–VII a little longer than wide. Lateral margins of IV–V gently rounded and lateral margins of II–VII armed with 5–6 fairly distinct, slender and acutely pointed but somewhat unequally sized spines. Tergum VIII only with four fairly small spines along lateral margins. Dorsal surface of II–V armed with a pair of median spines, which is very prominent on II and decrease in size towards V; otherwise set with a few minute paired spinules. VI and VII with several small paired spines and II–VII each with a transverse row of six small spines close to posterior margin. VIII–X only set with a variable number of small paired spinules; the median ones on VI–VII arranged in two longitudinal rows. Sternites II–VI armed with eight spines, which are most prominent on III and decrease in size towards VI. The anterolateral spine pale cream. A fine longitudinal median carina on III–V and the lateral invaginations of II–IV fairly indistinct and slender. Sternum VIII only with 4–6 small paired spines. Anal segment roughly parallel-sided with posterior margin broadly rounded and roundly indented medially (Fig. 241). Epiproct very small and roundly triangular. Cerci small, club-like and distinctly compressed laterally in basal portion. Vomer broad at the base, distinctly curved dextrally with a distinct longitudinal median furrow and the outer portion considerably swollen; the terminal hook black, short, distinctly upcurving and directed towards the right (Fig. 405). Poculum roughly reaching to posterior margin of tergum IX with posterior margin very broadly rounded (Fig. 242). Legs all fairly long and moderately stocky, mesofemora almost as long as pro- and mesonotum combined, metafemora reaching about half way along abdominal segment VII and metatibiae projecting considerably beyond apex of abdomen. Armature generally as in ♀♀ but the ventral spines of the metatibiae smaller and less curved (Fig. 232). Variability: In addition to a variable colouration, considerable variability is seen in the mesothoracic armature of both sexes. While specimens from Bidoup-Núi Bà National park have the anterior-mesal mesonotals and anterior mesonotals very small to obsolete (Figs. 234–235), these spines and the anterior-mesal mesonotals in particular are strongly developed and very prominent in specimens from the Gia Lai Province and Phykhank Nhachang Province in ZIN. One ♀ from Gia Lai Province in ZIN even has the anterior-mesal mesonotals represented by two pairs of fairly distinct spines, each of which share a common base (Fig. 236). A ♂ from Gia Lai Province in ZIN has one of the anterior-mesal mesonotals accompanied by a further, somewhat cephalad directed spine (Fig. 237). The two specimens from Dak Lak Province in MMUM are roughly intermediate in aspect of the size of these spines. In general, the specimens in ZIN have the complete body armature comparatively more decided than specimens from the other localities. Interestingly there is also variability in the colouration of the eggs, those from Gia Lai Province and Phykhank Nhachang Province being mid to dull grey, while those laid by specimens from Bidoup-Núi Bà National Park are pale to mid brown (Figs. 420–421). Despite this different colouration no other important morphological difference is seen. Eggs (Figs. 420–421): Moderately sized, barrel-shaped with the polar-area rounded; 1.45x longer than wide. Capsule surface very rough and uneven with numerous irregular impressions and wart-like swellings; all over covered with very short hairy structures; short setae only present on operculum and on opercular collar. Below micropylar cup with a short longitudinal furrow. Microyplar plate broadly X-shaped with all four extensions fairly short; anterior extensions strongly incurving with the apical half almost parallel to each other; posterior extensions slightly directed downward with the apex somwhat expanded posteriorly. Outer margin of plate distinctly raised and aculte granulose. Micropylar cup a shallow impression with a fairly distinct and strongly extending cup-like swelling posteriorly. Operculum almost circular and slightly conical. Capsule either plain pale to mid brown with the outer margin of the micropylar plate contrasting dark brown to black or plain mid to dull grey. Measurements [mm]: length incl. operculum 8.2–8.9, length 7.5–7.8, width 5.5–6.0, height 5.9–6.2. Distribution (Fig. 493): South Vietnam: Prov. Gia Lai, An Khe district, Buôn Luoi, 20km N of Kannack [ZIN]; Prov. Gai Lia, An Khe district, Tram Lap, 40km N of Kannack [ZIN]; Prov. Gai Lia, Krông Pa district, Ka Bang [ZIN]; Prov. Phykhang Nhachang [ZIN]; Prov. Lâm Dông, Dam Rông district, Bidoup-Núi Bà National Park [RBINS, NHMW, coll. FH]; Prov. Dak Lak, Krong Kmar district, Chu Yang Sin National Park [MMUM]. Specimens examined: 28



71

TABLE 13. Measurements [mm] of Haaniella gorochovi n. sp. ♂ HT [ZIN]

♂♂ PT [ZIN]

♀♀ PT [ZIN]

Body

61.0

59.0–60.0

90.0–101.0

Head

5.0

5.0

6.0–8.0

Antennae

34.0*

>25.0 *

42.0–44.0

Pronotum

6.0

5.5–6.0

9.0–9.5

Mesonotum

6.5

6.0–7.0

11.0–13.0

Metanotum

4.5

4.5

4.5–5.0

Median segment

5.0

5.0

5.5–6.5

Tegmina

10.0

11.0–12.0

16.0–18.0

Alae

9.0

9.0–10.0

12.0–13.0

Profemora

14.0

14.0

18.0–20.0

Protibiae

15.0

15.0

19.0–21.0

Mesofemora

13.0

13.0

16.0–18.0

Mesotibiae

15.0

15.0

18.5–19.0

Metafemora

17.0

17.0–18.0

24.0–28.0

Metatibia

20.0

19.0–20.0

27.0–30.0

* tips broken ** seven female nymphs (28.0–65.0mm) have been excluded from the measurements given above

Haaniella grayii (Westwood, 1859) (Figs. 86, 95, 243–259, 387–388, 391, 407, 426–427, 429, 466–469, 493) [Gray’s Haaniella] Heteropteryx Grayii Westwood, 1859: 82, pl. 30: 2 (♂) & 3 (♀). LT, ♀: Borneo; 3/4, 1830–73 [UMO, 529]; [Not: PLT’s - all represent H. saussurei (Kirby): 2 ♂♂, 1 ♀: Borneo [UMO, No. 529]; 1 ♂: Borneo, 58–66; BMNH(E) #845210 [NHMUK]. Brock et al., (in press). [Type data] Heteropteryx grayi, Sharp, 1895: 262. Redtenbacher, 1906: 170. Dohrn, 1910: 402. Haaniella Grayii, Kirby, 1904b: 397. Haaniella grayii, Günther, 1932: 66. Rehn, 1938: 368. Bragg, 1998: 21, figs. 18–20 (♀) & 29–30 (egg). [Lectotype designation] Dräger, 2011: 51, figs. 10: 5-a–c (♀ abdomen), 11: 3 (♂ thorax), 25 (♀♀), 26 (♀), 27–28 (♂), 37: 6a–b (egg). Seow-Choen, 2016: 408, Figs. 1033-1037. Haaniella grayi grayi, Günther, 1935: 124. Günther, 1936: 124. Günther, 1943: 150. Günther, 1944: 71. Bragg, 1990: 157 Bragg, 1991: 8. Seow-Choen, 2000a: 151, fig. 5 (♂), 6 (♀). Bragg, 2001: 85, figs. 15 c–d (egg), 19 a–c (♀ abdomen), 20 (egg). Heteropteryx australe Kirby, 1896: 472. LT, ♂: Australia, 73-12; BMNH(E) #844211 [NHMUK]; PLT, ♀: Australia, 73-12; BMNH(E) #844212 [NHMUK]. [Synonymised by Günther, 1944: 71, Lectotype designated by Bragg, 1998: 21] Kirby, 1904b: 398. Heteropteryx australis, Redtenbacher, 1906: 170. [Not: Haaniella grayi dehaani (Westwood, 1859); listed as subspecies by Günther, 1944: 71 - corrected by Bragg, 1998: 21]. [Not: Haaniella grayi dipsacus (Redtenbacher, 1906); listed as subspecies by Günther, 1944: 71 - corrected by Bragg, 1998: 21].


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Material examined (61 ♂♂, 72 ♀♀, 29 nymphs, eggs): SARAWAK: 1 ♂, 1 ♀: bred by Potvin, Sarawak, 1994 [RBINS]; 3 ♀♀: Borneo, Sarawak, 1865–66, coll. S. Doria (1 ♀: Best. Verz. No. 158) [MSNG]); 1 ♀: Haaniella grayi grayi (Westwood), Sarawak, 1989, P.E. Bragg [NHMUK]; 1 ♂: Male, Haaniella grayi grayi (Westwood) Mt. Sebakam, Bengoh, Sarawak, P.E. Bragg, 29-07-1989 [NHMUK]; 1 ♂, 1 ♀: ex Zucht F. Hennemann, 1996, Sarawak [SMNS]; 1 ♂: ex Zucht. I Abercrombie I.1995, urspr.: Sarawak [coll. FH, No. 0116-1]; 1 ♂, 4 ♀♀: ex Zucht F. Hennemann 1996, urspr.: Sarawak [coll. FH, No’s 0116-3 to 6]; 15 ♂♂, 7 ♀♀, 4 ♂♂ (nymphs), 5 ♀♀ (nymphs), eggs: ex Zucht F. Hennemann, urspr.: NW-Sarawak, Mt. Serapi, 1998-2002 [coll. FH, No’s 0116-7 to 37, E & ED]; 1 ♂, 2 ♀♀: ex Zucht H. Dräger 2011-2013, Herkunft: NWSarawak, Mt. Serapi [coll. FH, No’s 0116-38 to 40]; 2 ♀♀: NW-Sarawak, Mt. Serapi, 200m, leg. F. Hennemann & O. Conle 28.VII.1996 [coll. OC]; 2 ♀♀: Borneo, NW-Sarawak, MT. Serapi, 200 m, 28.07.1996, leg. Conle & Hennemann [coll. OC]; 27 ♂♂, 22 ♀♀, 1 ♀ (nymph): Ex Zucht O. Conle, 2000–2001, Herkunft: Borneo, NWSarawak, Mt. Serapi, 200 m, leg. Conle & Hennemann 28.07.1996 [coll. OC]. KALIMANTAN: 7 ♂♂, 7 ♀♀, 2 ♂♂ (nymphs), 4 ♀♀ (nymphs), 31 small nymphs: Borneo, Exped. Dr. Nieuwenhuis, 1894–1898, Boven Mahakam, Long-Bloe-Oe [RMNH]; 2 ♀♀, 1 ♀ (nymph): Mus. Leiden, J. v. Veldhuijzen, Sintang, Goedhuis Borneo-Exped., Haaniella grayii (Westwood) Günther det.; Haaniella grayii (Westwood) Bragg det. [RMNH]; 1 ♂, 3 ♀♀ (nymphs): Borneo, Exped. Dr. Nieuwenhuis, Juli 1894, Boegan; Haaniella grayii (Westwood) Günther, det. [RMNH]; 2 ♀♀: Dr. J. Bosscha, Sambas, W-Borneo, Aug. 1891; Haaniella grayii (Westwood) K. Günther, det. [RMNH]; 2 ♂♂: H.C. Siebers, Midden O-Borneo, 31.8.1925; Heteropteryx grayii Karny det.; Haaniella grayii (Westwood) Klante det. [RMNH]; 1 nymph: H.C. Siebers, Midden O-Borneo, 10.8.1925; Haaniella grayii (Westwood) det. Klante, 1965 [RMNH]; 1 ♂: Tandjong-Amontay, Süd-Ost-Borneo, A.J. Speyer comm., 15.II.1896 [ZMUH]; 3 ♂♂ (nymphs): Aus Alkohol; Borneo, Obat-Fluss, linker Nebenfluss des Mendalam, Nebenfluss des oberen Kapuas, 27.-31.7.1925; K. Günther determ. Vers. Nr. 131, 1932; Haaniella grayi Westw., K. Günther det [ZMUH]; 1 ♂: E. Borneo 95m, Gunungsari, 21.VIII.1956, A.M.R. Wegner; Haaniella grayii (Westwood) det. Bragg, 1995 [RMNH]; 2 ♀♀ (nymphs): E. Borneo 125m, Tabang, 4.10.1956, Bengan River, A.M.R. Wegner; Haaniella grayii (Westwood) det. Klante, 1965 [RMNH]; 1 ♀: P. hasf., Indonesia, Borneo; Haaniella grayii (Westwood) K. Günther det. [RMNH]. “BORNEO”: 1 ♀: Borneo, Wahnes leg.; Nr. 162; Heteropteryx grayii Westw., Brunner v. W. det. [MNHU]; 1 ♂, 1 ♀: Borneo [MNHU]; 1 ♂, 2 ♀♀ (nymphs): Borneo, Coll. Giglio-Tos [MIZT]; 1 ♂: Borneo; Haaniella grayii (Westwood) det. K. Günther [RMNH]; 3 ♀♀: Museum Paris, Borneo, Chaper 1843-91, 1292-91 [MNHN]; 1 ♀: Museum Paris, Borneo, 286-62 [MNHN]; 1 ♀: Mus. Paris, Borneo, Chaper, 1843-91; Coll. Br. v. W. Borneo, Mus. Paris; det. Redtenb. Heteroptery grayii Westw. , 23. 334 [NHMW, No. 283]; 1 ♀ (nymph n5): Nord-Borneo, ex coll. Fruhstorfer; H. Fruhstorfer vend. 6.II.1904 [ZMUH]. NO DATA: 1 nymph: without data, ex Mus. Bogor [RMNH]. Diagnosis: Closely related to H. dehaanii (Westwood, 1859) which is here considered the sister-taxon. The lateral teeth of the epiproct of ♀♀ are synapomorphy of these two species. Both sexes however differ from H. dehaanii by the generally larger size, more slender habitus and relatively longer body segments, indistinct anteriormesal mesonotals, which are considerably smaller than the antero-lateral and anterior mesonotals, and having four rather short posteromedial mesonotals between the bases of the tegmina (two large spines in dehaanii). Furthermore, both sexes have the coxa-trochanter joints bright green (dull greyish blue in dehaanii) and the interior surface of the metacoxae dull red (black in dehaanii). Males may also be distinguished from those of H. dehaanii by the contrasting green spines of the thorax and characteristic colouration of the tegmina, which are white in the apical have and have the main veins in the interior portion marked with dark brown. Eggs differ from those of H. dehaanii by the larger dimensions, being somewhat more elongate, more unevenly arranged hairy structures of the capsule and shape of the micropylar plate. Etymology: Named in honour of the well-known English zoologist George Robert Gray (1808–1872). Description: ♀♀ (Figs. 243, 245). Large (body length 102.0–143.0.0 mm) and fairly slender with comparatively reduced thoracic armature and four distinct posteromedial mesonotals between the bases of the



73

tegmina. Colour very variable and ranging from dull ochre to dark brown, either plain (Figs. 243, 466) or entire body and tegmina irregularly mottled in a lighter brown or straw (Fig. 244). Occasionally specimens occur which have most of the mesonotum whitish to pale cream and all femora with a faint pale transverse band sub-apically (Figs. 245, 467–468). Head with two ± defined dark brown postocular stripes. Cephalic and thoracic armature with the tips dull orange to red and the basal portion dull brown; larger spines of the mesonotum often with green bases. Interior surface of metafemora more or less distinctly pale red or at least with a slight reddish hue in basal half. Metacoxae dull reddish interiorly and all coxal trochanters greyish black with bright green mottling. Head about 1.2x longer than wide with the vertex very gently convex in posterior portion and armed with 12 spines (Figs. 257–258). Vertex with four distinct and almost equally sized coronals which form the four corners of a square; anteriorly with a further pair of much smaller anterior coronals and a minute to obsolete pair of supraantennals (Fig. 257). Posterior margin with two prominent lateral coronals and behind the eyes with a distinct supra-orbital, which is roughly equal in size to the coronals. Supra-occipital very prominent and usually a little larger than the anterior and posterior coronals. Occasionally one or two minute gulars on the cheeks. Antennae consisting of 24 segments and ± reaching half way along abdominal tergum IV. Pronotum slightly longer and noticeably wider than head, a little longer than wide and trapezoidal in outline with posterior margin some 1.2x wider than anterior margin; the lateral margins concave, indented medially and set with 3–4 minute spines. Antero and postero-lateral pronotals well pronounced and roughly equal in size; inter-posterior pronotals minute or obsolete (Figs. 257–258). Just before transverse median sulcus with a very strong pair of pre-median pronotals (Fig. 257) and the posterior portion usually with one or two pairs of minute median pronotals. Mesothorax gently and gradually widening towards the posterior with posterior portion some 1.6x wider than anterior margin. Mesonotum slightly narrowing towards the posterior and about 1.7x longer than pronotum. Lateral margins set with 6–8 fairly small and slender, somewhat unequally sized spines, the antero-lateral spine much enlarged and somewhat backcurving (Fig. 258). Disc with a pair of distinct anterior-mesonotals similar in size to the anterolaterals and an almost equally sized pair of pre-median mesonotals; otherwise only set with a variable number of minute paired spinules or spiniform tubercles (Figs. 257–258). Sometimes a small pair of anterior mesal mesonotals may be present. Between bases of tegmina with four prominent and slender posteromedial mesonotals on a common base (Fig. 257). Tegminal spine moderately distinct. Mesopleurae with a prominent and fairly slender (sometimes backcurving) supra-coxal and three distinct, fairly slender laterals along lower margin; one or two much smaller spines between these enlarged spines (Fig. 258). Metapleurae with lateral margin gently rounded and armed with five long and slender spines, the 3rd largest and the 5th much smaller than previous; supra-coxal distinct but somewhat smaller than that of the mesopleurae. Sensory area of profurcasternum distinct and roundly triangular in outline; profurcasternum otherwise with one or two small spines laterally. Mesosternum with a lateral row of three short but strong spines, which increase in size towards the posterior; otherwise fairly irregularly set with a variable number of small spinules or spiniform tubercles (Fig. 259). Metasternum with two fairly distinct spines on each side of the median line. A fairly pronounced pair of spines on abdominal sternum I (Fig. 86). Tegmina reaching about half way along abdominal tergum II, oval in shape with the apical half increasingly narrowed and the apex fairly selnder; the radial vein strongly raised a forming a prominent longitudinal keel in basal portion of tegmen. Abdominal tergites II–IV almost parallel-sided, following gradually narrowing; II–VI transverse with lateral margins straight and armed with seven small, slender, sub-equally sized spines, the posteriormost being somewhat enlarged. VII only with 3–5 small spines on lateral margins. Dorsal surface of II smooth or at best with a very minute and distant pair of spinules posteriorly, III–VII with a transverse row of four very minute spinules close to posterior margin and III–IV with a ± enlarged pair of spines in centre. Sternites II–VI with eight spines (Fig. 86), the four median ones fairly prominent, larger than the outer ones and roughly forming a square. Lateral invaginations of sternites II–IV fairly distinct and elongate-oval in shape (Fig. 86). Sternum VII only with a few minute spines, the praeopercular organ formed by a rounded posteromedian incision, a rounded black median swelling in front and two backcurving spines at posterior margin of segment (Fig. 256). Tergum VIII strongly convex with lateral margins smooth. Anal segment with a blunt longitudinal median keel and the posterior margin with a wide and very shallow triangular emargination (Fig. 255). Epiproct about as long as anal segment, projecting over apex of subgenital plate and the apical portion with six triangular teeth (Figs. 254–255). Cerci small and compressed laterally. Subgenital plate with apex truncate and the posterior margin gently concave; not reaching apex of epiproct (Fig. 256). All legs fairly long and with all carinae spinose or dentate, the dorsal carinae of the tibiae only minutely denticulate although. Metafemora reaching to abdominal segment VI and metatibiae


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projecting very slightly beyond apex of abdomen. Ventral surface of femora with a longitudinal median row of small spines. Two ventral carinae of metatibiae armed with about seven spines, which strongly increase in size and become increasingly incurving towards the apex of tibia; the most apical spine of the posteroventral carina largest (Fig. 387). ♂♂ (Figs. 246–247). Large (body length 78.0–98.3 mm) and slender with very prominent bright green thoracic armature and gently narrowing abdominal segments III–VI. Colouration ranging from pale creamish to dark brown (usually darkening with age), rarely with a slight dull greenish hue. Head with a well defined black postocular stripe. Complete body armature bright green the tips dull orange to reddish mid brown. Mesonotum sometimes with two broad green slightly converging longitudinal stripes, the lateral portions usually of more pale colour than rest of mesonotum. Tegmina with basal 1/3 dark to blackish brown and the remaining portion white with a network of dark brown to black veins. All tibiae and apical portion of the femora mid to dark brown, the rest of femora with a greenish hue; armature mostly dull green. Metacoxae dull red interiorly. All coxal trochanters bright green (Fig. 391). Shape and armature of head as in ♀♀, but armature more prominent with all spines but the posterior and lateral coronals in particular much longer and more slender (Figs. 251–252). Antennae long, consisting of 23 segments and at least reaching to abdominal segment VII. Pronotum generally as in ♀♀ but more elongate, with the pre-median pronotals in particular much larger and longer. Antero and postero-laterals much more pronounced with the former being somewhat larger than the latter; usually with a minute pair of anterior-mesal pronotals and a more distant pair of very small inter-posterior pronotals (Fig. 251). Mesothorax moderately elongate, some 1.6x longer than pronotum and gently widening towards the posterior. Mesonotum slightly narrowing towards the posterior and about 2x longer than wide. Lateral margins set with 4–5 small subequally sized spines and in anterior portion with a very prominent, long and slender antero-lateral mesonotal. Disc armed with four very prominent spines, a very large pair of anterior mesonotals and a similarly sized pair of post-median mesonotals; surface otherwise sparsely tuberculose to spinulose (Figs. 251–252). Between the bases of the tegmina with four very large and slender posteromedial mesonotals on a common base, the anterior two spines of which are somewhat shorter than the posterior ones (Figs. 251–252). Tegminal spine small. Mesopleurae with 3–4 slender laterals, the most anterior and posterior ones being largest. Supra-coxal very prominent and almost equal in size to the posteromedial mesonotals (Fig. 252). Sensory area of profurcasternum as in ♀♀ and segment otherwise with one or two small spines laterally (Fig. 253). Mesosternum with a fine longitudinal median carina and armature as in ♀♀ (Fig. 253). Metasternum with armature as in ♀♀ but all spines considerably more prominent and occasionally with a further posterior spine (Fig. 89). Tegmina ± reaching to posterior margin of median segment, elongate-oval in shape and with the radial vein raised and forming a longitudinal keel in basal portion of tegmen. Abdominal segments II–VI very slightly gradually narrowing, almost equal in length and averaging 1.3–1.4x longer than wide. VII shorter than previous and widening towards posterior. Tergites II–IV with a central pair of prominent, elongate spines, which is most prominent on II and only indicated on V; surface otherwise irregularly set with small paired spinules. II–VII with 5–7 small and slender, subequally sized spines along lateral margins. Tergum VIII with lateral margins unarmed and widening towards posterior. Sternum I with a moderately distinct pair of spines. Sternites II–VI with eight spines, which are most prominent on II and gradually decrease in size towards VI; VII only with 2–4 minute spines; lateral invagionations of II and II slender (Fig. 89). Anal segment much narrower than IX, decidedly narrowed towards posterior and the posterior margin with a small but deep triangular median indention (Fig. 249). Epiproct very small and rounded. Cerci fairly large, compressed laterally in basal portion and with the apex subangularly and club-like. Vomer broadly triangular with a fairly long, almost straight terminal hook; dull orange with the apex black (Fig. 407). Poculum roughly reaching to posterior margin of anal segment with posterior margin broadly rounded and much labiate (Figs. 248, 250). Legs all long and fairly slender, mesofemora almost as long as pro- and mesonotum combined, metafemora almost reaching to posterior margin of abdominal segment VI and metatibiae projecting distinctly beyond apex of abdomen. Armature generally as in ♀♀ but all spines, except for those on the ventral carinae of the metatibiae, comparatively longer (Fig. 388). Nymphs: The sex of nymphs can already be defined when they are newly hatched, ♀♀ having a slightly elongated epiproct which is missing in ♂♂. Both sexes usually have abdominal tergum VI pale ochre to white during their entire development. The colour-forms of ♀♀ are only achieved in the later instars. Variability: Together with H. echinata (Redtenbacher, 1906) this species is the most variable of the Bornean representatives of the genus. In addition to the size (→ see Table 14 below) the colouration of ♀♀ shows considerable variability and also slight variability is seen in the armature of the thorax. A ♂ in the first author's REVISION OF HETEROPTERYGINAE


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collection (coll. FH 0116-26) exhibits an additional but smaller 5th posteromedial spine between the bases of the tegmina. Females generally occur in two colour-forms, one which is uniformly light to dark brown (Figs. 243, 466) and the other which exhibits more or less defined and variable pale mottling all over the body, tegmina and legs (Fig. 244). Some specimens may have most of the mesonotum whitish to pale cream and rarely there are specimens that are almost black, with reddish brown mottling and markings, and have the entire mesonotum contrasting white (Figs. 467–468). Specimens of both colour-forms often have a white spot at the antero-lateral corner of the mesonotum. While the tegmina are mostly plain light to dark brown, some specimens possess a washed black lateral marking in the basal half. No such remarkable colour variation is seen in ♂♂. Eggs (Figs. 426–427, 429): Large, barrel shaped with polar-area rounded; 1.4–1.5x longer than wide. Entire surface densely setose, the setae obscuring most of the capsule and micropylar details, comparably long and increasing in length towards anterior portion of capsule and on operculum; irregularly arranged in clusters. Micropylar plate X-shaped with all four extensions fairly long and slender; anterior extensions at an angle of about 80° to each other with the apical portion very slightly incurving; posterior extensions broader with the apex strongly expanded and roundly truncate. Entire plate surrounded by a distinctly raised bulge. Operculum almost circular and roundly convex. Micropylar cup a shallow impression with a cup-like swelling posteriorly. General colouration plain mid to dark brown. The length of setae and size show regional variation (Bragg, 1998: 23, 2001: 89), examples from localities throughout Kalimantan having the setae strongly reduced with all the capsule and micropylar details clearly visible. Measurements [mm]: length incl. operculum 8.3–9.7, length 7.5–9.1, width 5.7–6.4, height 7.0–7.2. Comments: Bragg (1998: 21) recognized that the five specimens contained in UMO and NHMUK and marked as Westwood’s types represented two distinct species, and that the ♂ illustrated by Westwood (1859, pl. 30: 2) is not present in either collection. Bragg (1998: 21) designated a ♀ in UMO as the lectotype with reference to Westwood’s illustration and since this agrees with the subsequent treatment of the species (e.g. Redtenbacher, 1906; Günther, 1944). The paralectotypes all represent another species that was subsequently described by other authors and based on other material as Haaniella saussurei Kirby, 1904 and Heteropteryx saussurei Redtenbacher, 1906, the latter of which is a homonym. H. grayii is the averaging largest and together with H. echinata (Redtenbacher, 1906) most variable of the Bornean species. Like H. echinata it is quite widespread and frequently found in numerous localities throughout Sarawak and Kalimantan (→ see distribution below, Fig. 493). Bragg (1990b) reported the indigenous people in Sarawak to eat the eggs of this species, the eggs being boiled in water for about 30 seconds and the shell removed prior to consumption. H. grayii is being successfully reared in Europe since 1990 and culture-stock frequently re-imported from various locations. The original stock originated from Mount Serapi in NW-Sarawak and was included on the Phasmid Study Group culture list as culture No. 125. It is fairly easy to rear in the standard conditions for the genus and accepts numerous alternative foodplants in captivity, including bramble (Rubus fruticosus, Rosaceae), raspberry (Rubus idaeus, Rosaceae), roses (Rosa spp., Rosaceae), oaks (Quercus spp., Fagaceae), pyracantha (Pyracantha coccinea, Rosaceae), elder (Sambucus nigra, Adoxaceae), flowering currant (Ribes spp., Rosaceae), hawthorn (Crataegus monogyna, Rosaceae), eucalyptus (Eucalyptus gunnii, Myrtaceae), ivy (Hedera helix, Araliaceae) and salal (Gaultheria shallon, Ericaceae). Also in the wild H. grayii is known to be fairly polyphagous, feeding on Rubus moluccanus (Rosaceae), Nepenthes ampullaria (Nepenthaceae), Dillenia suffruticosa (Dilleniaceae), Dicranopteris linearis (Gleicheniaceae) and an unidentified Zingiberaceae (→ 7.6). Distribution (Fig. 493): Borneo, endemic. NW-Sarawak: Bau nr. Wind Cave Nature Reserve [NHMUK, coll. PEB]; Kampong Bengoh [coll. PEB]; Mount Sebakam, Bengoh [NHMUK]; Dahan [NHMUK]; Ranchang waterfall, 2 km E of Serian [coll. PEB]; Mount Matang [SMSM]; Kubah National Park, Mount Serapi 90–300 m [SMSM, NMSC, coll. FH, coll. OC, coll. PEB]; Mount Santubong [SMSM]; Gunung Gading NP [photo by P. Bertner]. The fourth author found this speciesto be particularly plentyful in Kubah National Park, Ranchang National Park, Gunong Gading National Park and Serian. Kalimantan: Sungai Ratu Miri, Ratu Miri Logging Camp [coll. PEB]; Kapuas River [RMNH]; Bukit Obat, Mendalam River (affluent of the Upper Kapuas River) [ZMUH]; Mahakam River [RMNH]; Long Bagun “Boegan” [RMNH]; Bluu “Long Bloe oe” [RMNH]; Mount Sari 95 m [RMNH]; Bengen River, Tabang 125 m [RMNH]; Sambas [RMNH]; Sintang [RMNH]; Simatan [RMNH]; “Midden O-Borneo” [RMNH]; Tanjung Simpang N.P. “Tandjong” [ZMUH]. Specimens examined: 165


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TABLE 14. Measurements [mm] of Haaniella grayii (Westwood, 1859) ♂♂

♀♀

81.5–98.3 78.0–95.0

109.0–128.5 102.0–143.0

Head

7.0–8.8

11.5–13.5

Pronotum

7.8–8.8

10.8–12.5

Mesonotum

12.5–14.0

15.8–17.2


12.3–13.5

15.0–17.5

Tegmina

14.8–17.3

23.0–27.0

Profemora

22.0–25.2

23.0–26.5

Protibiae

13.5–18.0

24.2–27.3

Mesofemora

18.5–20.0

20.4–22.5

Mesotibiae

18.5–22.5

22.7–24.3

Metafemora

27.0–31.0

31.0–35.2

Metatibiae

29.5–35.0

35.7–39.0

Antennae

71.0–75.0

72.0–76.0


Haaniella parva Günther, 1944 rev. stat. (Figs. 81, 88, 92, 260–273, 422–423, 470–471, 494) [Dwarf Haaniella] Haaniella parva Günther, 1944: 73, fig. 2 (♂). LT [by present designation], ♂: N.O. Sumatra, Kwalu, Dr. Volz leg.; Haaniella parva K. Gthr. 1944 Syntypus Stett. Ent. Ztg. 10: 73, 1944; Haaniella parva Gthr. 1944 Syntypus [SMTD]; PLT, ♂: same data as LT [ZMUH—not traced].

Materal examined (5 ♂♂, 4 ♀♀, eggs): SUMATRA: 1 ♂: Sumatra, Medan, Env. De Dolok-Baros, 2° semestre 1905, M. Moissinac, Museum Paris, Sumatra 1906, M. Maindron, Heteropteryx vois de haani [MNHN]; 1 ♂, 2 ♀♀: NW-Sumatra: Provinz Sumatera Utara, Berastagi, Roadside leading to Mount Sibayak, 1220 m, leg. F. Seow-Choen 13.IV.2013 [coll. FH, No’s 0868-1 to 3]; 3 ♂♂, 2 ♀♀: N-Sumatra, Berastagi, Roadside leading to Mount Sibayak, 1220 m, leg. F. Seow-Choen 13.IV.2013 [coll. FSC]. Diagnosis: This is the smallest of all known members of Haaniella. The most closely related species and possible sister-taxon appears to be the Vietnamese H. gorochovi n. sp., with which it shares the lack of lateral teeth of the epiproct of ♀♀, a character is shared with the muelleri species-group. Both sexes of this very small species can be separated from H. gorochovi n. sp. by the namely much smaller size, considerably less developed body and leg armature, brown cephalic and thoracic spines (contrasting red in gorochovi n. sp.) and having four small instead of two prominent posteromedial mesonotals between the bases of the tegmina. Females also differ by the bluntly rounded apex of the subgenital plate. The characteristic eggs differ from those of all other members of Heteropteryginae by the shape of the micropylar plate, which has the four extensions very short and is almost roundly rectangular in shape. Etymology: The name (parvus lat. = small) refers to the remarkably small size of this species. Description: ♀♀ (Figs. 260–261). Very small (body length 45.0–52.0 mm) and stocky with fairly reduced and blunt armature. Colour ranging from mid over dark brown to almost black (usually darkening with age), either plain or with irregular mottling. Pro- and mesonotum sometimes with a faint, washed dark longitudinal median band in pale specimens. Dorsal armature of head and thorax black basally with the apical portion of all large spines



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dark reddish brown. Tegmina ranging from plain mid over dark brown to almost black, the anterior margin sometimes slightly greenish in the median portion and occasionally a faint pale basal stripe below radial vein. Interior surface of metafemora with a dull reddish hue; metacoxae dark reddish brown interiorly. Coxal trochanters of metacoxae dull greyish red. Head subglobose and a little longer than wide with the vertex convex in posterior portion; armed with eight stout spines (Figs. 270–271). Vertex with of moderately distinct, blunt and almost equally sized, conical anterior and posterior coronals which roughly form the four corners of a square (Fig. 271). Posterior margin with two similarly sized, blunt lateral coronals and behind the eyes with a fairly distinct supra-orbital (Fig. 271). Frons with a variable number of very small, paired conical spines or spiniform tubercles. Antennae consisting of 24 segments and ± reaching to posterior margin of median segment. Pronotum slightly longer and wider than head, a little wider than long and trapezoidal with posterior margin some 1.3x wider than anterior margin; the lateral margins concave, indented medially and set with about four tubercles. Antero-lateral pronotals moderately prominent, postero-lateral pronotals somewhat smaller and near posterior margin with a pair of very minute inter-posterior pronotals. Just before transverse median sulcus with a pair of prominent, conical pre-median pronotals; surface otherwise irregularly tuberculose (Fig. 270). Mesothorax slightly, gradually widening towards the posterior with posterior portion some 1.4x wider than anterior margin. Mesonotum slightly narrowing towards the posterior and only about 1.2x longer than pronotum. Lateral margins with 4–5 blunt tubercles, which decrease in size towards the posterior. Antero-lateral mesonotals fairly small, blunt and hardly larger than other laterals. Anterior mesonotals prominent, anterior-mesal mesonotals distinct, conical and often with a further, much smaller spine in front. Disc otherwise irregularly tuberculose and between bases of tegmina with two short, blunt posteromedial mesonotals on a rounded common base (Fig. 270). Tegminal spine small. Mesopleurae with a strong and comparatively slender supra-coxal spine (largest of all thoracal spines) and a row of about four blunt spines along lower margin, which decrease in size towards the posterior. Metapleurae with lateral margin moderately rounded and armed with five spines, the anterior two spines of moderate size, the third much smaller and the last two larger than previous ones; supra-coxal prominent, conical and shorter than that of mesopleurae. Sensory area of profurcasternum distinct, triangular in outline and posteriorly bordered by a blunt V-shaped bulge (Fig. 81). Mesosternum with two lateral spines, otherwise finely granulose. Metasternum with four small and blunt spines on each side of the median line; the outermost largest. Tegmina roundly rhomboidal in shape and slightly projecting over posterior margin of median segment; the radial vein strongly raised a forming a prominent longitudinal keel in basal portion of tegmen. Abdominal tergites II–III very slightly widening, IV widest and V–VII gradually narrowing; all considerably wider than long, II and VII somewhat longer than III–VI. Lateral margins of III–VI very gently deflexed and rounded and armed with four very short and blunt spines. III–V with a pair of small, conical median spines; tergites otherwise irregularly rugulose and sparsely tuberculose. Sternites II–V with four short spines on each side of the median line, which roughly form a square; VI only with 4–6 very minute spines. II and III with large, oval, deeply impressed lateral invaginations (Fig. 88). Praeopercular organ on sternum VII formed by a conical median swelling and two diverging carinae behind, which mark an impresssed and smooth triangular area; a short, blunt lateral spine near each carina (Fig. 266). Tergum VIII with lateral margins smooth. Anal segment narrowing towards posterior and with a blunt longitudinal keel; the posterior margin broadly rounded (Fig. 265). Epiproct somewhat longer than anal segment and projecting greatly over apex of subgenital plate (Fig. 264); the apex bifid with a fairly deep triangular incision (Figs. 265–266). Cerci small and compressed laterally. Subgenital plate short and hardly reaching to posterior margin of anal segment with apex bluntly rounded (Fig. 266). All legs moderately long and stocky with carinae of all femora dentate. Pro- and mesotibiae only with a few very indistinct tooth-like structures ventrally. Metafemora reaching to abdominal segment VI and metatibiae roughly reaching to apex of abdomen. Ventral surface of femora with an indicated longitudinal median row of very indistinct granules. Two ventral carinae of metatibiae armed with 7–8 short teeth, which gradually increase in size towards the apex of tibia (Fig. 260). ♂♂ (Figs. 262–263). Very small (body length 25.0–37.0 mm) and fairly stocky with a prominent pair of spines on abdominal tergites II–V. Colouration ranging from mid to very dark brown (usually darkening with age). Armature of head and thorax dark brown and ochre to reddish brown towards the tips. Tegmina with most of the basal half white to ochre, the radial vein contrasting pale reddish brown and the anterior portion ± green (Fig. 263). Interior surface of metafemora dull reddish brown and metacoxae dull reddish brown interiorly. Coxal trochanters as in ♀♀.


HENNEMANN ET AL.

Shape and armature of head as in ♀♀, but armature more prominent with all spines comparatively longer and more slender (Figs. 272–273). Frons with two pairs of small, blunt supra-antennals, which roughly form a square. Antennae long, consisting of 22 segments and reaching to abdominal segment IV. Pronotum generally as in ♀♀ but the pre-median pronotals in particular much longer and more slender (Fig. 272). Portion behind transverse median sulcus usually with 4–6 blunt, paired median pronotals, the most anterior pair of which is largest. Mesothorax short, some 1.4x longer than pronotum, roughly parallel-sided with the posterior portion somewhat widened. Mesonotum gently narrowing towards the posterior and about 1.4x longer than width of anterior margin; armature as in ♀♀ with all spines comparatively longer and more slender and the pair of posteromedial mesonotals in particular much larger and very prominent (Figs. 272–273). The latter usually with a pair of smaller blunt spines in front. Tegminal spine minute. Mesopleurae with armature as in ♀♀ but the spines averaging longer and the supracoxal very long and fairly slender. Sensory area of profurcasternum as in ♀♀. Mesosternum with a lateral row of five short, blunt spines, metasternum only with three spines on each side of the median line. Tegmina small, not reaching posterior margin of median segment, oval in shape and with the radial vein strongly raised and forming a prominent longitudinal keel in basal portion of tegmen. Abdominal segments II–VI gradually narrowing with VII narrowest segment and almost parallel-sided. VIII slightly widening towards the posterior. II–VI decidedly wider than long, VII almost as long as wide. Lateral margins of tergites II and III with four very short and blunt spines, IV–VI only with two minute spines posteriorly, VII with lateral margins unarmed. II–V with a prominent median pair of spines which is most decided on II and gradually decreases in size towards V. Dorsal surface of II–VI armed with a pair of median spines, which are very prominent on II and decrease in size towards VI; otherwise sparsely tuberculose and with a pair of small but spiniform posteromedian tubercles. VII–X with a blunt longitudinal median carina and unarmed. Sternites II–IV with eight blunt spines, the outer ones being larger than the inner ones; V and VI only with four very minute spines. II–III with very prominent, deeply impressed oval invaginations, which are also present in IV but on this segment are much more slender and slit-like (Fig. 92). Anal segment somewhat narrowed towards the posterior, the posterior margin rectangular with a small median indention (Fig. 268). Epiproct small and slightly tectiform. Cerci small, club-like and distinctly compressed laterally in basal portion. Vomer roundly triangular with a short and almost striaght terminal hook. Poculum very bulgy, projecting over posterior margin of tergum IX with posterior margin labiate, somewhat downcurving and broadly rounded (Figs. 267, 269). Legs all fairly long and slender, mesofemora almost as long as pro- and mesonotum combined, metafemora reaching to abdominal segment VI and metatibiae projecting distinctly beyond apex of abdomen. Armature as in ♀♀ but the ventral spines of the metatibiae somewhat smaller (Fig. 262). Variability: Based on the few known specimen only slight variability is seen in the colouration and size of the cephalic and thoracic armature. Eggs (Figs. 422–423): Small, barrel-shaped with the polar-area rounded; 1.4x longer than wide. Surface minutely granulose, sparsely set with minute pointed tubercles and irregularly and sparsely setose, the setae very short and most numerous at polar-area. Microyplar plate roundly rectangular with all four angles slightly expanded and rounded, the posterior extensions larger and more distinctly broadened than anterior angles. Outer margin of plate distinctly raised. Micropylar cup a shallow impression with a fairly distinct, cup-like swelling posteriorly. Operculum almost circular and moderately convex; surface like in capsule. Capsule mid to dark brown the outer margin of the micropylar plate black. The illustrated egg is somewhat discoloured. Measurements [mm]: length incl. operculum 5.5, length 5.0, width 3.2, height 3.8. Comments: The syntype in SMTD is here selected as the lectotype, since the second ♂ recorded by Günther from ZMUH could not be traced and hence is considered lost. The lectotype in SMTD is fairly incomplete, lacking both fore legs, the right hind leg, left metatatarsus and parts of both antennae. Zompro (2004: 198) placed H. parva in the newly described genus Miniopteryx, which is here shown to be a synonym of Haaniella Kirby, 1904 (n. syn.), H. parva merely being a remarkably small member of this genus. The original description by Günther (1944: 73) is quite detailed but Zompro (2004: 199) accused Günther of “highly superficial examination”, stating it was difficult to understand why the author did not mention the “laterosternal excavations” of the abdominal segments in H. parva. These were interpreted as a unique generic character by Zompro (2004: 199), who established and distinguished Miniopteryx from the other genera of Heteropteryginae exclusively on the basis of this character. However, these lateral invaginations of the basal abdominal sternites are present in all species of Haaniella (although indistinct in most species the muelleri species-group) and Heteropteryx Gray, 1835, hence are a key-feature of the subfamily Heteropteryginae.



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Distribution (Fig. 494): Sumatra, endemic. NE-Sumatra: Prov. Sumatera Utara, Kwalu River [SMTD—type locality]; NE—Sumatra: Prov. Sumatera Utara, Medan, Dolok-Baros [MNHN]; NW—Sumatra: Province Sumatera Utara, Berastagi 1220 m [coll. FH, coll. FSC]. Specimens examined: 10 TABLE 15. Measurements [mm] of Haaniella parva Günther, 1944. LT, ♂ [SMTD]

♂♂ [coll. FH, FSC]

♀♀ [coll. FH, FSC]

Body

34.3

25.0–37.0

45.0–52.0

Head

4.7

3.0–4.5

5.0–6.0

Pronotum

4.4

3.5–4.5

4.0–5.0

Mesonotum

4.8

4.5–5.6

5.0–6.4

Metanotum

4.3

4.5–5.4

–

Median segment

2.0

–

–

Tegmina

6.5

6.0–7.0

9.0–10.0

Profemora

9.5*

9.0–9.6

8.2–10.0

-

9.0–10.0

10.0–10.5

Mesofemora

8.0

7.3–9.0

7.6–9.0

Mesotibiae

8.5

8.6–9.8

8.5–9.6

Metafemora

11.1

8.0–10.2

11.3–13.5

Metatibia

12.6

11.0–12.0

13.0–14.5

Antennae

-

22.5–25.0

21.0–22.5

Protibiae

* according to Günther (1944: 73)

4.1.3 The echinata species-group The three species of the echinata species-group of Haaniella are endemic to Borneo and are sister to the grayii species-group, with which they share the longitudinal median row of spines on the ventral surface of the meso- and metafemora, prominent lateral invaginations of abdominal sternites II–IV, rather flattened head, specialized apex of the epiproct of ♀♀ and setose eggs. The following characters distinguish it from the members of the grayii-group: Anal segment of ♀♀ protuded into a spiniform projection posteromedially; epiproct of ♀♀ shortened and not reaching apex of subgenital plate; subgenital plate of ♀♀ projecting over epiproct, acute apically and at the base with an angular median projection; abdominal tergites II–IV of ♂♂ without an enlarged central pair of spines; lateral margins of anal segment of ♂♂ with a triangular tooth sub-basally. Another characteristic of the echinatagroup are the bright blue or blueish green coxal trochanters and intersegmental membranes between abdominal sternites I–III. There are 26 antennomeres in ♀♀ and 24 in ♂♂. This group and the grayii-group together are the sister-taxon of the muelleri species-group (Fig. 355). The lack of an enlarged central pair of spines on abdominal tergites II–IV of ♂♂ is shared with the muelleri-group and one species has distinctly lemon-shaped eggs with a distinctly conical polar-area and operculum, a character typical for the muelleri-group. The same species, i.e. H. saussurei (Kirby, 1904), also has a strikingly elongate and slender epiproct in ♂♂, a character unique in the echinata-group and otherwise only found in certain members of the muelleri-group. These characters suggest members of the echinata-group are more closely related to the muellerigroup than are the species of the grayii-group. For a comparison and distinction from the other two species-groups of Haaniella see Table 1.


HENNEMANN ET AL.

Haaniella echinata (Redtenbacher, 1906) (Figs. 78, 90, 274–290, 348–349, 379–380, 392, 409–410, 437–440, 445–446, 472–476, 495) [Prickly Haaniella] Heteropteryx echinata Redtenbacher, 1906: 171, pl. 6: 10 (♂) & 11 (♀). LT [by present designation], ♂: Nordborneo, Brunei, Coll. Br. v. W.; Brunei, Borneo, Fruhstorfer; det. Redtenb., Heteropteryx echinata; 19.883 [NHMW, No. 284]; PLT, ♂: Coll. Br. v. W.; Labuan, Burr; det. Redtenb., Heteropteryx echinata; Labuan, B.N. Borneo, ex coll. Swinhoe; 25.062 [NHMW, No. 284]; PLT, ♀: Coll. Br. v. W.; Borneo, Higgins; det. Redtenb. Heteropteryx echinata; 8654 [NHMW, No. 284]; PLT, ♀ (nymph): Coll. Br. v. W., Sarawak (Borneo), Stevens; det. Redtenb. Heteropteryx echinata; 5234 [NHMW, No. 284]; PLT, ♀ (nymph): Labuan, B.N. Borneo, ex coll.: Swinhoe; Collectio Br. v. W., det. Redtenb. Heteropteryx echinata; 25.063 [NHMW, No. 284]; PLT, ♀: N. Borneo Rolle [MNHU]; PLT, ♂: Marapok, Nd. Borneo; Heteropteryx grayi ?; Heyne 901.; 162.; H. echinata Redt., Brunn. Redt. det. [ZIN]. Dohrn, 1910: 403. Giglio-Tos, 1910: 14. Haaniella echinata, Günther, 1932: 313. Rehn, 1938: 368. Günther, 1944: 71. Cocking, 1986: 7. Harman, 1987: 380–382, figs. 1 (♂), 2 (♀) & 3 (egg). Hausleithner, 1991: 219. Chan, 1996: 31. Schulten, 1995: 96, figs. 58 (♀) & 59 (mating pair). Bragg, 1998: 18, figs. 15–17 & 27–28. Brock, 1998: 27. Seow-Choen, 1998: 89, table 1. Lipinski et al., 1999: 96, fig. 44. (REM-study of egg) Bragg, 2001: 80, figs. 15 a–b (egg), 17 a (♂), b (♀), 18 a–c (♀ abdomen), d–e (egg). Seow-Choen, 2000: 152, figs. 3 (♂) & 4 (♀). Zompro, 2005b: 260. Brock, 2007: 50. Dräger, 2011: 56, figs. 10:1a–c (♀ abdomen), 11: 5 (♂ thorax), 33 (♀), 34 (♂), 35 (♀ defensive reaction), 36 (♂), 37: 5a–c (eggs), 38–39 (♀). Seow-Choen, 2016: 407, figs. 1029-1032. Haaniella echinata echinata, Günther, 1944: 71. [not Haaniella saussurei (Redtenbacher, 1906); errorneous synonym by Rehn, 1938: 368—corrected by Bragg, 1998: 19]. [not Haaniella echinata echidna (Rehn, 1938); listed as subspecies by Günther, 1944: 72—corrected by Bragg, 1998: 19]. [not Haaniella echinata saussurei (Redtenbacher, 1906); listed as subspecies by Günther, 1944: 72—corrected by Bragg, 1998: 19]. [not Haaniella echinata scabra (Redtenbacher, 1906); listed as a subspecies by Günther, 1944: 72—corrected by Bragg, 1998: 19].

Material examined (62 ♂♂, 49 ♀♀, 38 nymphs, eggs): BRUNEI: 1 ♀ (penultimate instar): Borneo, Brunei, Lumut District, Sungei Liang Arboretum, 23rd June 1990, Coll. C.L. Chan & Salleh [NHMUK]; 1 ♂, 1 ♀: Borneo, Brunei, Temburong, Kuala Belalong, 19th June 1989, Coll. K.M. Wong [NHMUK]; 1 ♀, 1 ♀ (n3): Borneo, Brunei, Beliat District, Rampayoh Waterfalls Trail, 8 January 1994, Coll. C.L. Chan, Seow-Choen & Maraina Wong [NHMUK]; 1 ♀ (n4): Brunei, Sg. Liang, 30/5/89, Coll: Chan & Wong [NHMUK]; 3 ♂♂, 3 ♀♀: Borneo, Brunei, Temburong District, Kuala Belalong, Badas, Seria (swamp forest), leg. F. Hennemann & M. Herbert, VIII.1994 [coll. , No’s FH 0230-8 to 13]; 5 ♂♂, 6 ♀♀, eggs: ex Zucht F. Hennemann, 1995-1998, Herkunft: Brunei, Temburong District, Kuala Belalong, Badas (F1) [coll. FH, No’s 023014 to 24, & E]; 1 ♂, 1 ♀: ex Zucht H. Dräger 2011, Herklunft: Brunei [coll. FH, No. 0230-25]; 1 ♂, 1 ♀: ex Zucht O. Conle, 1999, Brunei [coll. OC]. SABAH: 1 ♀, 1 ♂, 1 ♂ (penultimate instar): Borneo: Sabah, Ulu Padas, Long Pasia Ladang area, 950 m, Coll. Pengiran Balang, 15–18 August 2000 [NHMUK]; 1 ♂: Borneo: Sabah, Ulu Padas, Long Pilang Campsite, 1000 m, Riverine forest (Loc. name: Lemut), Coll. Justine Vaz, 27 Nov 2000 [NHMUK]; 1 ♂: Borneo: Sabah, Long Pasia, 3 km NW of Long Pasia Village, 3700 ft a.s.l., 8 November 1997, Coll. Leopold Madani [NHMUK]; 1 ♂, 1 ♀: Borneo, Sabah, Penampang District, Ulu Moyog c. 650 m, 29th April 1989, Coll. Natives [NHMUK]; 1 ♀: Moyog, 22/IV/



81

89, Coll: Native [NHMUK]; 1 ♂ (n3): Borneo, Sabah, Gunung Trus Madi, c. 850 m, 17th May 1991, Coll. Tan Fui Lian; Logged over forest ridge, on shrubby vegetation [NHMUK]; 1 ♂, 2 ♀♀: Borneo, Sabah, Crocker Range, Tenom, Sungei Purulon, 300 m, 4th September 1990, Coll. Tan Fui Lian [NHMUK]; 1 ♂: Kallang, Tenom, Sabah (N. Borneo), 25.VI.1986, Coll. C.L. Chan & J. Huisman [NHMUK]; 2 ♀♀ (n3): Borneo, Sabah, Tenom, Kallang Water Fall, 25th June 1986, Coll. Chan & Huisman [NHMUK]; 1 ♂ (penultimate instar): Borneo, Sabah, Crocker Range, Ridge above Sungei Purulon c. 350 m, 28th June 1989, Coll. Tan Fui Lian [NHMUK]; 1 ♀ (n5): Sepilok, Sabah (N. Borneo), 8 IX 1982, Coll. C.L. Chan; Haaniella echinata (Redt.) ♀ nymph [NHMUK]; 2 ♂♂, 1 ♀ (n3): Sepilok, Sabah (N.Borneo), 8.X.1982, Coll. C.L. Chan [NHMUK]; 1 ♀, 2 ♂♂, 3 eggs: Sepilok, Sabah (N. Borneo), 13 June 1982, Coll. C.L. Chan & S. Nagai [NHMUK]; 1 ♂ (penultimate instar), 1 ♀ (n5): Borneo, Sabah, Sepilok Forest R., Orang Utan R. Centre, 8th October 1981, Coll. C.L. Chan [NHMUK]; 1 ♂, 1 ♂ (penultimate instar), 1 ♀ (n4), 1 ♂ (n5): Borneo, Sabah, Sepilok Forest R., Orang Utan R. Centre, 8th October 1982, Coll. C.L. Chan [NHMUK]; 1 ♀ (n5): Sepilok Sandakan, (N. Sabah), 9 July 1983, Coll. C.L. Chan & Ronnie Young [NHMUK]; 2 ♂♂: Borneo, Sabah, Sandakan District, Sepilok Forest Reserve, 13th June 1988, Coll. C.L. Chan & R. Young [NHMUK]; 2 ♀♀: Ulu Dusun, Sabah (N. Borneo), 18 May 1984, Coll. C.L. Chan & Wong [NHMUK]; 6 ♂♂: Ulu Dusun, Sabah (N. Borneo), 18 May 1984, Coll. C.L. Chan & W. Wong [NHMUK]; 1 ♂: Poring, Sabah (N. Borneo), 2.VII.1983, Coll. C.L. Chan & Michael Chan [NHMUK]; 1 ♂ (n5): Poring Hot Spring, Sabah (N. Borneo), 2 July 1983, Coll. C.L. Chan & Michael Chan [NHMUK]; 1 ♀: Poring 31/7/90 [NHMUK]; 1 ♀: Ppring Hot Springs, 31/7/90, Chan, Bragg & Inglis [NHMUK]; 1 ♂: Borneo, Sabah, Keningau, Apin Apin c. 2000' a.s.l., January 1993, Coll. A. Lamb & J. Mood [NHMUK]; 1 ♂: Mt. Silam 2200', Laha Datu, Sabah (N. Borneo), April 1982, Coll. W. Mrazek [NHMUK]; 1 ♀ (n3): Batu Putih, Kinabatangan, Sabah; P.S. Shim Coll. 20.IX.1981 [NHMUK]; 1 ♀: Borneo, Sabah, Sipitang District, Mendolong, SFI area, Secondary cum Acacia plantation, 800 m, June 1989, Coll. F.L. Tan & R. Inger [NHMUK]; 2 ♂♂: N-Sabah, Mt. Kinabalu, Poring Hot Springs 480 m, leg. F. Hennemann & O. Conle, 4.–8.VIII.1996 [coll. FH, No’s 0230-25 & 26]; 10 eggs: ex Zucht T. Guttmann, 1996, Herkunft: Sabah, Poring Hot Springs [coll. FH, No. 0230-ED]; 1 ♂, 1 ♀: B.N. Borneo, nr. Kinabalu, Kabayan, 600m, 9.5.1929; Haaniella echinata (Redt.) det. Klante [RMNH]; 1 ♀ (nymph): B.N. Borneo, Bettotan, nr. Sandakan, 21.8.1927¸ Haaniella echinata (Redt.) det. Klante [RMNH]; 1 ♂: Sabah, Beaufort, 105km S of Long Pa Sia and surroundings, 4°24N 115°43´E, Nov. Dec. 1987, leg. Sigar Bin Baru; Haaniella echinata (Redtenbacher) det. Bragg, 1993 [RMNH]; 3 ♂♂, 3 ♀♀: leg FSC, Sepilok Orang Utan Rehabiliation Centre [coll. FSC]; 1 ♀: Sabah, 2004 [coll. OC, No. 0031-2]. SARAWAK: 2 ♀♀, 2 nymphs: N. Borneo, Exp. 1912, Mohari; det. Klante [RMNH]; 1 ♀, 5 ♂♂: Sarawak, Batu Niah, XI.– XII.1980, A. Harman [NHMUK, 1982-387]; 2 ♂♂: Sarawak, Ulu Dusun, 1977, A. Harman; „Haaniella echinata“ det. C.L.Chan, 1989 [NHMUK, 1982-387]; 1 ♀ (nymph): Sarawak, 4th, Division, Gunung Mulu NP, Camp 6, Hidden Valley, 20.–28.III.1978, P. Chapman [NHMUK, 1982-206]; 1 ♀: 87-14, Rejang River, Sarawak, Pres. By Brooks Lous. Esq [NHMUK]; 1 ♂ (nymph): Sarawak, 87-14 [NHMUK]; 2 ♀♀, 1 ♂, 1 ♂ (penultimate instar), 1 ♀ (penultimate instar), 1 ♂ (n3), 4 ♀♀ (n3): Borneo, Sarawak, Mulu National Park, Gunung Mulu Trail, 15–16 March 1994, Coll. C.L. Chan & Poon [NHMUK]; 1 ♀: N. Borneo, ex Coll. Fruhstorfer [SMNS]; 1 ♂, 1 ♀: Museum Paris, Nhatrang, A. Kempf, 1913, Heteropteryx Erringtoniae Redtb., det. L. Chopard & L. Berland [MNHN]; 1 ♀: Museum Paris, Borneo, Mt. Bahan, Pfeiffer III.57 [MNHN]; 3 ♂♂, 2 ♀♀, 2 ♀♀ (nymphs): OSarawak, Batu Niah NP, leg. F. Hennemann & O. Conle 2.–4.VIII.1996 [coll. FH, No’s 0230-1 to 7]; 5 ♂♂, 5 ♀♀, 1 ♀ (nymph): O-Sarawak, Batu Niah NP, 2.–4.VIII.1996, leg. Conle & Hennemann [coll. OC]; 1 ♀, 2 ♀♀ (nymphs): Borneo, N-Sabah, Tawau, 5.2013, leg. Berezin [coll. OC]; 1 ♂: Ex Zucht B. Kneubühler 2013, Herkunft: Sarawak, Niah N.P. Und Lambir Hills N.P., leg. Francis Seow-Choen (F1) [coll. OC]; 1 ♀: Borneo, Sarawak; det. Ebner 1925 Heteropteryx echinata [NHMW, No. 284]; 3 ♂♂, 3 ♀♀: leg. FSC Niah Caves NP [coll. FSC]. KALIMANTAN: 1 ♀, 1 nymph: H.C. Siebers, Midden-O-Borneo, 14.9.1925; Heteropteryx echinata Redt. det. Karny; Haaniella echinata (Redt.) det. Klante [RMNH]; 1 ♀ (nymph): M. O. Borneo, Exped., Oeloe Kelai, Kapt. Buys, 9.10.1925; Haaniella echinata Redt. det. K. Günther [RMNH]; 1 ♂: E-Borneo, Kalimantan, Timur, Mount Beratus IV.2009 [coll. OC., No. 0031-1]. “BORNEO”: 1 ♀: Borneo, Juli 1912; Haaniella echinata Redt. det. K. Günther; Haaniella echinata (Redtenbacher)det. Bragg


HENNEMANN ET AL.

[RMNH]. Diagnosis: Very closely related and the sister-taxon of H. scabra (Redtenbacher, 1906). The enlarged anterolateral mesonotals, distinct posterior pair of spines on the pronotum as well the gradually tapered abdominal segments II–V and short, rounded epiproct of ♂♂ are shared with this species. Both sexes are however frequently distinguished from H. parva by: the distinctly larger size; the anterior coronals being distinctly smaller than the remaining; presence of supra-orbitals, considerably more elongate antennomeres, spinulose abdominal tergites II– VI (scabrous or rugulose in scabra); distinctly shorter spination of abdominal sternites II–VII and comparatively shorter tegmina, which at best reach to the posterior margin of abdominal tergum II. The eggs differ from those of H. scabra by the much larger dimensions, relatively more slender micropylar plate and plain brown colouration. From H. saussurei (Kirby, 1904) it differs by: the unequally sized spines on the lateral margins of the mesonotum (anterior lateral mesonotals strongly enlarged), more prominent posterior pronotals of both sexes, as well as the gradually tapered abdominal segments II–VI and short epiproct of ♂♂. Males furthermore differ by the stockier habitus and different colouration of the tegmina. The eggs at once differ from those of H. saussurei by the barrel-like shape and lack the conical polar-area and operculum typical for H. saussurei. Etymology: The name (echinus gr. = sea urchin) refers to the spiny body and legs of this species. Description: ♀♀ (Figs. 274–277). Large (body length 91.0–135.0 mm) and massive species with prominent antero-lateral mesonotals and four distinct posteromedial mesonotals between the bases of the tegmina. Colour very variable and ranging from dull ochre (Fig. 476) over various shades of brown to almost black, either plain or prettily mottled. Occasionally pale ochre specimens occur, which have the head, pleurae and ventral body surface dark brown (Fig. 472), others may have a distinct and contrasting pale cream to white marking on the mesonotum and abdominal tergum VI (Fig. 475). Mesonotum often with the anterolateral angles pale cream to white. Dorsal armature of head and thorax dull green basally, then becoming dull red and the tips black. Abdominal tergites VII– VII usually with several paired black markings or bold spots. Head with a ± defined dark postocular stripe. Tegmina ranging from plain dull ochre over brown to almost black, sometimes with a bold black spot or marking roughly in centre (Fig. 275) and often with the basal portion pale cream to whitish; these pale portions forming a roughly heart-shaped markiong when the tegmina are closed (Fig. 475). More rarely the pale basal portion posteriorly bordered by a diagonal white stripe (Fig. 474). Legs either plain or with with pale mottling in the apical half. Interior surface of metafemora more or less distinctly pale red or at least with a slight reddish hue; metacoxae dull red interiorly. Coxal trochanters of metacoxae and membranes between abdominal sternites I–III bright blue (Fig. 348). Head about 1.2x longer than wide, parallel-sided with the vertex very gently convex in posterior portion; armed with 12 spines (Figs. 282–283). Vertex with four distinct and almost equally sized median coronals which formthe four corners of a square. Anteriorly with a much smaller pair of supra-occipitals and a minute pair of supra-antennals (Fig. 282). Posterior margin with two prominent lateral coronals, which are roughly equal in length but somewhat stronger than the anterior and posterior coronals. Behind the eyes with two fairly distinct supraorbitals, the anterior one noticeably smaller than the posterior one. Usually with one or two minute gulars on cheeks. Antennae consisting of 26 segments and ± reaching half way along abdominal tergum IV. Pronotum slightly longer than head, a little longer than wide and trapezoidal with posterior margin some 1.4x wider than anterior margin; the lateral margins concave, indented medially and set with 4–6 minute spines. Antero and postero-lateral pronotals distinct and near posterior margin with a somewhat less pronounced pair of inter-posterior pronotals. Just before transverse median sulcus with a prominent and strong pair of pre-median pronotal spines (Fig. 283). Mesothorax gently and gradually widening towards the posterior with posterior portion some 1.2x wider than anterior margin. Mesonotum slightly narrowing towards the posterior and only about 1.2x longer than pronotum. Lateral margins with 2–6 fairly small and slender, unequally sized spines and a very prominent and long antero-lateral mesonotal. Disc with a pair of distinct anterior-mesonotals, which are slightly smaller to almost equal in size to the antero-lateral mesonotals; otherwise only set with a few minute paired spinules in the median portion (Fig. 282). Between bases of tegmina with four distinct posteromedial mesonotals; the posterior pair usually somewhat longer than the anterior pair (Fig. 283). Tegminal distinct. Mesopleurae with a strong supra-coxal spine and 6–7 unequally sized but fairly distinct, slender laterals along lower margin; the most anterior one larger than the remaining (Fig. 282). Metapleurae with lateral margin rounded and armed with six spines, the anterior five spines very long and sometimes gently backcurving, the posterior spine considerably smaller; supra-coxal very prominent and the largest of all the thoracic spines. Sensory area of profurcasternum moderately distinct, oval in



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outline and with a longitudinal furrow medially; segment otherwise with a very distant pair of small spines (Fig. 78). Mesosternum with a lateral row of three stout but strong spines, which increase in size towards the posterior; usually a second row of 2–3 much smaller spines interiorly. Metasternum with 4–5 short but fairly strong spines on each side of the median line; the outermost strongest. Tegmina ± reaching to posterior margin of abdominal tergum II, oval with the apical portion increasingly narrowed; the radial vein strongly raised a forming a prominent longitudinal keel in basal portion of tegmen. Abdominal tergites II–III widening, IV widest and V–VII narrowing, all transverse with lateral margins deflexed and either straight to gently rounded and armed with 7–8 slender, unequally sized spines. II smooth, III–VII with a few very minute paired spinules near median line and a transverse row of six minute spines close to posterior margin; these indistinct or obsolete on III and becoming increasingly pronounced towards VII. Sternites II–VI with a transverse row of six fairly distinct spines close to posterior margin and 1–2 pairs of spines near anterior margin, V–VI only with eight spines. Praeopercular organ formed by a pair of short but very prominent, distant outward pointing post-median spines on sternum VII (Fig. 287). Tergum VIII with about three very minute spines on lateral margins. Anal segment with a blunt longitudinal keel, which is posteromedially protuded into a blunt spiniform projection (Fig. 286). Epiproct shorther than anal segment with posterior margin angular, notched medially and armed with 2–6 minute teeth (Figs. 285–286). Cerci small and compressed laterally. Subgenital plate gently downcurving in the apical portion, the apex bluntly triangular and projecting considerably over apex of epiproct (Fig. 286). All legs strong and with all carinae spinose or dentate, the dorsal carinae of the tibiae only minutely denticuate. Metafemora reaching to abdominal segment VI and metatibiae projecting somewhat beyond apex of abdomen. Ventral surface of femora with a longitudinal median row of small spines. Two ventral carinae of metatibiae armed with about eight large spines, which strongly increase in size and become increasingly incurving towards the apex of tibia; the most apical spine of the posteroventral carina largest (Fig. 379). ♂♂ (Figs. 278–281). Medium sized to large (body length 68.0–94.0 mm) and fairly stocky with prominent antero-lateral mesonotals and four very large posteromedial mesonotals between the bases of the tegmina. Colouration ranging from pale to very dark brown (usually darkening with age), rarely with a slight dull greenish hue. Sometimes with a large whitish marking in anterior portion of mesonotum and occasionally with a bold cream to ochre marking on abdominal tergum VI. More rarely pale specimens may have a faint dark brown longitudinal median stripe running along most of the dorsal body surface. Head with a ± defined dark postocular stripe. Armature of thorax green and dull red towards the black tips. Tegmina ranging from plain dull ochre over brown to almost black, often with the basal portion pale cream to white (these pale portions forming a roughly heart-shaped marking when the tegmina are closed, Figs. 278–279) or with a pale cream to white diagonal stripe (Figs. 280– 281). Legs either plain or with with pale mottling in the apical half. Interior surface of metafemora more or less distinctly pale red or at least with a slight reddish hue; metacoxae dull red interiorly. Coxal trochanters of metacoxae and membranes between abdominal sternites I–III bright blue (Figs. 349, 392); coxal trochanters of proand mesocoxae bright green. Shape and armature of head as in ♀♀, but armature more prominent with all spines comparatively longer and more slender (Fig. 284). Antennae long, consisting of 24 segments and reaching to abdominal segment V or VI. Pronotum generally as in ♀♀ with but the pre-median pronotals in particular longer and considerably more prominent. Mesothorax moderately elongate, some 1.6x longer than pronotum, roughly parallel-sided with the posterior portion somewhat widened. Mesonotum gradually narrowing towards the posterior and about 1.6x longer than width of anterior margin; armature as in ♀♀ with all spines comparatively longer and more slender and the four posteromedial mesonotals in particular much larger and very prominent (Fig. 284). Tegminal spine distinct. Mesopleurae with armature as in ♀♀ but the spines averaging longer and the supra-coxal very prominent. Sensory area of profurcasternum as in ♀♀. Armature of meso- and metasternum as in ♀♀, but all spines considerably more pronounced. Abdominal sternum I with a very prominent pair of spines close to posterior margin (Fig. 90). Tegmina ± reaching half way along abdominal tergum II, oval in shape with the apical portion increasingly narrowed; the radial vein strongly raised a forming a prominent longitudinal keel in basal portion of tegmen. Abdominal segments II–V gently and gradually narrowing, VI narrowest segment and parallel-sided, VII and VIII slightly widening. II, III and VII somewhat wider than long, IV–VI roughly as wide as long. Lateral margins of tergites II–V with about seven small and short, unequally sized spines, VI and VII only with five small spines and only 2–4 minute spines present on lateral margins of VIII. Dorsal surface of II–VII set with a variable number of minute spines or spiniform tubercles mostly arranged as two longitudinal rows in median portion and a transverse row close to posterior margin, VIII–X unarmed but VIII occasionally with a single pair of minute spines close to


HENNEMANN ET AL.

anterior margin. Tergum IX with posterolateral angles protuded into a blunt, posteriorly directed spiniform process. Sternites II–VI with eight spines, V–VI only with six spines and VII unarmed except for a minute pair of spines close to posterior margin (Fig. 290). The lateral invaginations of II–IV very distinct (Fig. 90). Anal segment with lateral margins sub-basally deflexed into a blunt, triangular, tooth-like lobe (Fig. 288), the posterior margin roundly excavated. Epiproct short, roundly triangular to semicircular in shape and slightly projecting over posterior margin of anal segment (Fig. 289). Cerci small, club-like and distinctly compressed laterally in basal portion. Vomer broad, triangular with the central portion somewhat concave and the fairly short terminal hook ± decidedly curved dextrally; ochre to dull orange with the basal portion brown (Figs. 409–410). Poculum projecting somewhat over posterior margin of tergum IX with posterior margin broad and roundly angular (Fig. 290). Legs all fairly long but stocky, mesofemora almost as long as pro- and mesonotum combined, metafemora reaching about half way along abdominal segment VII and metatibiae projecting distinctly beyond apex of abdomen. Armature as in ♀♀ but the ventral spines of the metatibiae comparatively smaller and less curved (Fig. 380). Nymphs: The sex of nymphs can already be defined when they are newly hatched, ♀♀ already having a slightly elongated epiproct which is missing in ♂♂. In general, the premature stades are more socky and robust than the adult insects and the body armature is averaging more pronounced. Variability: In accordance to its wide distribution H. echinata is remarkably variable in size, length and shape of the spination and colouration. It has developed several regional variations, and even the eggs show differences between certain localities. Eggs laid by cultured specimens originating from Poring Hot Springs (Sabah, Mt. Kinabalu) are strikingly different from those laid by specimens from Batu Niah NP (Sarawak) or Brunei, those from the latter two localities being virtually identical. The two main varieties of the eggs are described below and illustrated (Figs. 437–440). Females range in colour from dull ochre over dull brown to almost black but usually tend to darken with rising age. They are either plain in colour (e.g. Figs. 274–276) or attractively mottled with pale ochre, white and black (Fig. 475). The mesonotum often bears a bright white spot at the antero-lateral corner (Fig. 283) and occasionally there are paired dull black markings and/or black speckles on abdominal tergites VII–IX (Fig. 275). More rarely specimens occur that have the anterior portion of the mesonotum and abdominal tergum VI contrasting white (Fig. 475). The tegmina range from plain dull ochre to dark brown (e.g. Figs. 276–277) and often bear a black central spot or marking (Fig. 275). Some specimens have a large pale marking in the basal portion which produces a roughly heart-shaped marking when the tegmina are closed (Fig. 475) and more rarely both colour patterns occur simultaneously. Sometimes specimens may even have a whitish diagonal stripe across the tegmina (Fig. 474), which is more frequent in ♂♂. Females from Batu Niah have the lateral margins of abdominal tergites IV–VI more strongly dilated and set with relatively larger spines than e.g. specimens from Brunei. A ♀ from an unspecified locality in NHMW has all the body and leg armature much more strongly developed than all other examined material (Fig. 274). Males mostly show variability in the colouration of the tegmina. These are either plain brown in colour or show various different colourations. Frequently there is a large, roughly heart-shaped pale ochre to white marking similar to that occuring in ♀♀ (Figs. 278–279); sometimes this is reduced to either a whitish diagonal stripe (Figs. 280–281) or a weakly defined central white marking. Slight variation is seen in the shape of the vomer of ♂♂, which has the apex somewhat more curved to the right and the ventral surface more depressed in examples from Poring Hot Springs (compare Figs. 409 & 410). An interesting colouration is seen in specimens from Danum Valley Conservation Area in eastern Sabah and Tenom in western Sabah. These have all coxal trochanters and the intersegmental membranes between abdominal sternites I–III dull pink to red instead of the usual blue colour seen in specimens from all other localities. Eggs: Eggs of H. echinata show considerable regional variation. Eggs from Brunei (Badas) and Sabah (Mount Kinabalu, Poring Hot Springs) differ strikingly in the shape of the micropylar plate and operculum as well as the surface texture, hence these two varieties are here described separately. Brunei (Figs 439–440, 445): Very large, barrel shaped, polar-area rounded; 1.4x longer than wide. Entire surface densely setose, the setae obscuring most of the capsule and micropylar details, fairly long and arranged in irregular clusters; somewhat longer on operculum.. Micropylar plate basically X-shaped; anterior extensions at an angle of about 65° to each other with the apical portion gently incurving; posterior extensions with the apex strongly expanded towards the anterior, producing a roundly hooked shape. Outer margin of plate slightly raised. Microyplar cup a shallow impression with a rounded tubercle posteriorly. Operculum circular and slightly convex. General colour plain mid to dark brown. REVISION OF HETEROPTERYGINAE


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Measurements [mm]: length incl. operculum 10.7–11.3, length 9.8–10.5, width 6.9–7.8, height 7.8–8.2. Sabah (Figs 437–438, 446): Very large, barrel-shaped and slightly narrowing towards polar end, polar-area rounded; 1.3x longer than wide. Entire surface irregularly setose, the setae obscuring much of the capsule and micropylar details, uneven in length and arranged in clusters, which form an irregular net-like texture; setae longest in anterior half of capsule and along outer margin of micropylar plate. Spaces between the clusters of setae very minutely granulose. Micropylar plate basically X-shaped; anterior extensions at an angle of 90° to each other with the apex strongly expanded and club-shaped; posterior extensions with the apex strongly expanded towards the anterior, producing a roundly hooked shape. Outer margin of plate raised and covered with a rim of long setae. Micropylar cup a shallow impression and a distinct, black cup-shaped structure posteriorly. Operculum circular and conically elevated in centre. Colouration mid brown with the setae greyish brown. Measurements [mm]: length incl. operculum 9.2–10.0, length 8.7–9.2, width 6.2–6.8, height 6.2–7.1. Comments: This species was originally described from a fairly large series of specimens deposited in several European museums from Brunei, Sabah, Sarawak and the island of Labuan but so far no lectotype has been designated. Hence, a ♂ from Brunei in NHMW is here selected as the lectotype in order to guarantee stability of the name, although all type-specimens are conspecific. Several papers have since dealt with the captive breeding, biology and behaviour of H. echinata (e.g. Cocking, 1986; Harman, 1987; Schulten, 1995). Lipinski et al. (1999: 96) have provided a REM-study of the egg. H. echinata is a widespread lowland species, which is quite common throughout much of Borneo (→ see list of localities below) and has developed several regional varieties. In accordance to its wide distribution, H. echinata is similarly variable as H. grayii. Bragg (2001: 85) reported the insects to be eaten by the local Iban people and Chan (1996: 31) recorded monitor lizards (Varanus sp.) as predators of H. echinata. Culture stock has been imported into Europe on many occasions and from different localities since the early 1980’s. The original stock originated from a pair collected in 1979 by Allan J.E. Harman (Essex, England) in Ulu Dusun, Sabah and was included on the Phasmid Study Group culture list as culture No. 26. Subsequently, stock has been imported and successfully reared from Sarawak (Batu Niah National Park and Lambir National Park), Brunei (Temburong District, Kuala Belalong) and Sabah (Mount Kinabalu, Poring Hot Springs). H. echinata is fairly easy to rear in the standard conditions for this genus and accepts a variety of alternative foodplants, including bramble (Rubus fruticosus, Rosaceae), raspberry (Rubus idaeus, Rosaceae), roses (Rosa spp., Rosaceae), pyracantha (Pyracantha coccinea, Rosaceae), flowering currant (Ribes spp., Rosaceae), hawthorn (Crataegus monogyna, Rosaceae), ivy (Hedera helix, Araliaceae), oaks (Quercus spp., Fagáceae), silver birch (Betula pendula, Betulaceae) and salal (Gaultheria shallon, Ericaceae). In the wild H.echinata is known to be fairly polyphagous, feeding on a variety of different plants (→ 7.6). Distribution (Fig. 495): Widely distributed throughout Borneo, endemic. Sabah: Mount Kinabalu, Poring Hot Springs 480–760 m [NHMUK, coll. FH, coll. OC, coll. PEB]; Kabayau nr. Kinabalu 600 ft. [RMNH]; Kinabatangan District, Sepilok [coll. PEB]; Kinabatangan District, Batu Putih [NHMUK]; Kinabatangan District, Sepilok, Orang Utan Rehabilitation Centre [NHMUK, coll. FSC]; Kinabatangan District, Batu Putih [NHMUK]; Long Pasia ladang area, Ulu Dusun, 950 m [NHMUK]; Bettotan nr. Sandakan [RMNH]; Marapok [ZIN]; Beaufort, 105km S of Long Pa Sia [RMNH]; Bungal [NHMUK]; Penampang District, Ulu Moyog [NHMUK]; Crocker Range, Tenom District, Sungai Purlon, ca. 350 m [NHMUK]; Crocker Range, Gunung Trus Madi, 850 m [NHMUK]; Tenom, Kallang Waterfall [NHMUK]; Mount Silam, Mahad Datu 2200 ft. [coll. CLC]; Keningau, Apin Apin [NHMUK]; Laha Datu, Mount Silam, 670 m [NHMUK]; Sipitang District, Mendolog, SFI area 800 m [NHMUK]; Sandakan [NHMUK, Rehn, 1938: 368]; Ulu Padas, Long Pilang Campsite, 1000 m [NHMUK]; 3km NW of Long Pasia Village, 1100 m [NHMUK]; Tawau Hill Park [coll. OC]; Danum Valley Conservation Area, edge of Danum valley along the neighbouring oil palm estate [observed by the fourt author]. Brunei: “Brunei” [NHMW]; Temburong District, Kuala Belalong, Badas, swamp forest [coll. FH, coll. PEB, coll. OC]; Temburong District, Kuala Belalong 100 m [BMKP, NHMUK, coll. PEB]; Teraja, waterfall trail [coll. PEB]; Beliat District, Rampayah waterfall trail [NHMUK]; Sungai Liang Forest Reserve [NHMUK]; Lumut District, Sungai Liang Arboretum [NHMUK]. Sarawak: Rajang River [NHMUK]; 20 km upstream of Kapit [coll. PEB]; Pelagus Logging Camp, 40 km from Kapit [coll. PEB]; Kapit area [coll. PEB]; Mount Dulit, junction of Tinjar and Lejok Rivers [NHMUK]; Lio Matah [SMSM]; Batu Niah National Park [NHMUK, SFDK, coll. FH, coll. FSC, coll. PEB]; Mount Bahan [MNHN]; Nhatrang [MNHN]; Mulu National Park, Gunung Mulu trail [NHMUK]; Mulu National Park, near Park Headquarters [photo by Paul Bertner]; Lambir National Park [coll. FSC]. Kalimantan: “Midden O-


HENNEMANN ET AL.

Borneo” [RMNH]; Balekpapan Road [RMNH]; Ulu Kelai, upper River Kelai “Oeloe Kelai” [RMNH]; Samarinda [MIZT]; Selatan, Mount Bakayan [coll. OC]; Timur, Mount Banyutawar [coll. OC]; Timur. Mount Beratus [coll. OC]. Labuan Island [ANSP, NHMW]. The fourth author found this species to be particularly plentyful at Tawau Hill Park in Sabah. Specimens examined: 156 TABLE 16. Measurements [mm] of Haaniella echinata (Redtenbacher, 1906) ♂♂

♀♀

71.5–87.0 68.0–94.0

100.0–117.0 91.0–135.0*

Head

7.4–9.0

10.5–12.0

Pronotum

6.8–8.0

10.7–13.2

Mesonotum

9.8–11.2

13.5–17.0


10.8–12.6

9.3–15.5

Tegmina

15.0–17.2

21.4–27.0

Profemora

19.5–23.0

20.6–23.0

Protibiae

21.2–25.0

25.0–28.0

Mesofemora

16.0–17.4

18.3–21.0

Mesotibiae

18.0–21.0

22.5–23.7

Metafemora

25.0–27.3

27.3–31.0

Metatibiae

27.3–31.0

33.2–36.0

Antennae

60.0–62.5

56.5–68.5


* Bragg (1998: 92 & 2001 : 76) listed a 84.0 mm ♀ from OXUM, which is here excluded.

Haaniella saussurei Kirby, 1904 (Figs. 83, 85, 291–305, 364, 383–384, 431–432, 477–481, 496) [Saussure’s Haaniella] Haaniella Saussurei Kirby, 1904b: 397. HT, ♀: Coll Guérin; Museé de Genève No. 127; Heteropteryx saussurei Redtb., 127. [MHNG]. (Description of new species based on Saussure, 1869: 310 - not Westwood, 1859) Heteropteryx grayi var., Saussure, 1869: 310. Haaniella saussurei, Günther, 1932: 313. Bragg, 1991: 76, fig. 1 (spermatophore) Bragg, 1992: 297, fig. 2 (egg). Bragg, 1998: 23, figs. 21–23 (♀ abdomen) & 35–36 (egg). Bragg, 2001: 90, figs. 15 i–j (egg), 21 a–b (♀ abdomen), d–e (egg), pl. 3: b (♀). Dräger, 2011: 55, figs. 10: 2a–c (♀ abdomen), 11: 4 (♂ thorax), 29 (♀ defensive reaction), 30 (♀), 31 (♂, ♀♀), 32 (mating couple), 37: 3a–b (egg). Seow-Choen, 2016: 409, figs. 1038-1042. Heteropteryx saussurei Redtenbacher, 1906: 171. LT [by present designation], ♀: Coll Guérin; Museé de Genève No. 127; Heteropteryx saussurei Redtb., 127. [MHNG]; PLT, ♀: Borneo, Sarawak, 1865 66, Coll. S. Doria; Best. Verz. No. 84 [MCSN]; HNHM-material destroyed. [Not: PLT, ♀: Bruner, Borneo, 1871; Mus. Caes. Vind. Borneo Brunner ded.; det. Redtenb. Heteropteryx saussuei, Heteropteryx saussurei Redt. [NHMW, No. 285] = H. echinata (Redtenbacher, 1906)]. (Objective synonym & homonym, synonymised by Bragg, 1992: 297) Haaniella echinata saussurei, Günther, 1944: 72. Haaniella echidna Rehn, 1938: 369, pl. 14: 1 (♂). HT, ♂: Sarawak, Sadong, H.W. Smith leg.; Hebard Type, No. 1294 [ANSP]; PT, ♂: same data as holotype [MCZC]. (Synonymised by Bragg, 1992: 297) Haaniella echinata echidna, Günther, 1944: 72. Heteropteryx grayii Westwood, 1859: 82, pl. pl. 30: 2 (♂) & 3 (♀) (in part—only PLT's: 2 ♂♂, 1 ♀: Borneo [UMO, No. 529]; 1 ♂: Borneo, 58–66; BMNH(E) #845210 [NHMUK];



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Material examined (14 ♂♂, 16 ♀♀): SARAWAK: 5 ♂♂, 3 ♀♀, 7 eggs: ex Zucht F. Hennemann, 1997/1998, Herkunft: NW-Sarawak, Bako NP [coll. FH, No’s 02311 to 8 & E]; 2 ♀♀: ex Zucht H. Dräger 2011–2013, Herkunft: NW-Sarawak, Tarum nr. Debak [coll. FH, No’s 0231-9 to 11]; 1 ♂, 1 ♀: Ex Zucht O. Conle, 2006, Herkunft: Borneo, NW-Sarawak, Bako N.P. (PSG No. 177) [coll. OC]; 3 ♂♂, 4 ♀♀: Ex Zucht O. Conle, 2006, Herkunft: Borneo, NW-Sarawak, Tarum nr. Debak (PSG No. 177) [coll. OC]; 3 ♂♂, 3 ♀♀: Batang Ai National Park, leg. FSC [coll. FSC]; 2 ♂♂, 3 ♀♀: Bako National Park, leg. FSC [coll. FSC]. Diagnosis: Related to H. echinata (Redtenbacher, 1906) and H. scabra (Redtenbacher, 1906) and here regarded as the sister-taxon of these two species, with which it shares the typical features of the echinata speciesgroup, e.g. the short epiproct and long apically pointed subgenital plate of ♀♀ as well as the sub-basal tooth on the lateral margins of the anal segment of ♂♂. H. saussurei however differs from both these species by: the equally sized and fairly small spines along the lateral margins of the mesonotum; not enlarged antero-lateral mesonotals and very small pair of inter-posterior pronotals of both sexes, very small posteromedial mesonotals of ♀♀, as well as the more slender and elongate body, almost parallel-sided abdominal segments II–VI (gradually narrowing in echinata and scabra) and strongly elongated, slender epiproct of ♂♂ (Fig. 297). The eggs are readily distinguished from all members of the echinata and grayii species-groups by their lemon-like shape, having the polar-area and operculum distinctly conical as well as the typical and complex shape of the micropylar plate (Figs. 431–432). The lemon-shaped eggs with the strongly reduced setae resemble those of the muelleri species-group and also the slender, elongated epiproct of ♂♂ is otherwise only found in members of that species-group. Other features indicate close relation to the grayii species-group, i.e. ♂♂ at first glance strongly resembling those of H. grayii (Westwood, 1859) with which they share the contrasting green spines of the body, bright green coxa-trochanter, red interior surfaces of the metacoxae and distinctive colouration of the tegmina. Etymology: Named in honour of the well-known Swiss zoologist and mineralogist Henri Louis Frédéric de Saussure (1829–1905), who first described this species as a variation of H. grayii (Westwood) in 1869. Description: ♀♀ (Figs. 291–293). Large (body length 104.0–132.0 mm) and massive species with fairly reduced and indistinct cephalic and thoracic armature. Colour very variable and ranging from dull ochre (Fig. 293) over various shades of brown to almost black, either plain (e.g. Fig. 477) or prettily mottled (Fig. 480). Often pale ochre specimens occur, which have the head, posterior portions of the pro- and mesonotum, pleurae and ventral body surface dark brown (Figs. 478, 481). More rarely specimens may have a faint dark longitudinal line over most of the dorsal body surface. Dorsal armature of head and thorax dull green with the tips black. Abdominal tergum VIII occasionally with a black lateral marking (Fig. 479). Head with a ± defined dark postocular stripe. Tegmina ranging from plain dull ochre over brown to almost black, sometimes with a bold black spot or marking roughly in centre (Fig. 479) and often with the basal portion pale cream to whitish; these pale portions forming a roughly heart-shaped marking when the tegmina are closed (Figs. 291–293, 478–481). Legs either plain or with with pale mottling in the apical half. Interior surface of metafemora more or less distinctly pale red or at least with a slight reddish hue; metacoxae dull red interiorly. All coxal trochanters greyish blue (Fig. 393). Head about 1.2x longer than wide, very slightly narrowed towards the posterior with the vertex very gently convex in posterior portion; armed with 12 spines (Figs. 304–305). Vertex with four distinct and almost equally sized coronals which roughly form the four corners of a square; anteriorly with a further pair of much smaller supra-occipitals and a minute to obsolete pair of supra-antennals (Fig. 304). Posterior margin with two prominent lateral coronals, which are roughly equal in length but somewhat more slender than the anterior and posterior coronals. Behind the eyes with a fairly distinct supra-orbital, which is somewhat smaller than the coronals. Antennae consisting of 26 segments and ± reaching half way along abdominal tergum IV. Pronotum slightly longer and noticeably wider than head, a little longer than wide and trapezoidal in outline with posterior margin some 1.3x wider than anterior margin; the lateral margins concave, indented medially and set with 4–5 minute spines. Antero and postero-lateral pronotals well pronounced but fairly small; inter-posterior pronotals obsolete or lacking (Fig. 304). Just before transverse median sulcus with a moderately sized pair of stout pre-median pronotals (Fig. 304). Mesothorax gently and gradually widening towards the posterior with posterior portion some 1.7x wider than anterior margin. Mesonotum very slightly narrowing towards the posterior and almost 2x longer than pronotum. Lateral margins set with 6–8 fairly small and slender, almost equally sized spines, the antero-lateral spine not larger


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than the remaining. Disc with a pair of fairly distinct anterior-mesonotals and otherwise only set with a few minute paired spinules or spiniform tubercles in the median portion (Figs. 304–305). Between bases of tegmina with two (rarely four) small and short posteromedial mesonotals (Fig. 304); if four spines present, the anterior pair decidedly smaller than the posterior pair. Tegminal small. Mesopleurae with a moderate, slender supra-coxal spine and about eight unequally sized but fairly distinct, slender laterals along lower margin; one of the median spines somewhat enlarged and comparatively longer than the others (Fig. 305). Metapleurae with lateral margin gently rounded and armed with six spines, the anterior five spines long and sometimes gently backcurving, 4th longest and the posteriormost spine considerably smaller; supra-coxal small. Sensory area of profurcasternum distinct, roundly triangular in outline and divided by a transverse furrow (Fig. 83). Mesosternum with a lateral row of three fairly distinct and strong spines, which increase in size towards the posterior. Metasternum with two small spines on each side of the median line and a fairly strong lateral spine. Tegmina ± reaching 2/3 along abdominal tergum II, oval with the radial vein strongly raised a forming a prominent longitudinal keel in basal portion of tegmen. Abdominal tergites II–III widening, IV widest and following narrowing, II–VII transverse with lateral margins moderately deflexed, straight and armed with seven small, slender, almost equally sized spines; the posteriormost somewhat enlarged. II smooth, III–VII with a few very minute paired spinules near median line and a transverse row of six minute spines close to posterior margin; these indistinct or obsolete on III and becoming increasingly pronounced towards VII. Sternites II–VI with a transverse row of 4–6 fairly distinct spines close to posterior margin and 1–2 pairs of spines in anterior portion, V–VI only with eight spines. The lateral invaginations of II–IV moderate and elongate oval (Fig. 85). Praeopercular organ formed by a pair of short but prominent, stout outward pointing postmedian spines on sternum VII (Fig. 303). Tergum VIII with 3–4 very minute spines along lateral margins. Anal segment with a blunt longitudinal keel, which is posteromedially protuded into a short, blunt spiniform projection; posterior margin rounded (Fig. 302). Epiproct shorter than anal segment with posterior margin roundly angular, notched medially and armed with 2–6 minute teeth (Figs. 301–302). Cerci small and compressed laterally. Subgenital plate gently downcurving in the apical portion (Fig. 301), the apex bluntly triangular and projecting considerably over apex of epiproct (Fig. 303). All legs strong and with all carinae spinose or dentate, the dorsal carinae of the tibiae only minutely denticulate. Metafemora reaching to abdominal segment VI and metatibiae projecting very slightly beyond apex of abdomen. Ventral surface of femora with a longitudinal median row of small spines (Fig. 364). Two ventral carinae of metatibiae armed with about seven large spines, which strongly increase in size and become increasingly incurving towards the apex of tibia; the most apical spine of the posteroventral carina largest (Fig. 383). ♂♂ (Figs. 294–295). Large (body length 82.0–92.0 mm) and slender with very prominent and long, bright green thoracic armature and parallel-sided abdominal segments III–VI. Colouration ranging from pale creamish to dark brown (usually darkening with age), rarely with a slight dull greenish hue. Head with a ± defined dark postocular stripe. Cephalic and thoracic armature bright green and gradually becoming brown towards the tips. Tegmina ranging from dull ochre to dark brown and usually with the veinature dark brown; either with a small white medio-lateral marking or great parts of the posterior portion white with a network of dark veins. Legs plain pale creamish to dark brown, the tibiae usually with a ± dull greenish wash; armature mostly dull green. Metacoxae dull red interiorly. All coxal trochanters bright blueish green (Fig. 394). Shape and armature of head as in ♀♀, but armature more prominent with all spines but the six coronals in particular much longer and more slender (Fig. 299). A fine longitudinal median coronal line present in most specimens. Antennae long and reaching to abdominal segment VI or VII; with 24 antennomeres. Pronotum generally as in ♀♀ but more elongate, with the pre-median pronotals in particular very much longer, upright and very large (Fig. 299). Antero and postero-lateral pronotals much more pronounced than in ♀♀ with the latter being somewhat larger than the former; usually with a minute pair anterior mesal pronotals and a more distant pair of small inter-posterior pronotals (Figs. 299–300). Mesothorax moderately elongate, some 1.8x longer than pronotum and gently widening towards the posterior (Fig. 300). Mesonotum slightly narrowing towards the posterior and about 2.2x longer than wide. Lateral margins set with 6–8 small almost equally sized spines and in anterior portion with a very prominent pair of long and slender anterior mesonotals; disc otherwise set with a variable number of irregularly dispersed minute spines. Rarely there may be a further strong pair of post-median mesonotals (e.g. a specimen photographed at Batang Ai National Park, Sarawak by the fourth author). Between the bases of the tegmina with four very large and strong posteromedial mesonotals on a common base, the anterior two of which are decidedly shorter than the posterior ones (Fig. 299). Tegminal fairly small. Meso- and metapleurae with armature



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as in ♀♀ but the spines averaging longer and the supra-coxal much more prominent and almost equal in size to the posteromedian mesonotals (Fig. 300). Sensory area of profurcasternum as in ♀♀. Armature of meso- and metasternum as in ♀♀ but metasternum with 4–5 distinct spines on each side of the median line. Tegmina ± reaching half way along abdominal tergum II, oval in shape and with the radial vein strongly raised a forming a prominent longitudinal keel in basal portion of tegmen. Tegminal spine distinct and fairly long and slender. Abdominal segment II gently narrowing, III–VI roughly uniform in width, parallel-sided and almost equal in length being averaging 1.3–1.4x longer than wide. VII shorter than previous and slightly widening towards posterior. Tergites II–VII all with 6–7 slender, somewhat unequally sized spines along lateral margins, the dorsal surface of III–VII set with a variable number of minute spinules. Tergum VIII only with three very minute spines on lateral margins and widening towards posterior. Abdominal sternum I with a fairly prominent pair of spines. Sternites II–VI with eight spines, which are most prominent on II and gradually decrease in size towards VI; VII only with 2–4 minute spines (Fig. 298). Lateral invaginations of II–IV moderate and slit-like. Anal segment with lateral margins sub-basally deflexed into a large, triangular, tooth-like lobe (Figs. 296–298), the posterior margin roundly excavated. Epiproct slender, elongate, narrowed towards a notched apex and considerably projecting beyond posterior margin of anal segment (Fig. 297, 411). Cerci small, club-like and distinctly compressed laterally in basal portion. Vomer broad, triangular with a fairly short and slightly curved terminal hook, which is mostly concealed by the poculum; dull orange with the apex black (Fig. 411). Poculum reaching almost half way along anal segment with posterior margin broadly rounded (Fig. 298). Legs all fairly long but moderately slender, mesofemora almost as long as pro- and mesonotum combined, metafemora roughly reaching to posterior margin of abdominal segment VI and metatibiae projecting distinctly beyond apex of abdomen. Armature generally as in ♀♀ but the ventral spines of the metatibiae considerably smaller and less curved (Fig. 384). Variability: A quite frequent species with variability mainly restricted to colouration. Females in particular are either plain pale to dark brown (e.g. Fig. 477) or may have the complete body pale straw or ochre with only the head, lateral margins of the pronotum, meso- and metapleurae and basal half of femora dark brown (Figs. 478, 481). While the tegmina are mostly plain brown (Fig. 477), some specimens possess a black central spot (Fig. 479) or have a weakly defined pale basal marking, which produces a roughly heart-shaped pattern (Figs. 478, 480–481) similar to that often seen in H. echinata (Redtenbacher, 1906). Occasionally there is a white spot at the anterolateral corner of the mesonotum or specimens possess a slight dark longitudinal median stripe along the entire dorsal surface of the body. Males show variability in the colouration of the tegmina. These are typically dark brown with the posterolateral portion broadly white and the veins marked by dark brown or black. Rarely however, specimens occur which have a white diagonal stripe on the tegmina and in others the white portion may be reduced to a fairly small and more defined area (Fig. 479). A ♀ in UMO is exceptionally small (body length 84.0 mm) and has a very finely spinose mesonotum. The spination of the thorax shows only slight variation but the paralectotype of H. grayii (Westwood, 1859) in NHMUK deserves particular mention, since the specimen has two instead of one pair of anterior mesonotals. A ♂ photographed by the fourth author at Batang Ai National Park is remarkable for having a strong pair of postmedian mesonotals (Fig. 479). Eggs (Figs. 431–432): Very large, distinctly lemon-shaped, with the polar-area acutely conical; capsule 1.3– 1.4x longer than wide. Capsule surface very minutely granulose and irregularly but sparsely covered with wart-like ventrally pitted structures, which are arranged in clusters and bear some very short setae; the setae somewhat longer on operculum. Micropylar plate very large and basically X-shaped but of a fairly complex shape. Anterior extensions very long, at the base with an angle of about 55° to each other, the median portion outcurving and the apex strongly expanded and angular with the exterior angle short and rounded but the interior angle elongated and gradually tapered; the interior angles facing towards each other. Posterior extensions more slender with the apex strongly expanded at both sides, forming a hammer-like shape. Outer margin of plate slightly raised, surface like capsule. Micropylar cup a small C-shaped swelling. Operculum round in cross-section and strongly conically raised. General colouration greyish mid to dull brown, the points of the conically raised operculum and polar-area black and the outer margin of the micropylar plate as well as the wart-like swellings very dark brown. Eggs of specimens from Batang Ai National Park (Sarawak) are remarkable for being almost entirely black and having the surface irregularly pitted. Measurements [mm]: length incl. operculum 10.6–12.1, length 9.0–10.2, width 5.9–6.1, height 6.9–7.0.


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Comments: There has been some confusion concerning the identity of this species, which has been clarified by Bragg (1992: 297). Kirby (1904b: 397) recognized the specimen in MHNG described as “Heteropteryx grayi var.” by Saussure (1869: 310) was a distinct species and named this as H. saussurei. Redtenbacher (1906: 171) was obviously unaware of Kirby’s publication and described his Heteropteryx saussurei based on the same specimen in MHNG, hence Redtenbacher’s name is a homonym and objective synonym of H. saussurei Kirby, 1904. Since Redtenbacher included two further specimens in his original type-series, the ♀ in MHNG is here selected as the lectotype in order to guarantee stability and validity of the synonymy established by Bragg (1992: 297). While the ♀ paralectotype in MCSN is conspecific with the lectotype, the second ♀ in NHMW (No. 285) is a remarkably strongly spinose specimen of H. echinata (Redtenbacher, 1906). The ♂ of H. saussurei was described as H. echidna by Rehn (1938) and synonymised by Bragg (1992: 297). The eggs of this species are very different in shape from the other Bornean representatives of the genus, the lemon-shaped capsule rather resembling eggs of the muelleri species-group, which is not found in Borneo. H. saussurei is being successfully cultured in Europe since 1994 from stock originating in Bako National Park and Tarum in NW-Sarawak. It was included on the Phasmid Study Group culture list as culture No. 177 but is not quite as easy to rear as other Bornean species of the genus. Alternative foodplants accepted in captivity include bramble (Rubus fruticosus, Rosaceae), raspberry (Rubus idaeus, Rosaceae), roses (Rosa spp., Rosaceae), hawthorn (Crataegus monogyna, Rosaceae), ivy (Hedera helix, Araliaceae), oaks (Quercus spp., Fagaceae) and eucalyptus (Eucalyptus gunnii, Myrtaceae). Distribution (Fig. 496): Restricted to coastal or near coastal areas in NW-Borneo. Brunei [UMO]. NWSarawak: Mount Sadong [ANSP, MCZC]; Tarum, near Debak [coll. PEB]; Sarikei [SMSM]; Simunjan [Bragg, 2001: 90]; Kuching [SFDK, SMSM, coll. PEB]; Kuching, 10th mile, Timber Research Centre [SFDK]; Bako National Park 50 m [coll. FH, coll. OC, coll. FSC, coll. PEB]; Batang Ai National Park [coll. FSC]. Specimens examined: 30 TABLE 17. Measurements [mm] of Haaniella saussurei Kirby, 1904 ♂♂

♀♀

82.0–89.0 82.0–92.0

104.0–112.0 127.0–132.0

Head

6.8–7.2

12.3–12.8

Pronotum

7.8–8.5

9.3–10.7

Mesonotum

11.2–13.5

15.0–16.3


11.5–12.3

14.5–16.0

Tegmina

10.2–17.0

26.0–27.0

Profemora

20.5–22.5

21.0–23.6

Protibiae

22.5–24.8

26.0–29.0

Mesofemora

17.0–18.5

19.0–21.2

Mesotibiae

18.3–19.5

23.0–24.2

Metafemora

24.5–28.0

30.0–31.5

Metatibiae

26.0–29.2

35.0–36.8

Antennae

64.0–71.5

66.0–68.0


Haaniella scabra (Redtenbacher, 1906) (Figs. 87. 91, 96, 306–322, 381–382, 395–396, 408, 433–436, 482–484, 496) [Small Haaniella] Heteropteryx scabra Redtenbacher, 1906: 171. LT [by present designation], ♂: Coll. Br. v. W., Kina-Balu, Rolle, det. Redtenb. Heteropteryx scabra, 25.63 [NHMW, No. 286]; PLT, ♀: Coll. Br. v. W., Kinabalu, Borneo, Staudinger, det. Redtb. Heteropteryx scabra, 20.832 [NHMW, No. 286]; PLT, ♀: Kina-Balu, ex. Heyne coll. [ZIN]; PLT, ♀: Kinabalu, Borneo;



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Brunner v. W. et Redtenbacher det. 1903 Type; Heteropteryx scabra Redt. [ETHZ]. Rehn, 1938: 369. Haaniella scabra, Cocking, 1985: 6 (figs. ♂, ♀). Bragg, 1998: 26, figs. 24–26 (♀ abdomen) & 33–34 (egg) Brock, 1998: 56. Seow-Choen, 1998: 89, table 1. Seow-Choen, 2000a: 151, figs. 11 (♂), 12 (♀). Bragg, 2001: 93, figs. 15 g–h (egg), 22 a (♂), b (♀), c (egg), 23 a–c (♀ abdomen), d–e (egg). Brock, 2007: 54. Dräger, 2011: 48, figs. 10: 3a–c (♀ abdomen), 11: 6 (♂ thorax), 19 (♂), 20 (♀), fig. 37: 7 (egg). Seow-Choen, 2016: 410, figs. 1043-1046. Haaniella grayi scabra, Günther, 1932: 313. Günther, 1935: 124. Günther, 1936: 124. Haaniella echinata scabra, Günther, 1944: 72.

Material examined (27 ♂♂, 41 ♀♀, 17 nymphs, eggs): N-SABAH: 1 ♀: B.N. Borneo, Mt. Kinabalu, Kiau, 3000ft., 2.5.1929 [RMNH]; 1 ♀, 3 nymphs: B.N.Borneo, Mt. Kinabalu, Lumu Lumu, 5500ft., 8.4.1929; Haaniella scabra (Redtb. det. Klante [RMNH]; 2 nymphs: B.N. Borneo, Mt. Kinabalu, Kenokok, 3300ft., 23.10.–26.4.1929; Haaniella scabra (Redtn. det. Klante [RMNH]; 1 ♂: Kina Balu, Borneo, Dobiasch, coll. R. Ebner det. Karny Heteropteryx scabra [NHMW, No. 286]; 5 ♀♀, 2 ♂♂: Sabah, Kinabalu NP, 5200ft, 10.–19.I.1981, A. Harman; “Haaniella scabra” det. C.L.Chan, 1989 [NHMUK, 1982-387]; 1 ♂: Kinabalu N.Park, Sabah (N. Borneo), c. 5200ft. a.s.l., 24 May 1984, Coll. C.L. Chan [NHMUK]; 1 ♀: Kinabalu N.Park, Sabah (N. Borneo), c. 5200ft. a.s.l., 24.V.1984, Coll. C.L. Chan [NHMUK]; 1 ♂, 2 ♀♀: Kinabalu N.Park, Sabah (N. Borneo), c. 5200ft. a.s.l., 9.X.1982, Coll. C.L. Chan [NHMUK]; 1 ♂, 1 ♀ (penultimate instar): Kinabalu N.Park, Sabah (N. Borneo), c. 5200ft. a.s.l., 1-V-1986, Coll. C.L. Chan & M.Y. Chan [NHMUK]; 1 ♂, 1 ♀ (penultimate instar): Kinabalu N.Park, Sabah (N. Borneo), c. 5200ft a.s.l., 22.V.1982, Coll. C.L. Chan & J. Lee [NHMUK]; 1 ♀, 2 ♀♀ (penultimate instar): Kinabalu N.Park, Sabah (N. Borneo), c. 5200ft a.s.l., 24.IV.1982, Coll. C.L. Chan & V.K. Lo [NHMUK]; 1 ♀ (penultimate instar): Kinabalu N.Park, Sabah (N. Borneo), c. 5200ft a.s.l., 14.II.1982, Coll. C.L. Chan et al [NHMUK]; 1 ♂: KNP-Sabah, 10–19 Jan 81, A. Harman no. 69 ♂ [NHMUK]; 1 ♀, 1 egg: Kinabalu N.Park, Sabah (N. Borneo), c. 5200ft. a.s.l., Jan. 1981, Coll. A. Harman [NHMUK]; 1 ♀: Silau, 13 July 1990, Chan [NHMUK]; 1 ♂: Borneo, Sabah, Mt Kinabalu, Silau Silau Trail c. 5200', 10th November 1991, Coll. C.L. Chan [NHMUK]; 1 ♂ (penultimate instar): Silau-silau Trail, Kinabalu N.Park, sabah (N. Borneo), c. 5200ft. a.s.l., 13-VII-1990, Coll. C.L. Chan [NHMUK]; 1 ♂: Marai-Parai, Mt. Kinabalu, Sabah (N.Borneo), II.1985, Coll. C.L. Chan [NHMUK]; 1 ♂, 3 ♀♀, 1 ♂ (nymph): Brit. NordBorneo, Kinabalu-Gebirge, 16.XII.1912, Grubauer leg.; “Heteropteryx scabra Redtb.” det. Klante, 1962 [SMNS]; 1 ♂, 2 ♀♀, 1 ♀ (n5), 1 ♀ (n3): Kinabalú, Borneo, ca. 1500 m, Waterstradt leg., H. Rolle vend. 25.IX.1904; K. Günther determ. Vers. Nr. 131, 1932; Haaniella scabra Redt., K. Günther det. [ZMUH]; 2 ♂♂, 4 ♀♀, eggs: NSabah, Mt. Kinabalu, near HQ, Silau-Silau trail, 1500m, leg. F. Hennemann & O. Conle 4.–8.VIII.1996 [coll. FH, No’s 0328-1 to 6 & E1]; 1 ♂, 2 ♀♀: N-Sabah, Mt. Kinabalu, near HQ, Silau-Silau trail, 1500m, leg. F. Hennemann & O. Conle 4.–8.VIII.1996 [coll. OC]; 2 ♂♂, 2 ♀♀, 1 ♀ (nymph n3): ex Zucht: Rob Krijns 2009, Herkunft: Sabah, Mount Kinabalu, 1500 m (PSG No. 71) [coll. FH, No’s 0328-7 to 11]; 4 ♂♂, 5 ♀♀, eggs: ex Zucht: F. Hennemann 2014, Herkunft: N-Sabah, Mount Kinabalu, 1500 m (PSG No. 71) [coll. FH, No’s 0328-12 to 20 & E2]; 1 ♂, 4 ♀♀, 2 ♀♀ (nymphs): ex Zucht O. Conle, 1999–2000, Herkunft: Borneo, Sabah, Mt. Kinabalu N.P., Silau-Silau trail, near headquarters, 1500 m, 04.–05.08.1996, leg. Conle & Hennemann [coll. OC]; 4 ♂♂, 3 ♀♀: ex Zucht O. Conle, 1999–2000, Herkunft: Borneo, Sabah, Mt. Kinabalu N.P., Silau-Silau trail, near headquarters, 1500 m, 04.–05.08.1996, leg. Conle & Hennemann [coll. OC]; 3 ♂♂, 3 ♀♀: leg FSC, Crocker Range National Park, 1200–1600 m asl. [coll. FSC]. Diagnosis: Very closely related and the adelphotaxon of H. echinata (Redtenbacher, 1906). The enlarged antero-lateral mesonotals of both sexes as well as the distinct inter-posterior pronotals, gradually tapered abdominal segments II–V and short, rounded epiproct of ♂♂ are shared with this species. Both sexes are however frequently distinguished from H. echinata by: the much smaller size; much larger supra-occipitals; conspicuously


HENNEMANN ET AL.

shortened antennomeres; having only two posteromedial mesonotals between the bases of the antennae (four in echinata); scabrous to rugulose abdominal tergites II–VI (spinulose in echinata); considerably longer spines of abdominal sternites II–VII and comparatively longer tegmina, which clearly project over the posterior margin of abdominal tergum II. The eggs differ from those of H. echinata by the much smaller dimensions, broader micropylar plate and characteristic colouration. Etymology: The name (scaber lat. = rough, scabrous) refers to the scabrous to rugulose but not spinose abdominal tergites of this species. Description: ♀♀ (Figs. 306, 308, 309–310). Small (body length 59.3–79.5 mm) and fairly stocky with long and slender legs, only two posteromedial mesonotals and strong ventral spination of the abdomen. Colour very variable and ranging from dull ochre (e.g. Figs. 309–310) over various shades of brown to almost black (usually darkening with age), either plain or prettily mottled (Figs. 306, 308). Occasionally specimens may have most of abdominal tergum VI more pale in colour or cream and sometimes specimens occur, which have a faint dark longitudinal median stripe running along most of the dorsal body surface (Fig. 483). Abdominal tergum VIII often with a pair of black markings at anterior margin (Fig. 306). Cepahlic and thoracic armature dark brown to black basally with the apical half red. Tegmina ranging from plain pale to dark brown and often with a bold black spot in centre (Figs. 309, 483) or more rarely with the basal portion pale cream to whitish and these pale portions forming a roughly heart-shaped marking when the tegmina are closed. Legs either plain or femora with a pale transverse band postmedially; tibiae often more pale in colour than femora. Interior surface of metafemora more or less distinctly pale red or at least with a slight reddish hue; metacoxae black interiorly with some pale red markings. Coxal trochanters of metacoxae and membrane between abdominal sternites I–III dark greyish blue (Fig. 395). Head about 1.2x longer than wide, parallel-sided with the vertex very gently convex in posterior portion; armed with 10–12 spines (Figs. 313–314). Vertex with four strong and almost equally sized coronals which roughly form the four corners of a square; anterior of these with a distinct pair of supra-occipitals that is just slightly shorter than the coronals. Supra-antennals minute (Fig. 313). Posterior margin with two prominent lateral coronals, which are roughly equal in length to the anterior and posterior coronals and behind the eyes with a somewhat shorter, stout supra-orbital. Antennae consisting of 26 segments and ± reaching half way along abdominal tergum III. Pronotum slightly longer and decidedly broader than head, a little longer than wide and trapezoidal in shape with posterior margin some 1.3x wider than anterior margin; the lateral margins concave, indented medially and set with 4–5 minute spines. Antero and postero-lateral pronotals distinct and near posterior margin with a similarly sized pair of inter-posterior pronotals. Just before the V-shaped transverse median sulcus with a prominent pair of upright pre-median pronotals (Fig. 313). Mesothorax very gently and gradually widening towards the posterior with posterior portion some 1.3x wider than anterior margin. Mesonotum slightly narrowing towards the posterior and about 1.6x longer than pronotum. Lateral margins with 5–6 very small equally sized spines and a fairly prominent and long antero-lateral spine. Disc with a pair of prominent anterior-mesonotals, which are somewhat larger than the antero-lateral mesonotals; otherwise set with a variable number of minute paired spinules or spiniform tubercles mostly in the median portion (Fig. 314). Between bases of tegmina with two moderately distinct, upright posteromedial mesonotals; rarely with a further much smaller pair anteriorly (Fig. 313). Tegminal small and stout. Mesopleurae with a strong supra-coxal spine and 6-7 unequally sized but fairly distinct laterals along lower margin, the most anterior one larger than the remaining and the 4th spine somewhat enlarged (Fig. 314). Metapleurae with lateral margin rounded and armed with five spines, the posteriormost spine considerably smaller than the remaining and the supra-coxal moderately prominent and slightly directed posteriad. Sensory area of profurcasternum covering more than half of segment and this segment otherwise with a small spine near each outer angle. Mesosternum with a lateral row of three short but strong spines, which increase in size towards the posterior; a few further but much smaller spines present in the median portion. Metasternum with 4–5 short but fairly strong spines on each side of the median line, the outermost of which is strongest. Tegmina ± reaching to posterior margin of abdominal tergum III, oval with the apical portion increasingly narrowed; the radial vein strongly raised a forming a prominent longitudinal keel in basal portion of tegmen. Abdominal tergites II–III widening, IV widest and the remaining narrowing, all transverse. Tergites II–VII with lateral margins deflexed and either straight to gently rounded and armed with 6–8 small but subequally sized spines. II smooth, III–VII with a few very minute paired spinules near median line and a transverse row of six minute spines close to posterior margin and otherwise scabrous; spinules indistinct or obsolete on III and becoming increasingly pronounced towards VII. Sternites II–VI with eight spines, which are very pronounced and often remarkably long on II and decrease in size towards VI.



93

Lateral invaginations very distinct and oval in II, obsolete on III–IV (Fig. 87). Praeopercular organ formed by a pair of short and stout closeby post-median spines on a convex median swelling on sternum VII (Fig. 319). Tergum VIII with 3–5 very minute spines on lateral margins. Anal segment with a blunt longitudinal keel, which is posteromedially protuded into a blunt spiniform projection; the posterior margin angular with the outer angles somewhat protuded (Fig. 318). Epiproct shorther than anal segment with posterior margin angular, notched medially and armed with 4–6 minute teeth (Fig. 318). Cerci small and compressed laterally. Subgenital plate downcurving in the apical portion, the apex bluntly triangular and projecting considerably over apex of epiproct (Figs. 317–319). All legs fairly long and slender with all carinae spinose or dentate, the dorsal carinae of the tibiae just very sparsely and minutely denticuate. Metafemora reaching to abdominal segment VII and metatibiae projecting somewhat beyond apex of abdomen. Ventral surface of femora with a few minute denticles arranged in a longitudinal median row (Fig. 365). Two ventral carinae of metatibiae armed with about six spines, four of which are are fairly distinct, somewhat incurving and roughly equal in size (Fig. 381). ♂♂ (Figs. 307, 311–312): Small (body length 45.5–57.0 mm) and stocky with prominent antero-lateral mesonotals, two strong posteromedial mesonotals and very prominent ventral armature of the abdomen. Colouration ranging from pale to very dark brown (usually darkening with age), mostly plain but often with most of abdominal tergum VI pale ochre. Sometimes specimens may have a faint dark longitudinal median stripe running along most of dorsal body surface. Cephalic and thoracic armature brown basally with the apical half orange to red (Figs. 315–316). Tegmina with the basal portion pale cream to white (these pale portions forming a roughly heart-shaped marking when the tegmina are closed (Fig. 482), a broad white to cream transverse median band (e.g. Fig. 307) or with a pale cream to white diagonal stripe. Interior surface of metafemora more or less distinctly dull red or at least with a slight reddish hue; metacoxae black interiorly. Coxal trochanters of metacoxae and membranes between abdominal sternites I–III dark greyish blue (Fig. 396). Shape and armature of head as in ♀♀, but armature more prominent with all spines comparatively longer and more slender (Fig. 315); the supra-occipitals distinct but noticeably smaller than the anterior and posterior coronals. Antennae long, consisting of 24 segments and reaching to abdominal segment V or VI. Pronotum generally as in ♀♀ with but all spines considerably more pronounced; often with one or two minute pairs of postmedian pronotals behind transverse median sulcus (Figs. 315–316). Mesothorax short, only some 1.4x longer than pronotum and gently widening towards the posterior. Mesonotum narrowing towards the posterior and only about 1.3x longer than width of anterior margin; armature as in ♀♀ with all spines much longer and more slender and also the antero-lateral mesonotals considerably more pronounced; usually with a variable number of short paired spines or spiniform tubercles in the median portion. Posteromedial mesonotals strong and slighly directed anteriad (Fig. 315). Tegminal comparatively longer and more slender than in ♀♀. Meso- and metapleurae with armature as in ♀♀ but the spines much longer and the supra-coxal very prominent and the longest of all the thoracic spines (Fig. 316). Sensory area of profurcasternum as in ♀♀. Armature of meso- and metasternum as in ♀♀ but with all spines more pronounced. Tegmina ± reaching to posterior margin of abdominal tergum II, oval with the apical portion increasingly narrowed; the radial vein strongly raised a forming a prominent longitudinal keel in basal portion of tegmen. Abdominal segments II–V gently and gradually narrowing, VI narrowest segment and almost parallel-sided, VII and VIII slightly widening; all wider than long. Lateral margins of tergites II–VI with about seven small and short, somewhat unequally sized spines, VII only with 4–5 small spines. Dorsal surface of II smooth, of IIIVII scabrous to granulose and at best with a very indistinct pair of posteromedian tubercles. Tergum IX with posterolateral angles protuded into a blunt, posteriorly directed spiniform process. Sternites II–VI with eight spines, which are very prominent on II but become increasingly smaller towards VI (Fig. 91), VII unarmed (Fig. 322). Lateral invaginations of II very large, those of III and IV much smaller and more slender (Fig. 91); occasionally II–III with a further pair of antero-median spines Anal segment with lateral margins sub-basally deflexed into a blunt, triangular, tooth-like lobe (Fig. 320), the posterior margin distinctly idented medially (Fig. 321). Epiproct short, transverse and broadly rounded (Fig. 321). Cerci small and compressed laterally. Vomer broad basally with the apical portion strongly narrowed and protuded into a fairly slender dextrally curved terminal hook; ochre with the apex black (Figs, 408). Poculum projecting somewhat over posterior margin of tergum IX with posterior margin broadly rounded (Fig. 322). Legs all fairly long and moderately slender, mesofemora almost as long as pro- and mesonotum combined, metafemora reaching to abdominal segment VII and metatibiae projecting considerably beyond apex of abdomen. Armature as in ♀♀ but ventral spines of metatibiae considerably smaller (Fig. 382).


HENNEMANN ET AL.

Variability: This species shows some variability in size, degree of body spination and colouration but is by far less variable than the closely related but widely distributed H. echinata (Redtenbacher, 1906). The type specimens of H. scabra in NHMW, ETHZ and ZIN and specimens in SMNS are averaging larger and slightly more robust with the body armature somewhat less developed than specimens collected more recently near the headquarters of Kinabalu National Park at an altitude of 1500 m. Presumably these specimens were collected at lower altitudes, since in general specimens tend to become smaller, less robust and exhibit relatively longer and more slender legs with increasing altitude. Furthermore, the specimens from the Kinabalu National Park Headquarters differ from the historical specimens mentioned above by the more prominent and longer ventral spination of the body, comparatively longer tegmina and ♀♀ often have the abdominal tergum VI straw to ochre and paler in colour than the remaining tergites (Fig. 310). Both sexes show considerable variability in the colouration of the tegmina, the colour patterns being similar to those seen in H. echinata (Redtenbacher, 1906). Females mostly have plain light to dark brown tegmina but often there is a variably sized blackish central spot (Figs. 309, 483) and occasionally there are specimens with slightly paler mottling in the apical half of the tegmina (e.g. Fig. 306). Males usually exhibit a large pale marking in the basal portion of the tegmina, which produces a roughly heart-shaped marking when these organs are closed (Fig. 482). More rarely these white markings are reduced to a diagonal stripe (e.g. Fig. 307). Females range in colour from dull straw (Fig. 482) over ochre and mid to dark brown (Fig. 482), but generally tend to darken with raising age. A specimen photographed by the fourth author is ochre and has a broad dull brown longitudinal stripe along the entire dorsal surface of the body (Fig. 483). Eggs (Figs. 433–436): Fairly small, barrel-shaped with the polar-area rounded; 1.3–1.4x longer than wide. Entire surface setose, the setae very long (on operculum in particular) and irregularly arranged in clusters which indicate hints of a net-like texture. Spaces between the hairy clusters very minutely granulose. Microyplar plate broadly X-shaped with all four extensions fairly short; anterior extensions at an angle of 75° to each other with the apex roundly widened; posterior extensions with the apex strongly expanded and club-like. Outer margin of plate gently raised. Micropylar cup a shallow impression with a small, cup-like swelling posteriorly. Operculum almost circular and moderately convex. Capsule pale to mid brown with all the setae dark brown, micropylar plate dark brown with the outer margin black. A single egg laid by a wild-caught ♀ from Silau-Silau trail (Mount Kinabalu NP) in coll. FH differs from those described above by the more elongate capsule (1.5x longer than wide) and comparatively less numerous and shorter hairy structures and setae (Figs. 433–434). Measurements [mm]: length incl. operculum 6.0–7.5, length 5.5–6.9, width 4.5–4.8, height 5.0–5.1. Comments: H. scabra was originally described from a total of four specimens and the ♂ in NHMW is here selected as the lectotype to guarantee stability of the name. This small mountainous species has its distribution restricted to the Mount Kinabalu massive and surrounding mountains in Northern Sabah and altitudes above 800 metres. Detailed examination and comparison shows H. scabra to be the sister-taxon of H. echinata (Redtenbacher, 1906), which is a widely distributed and averaging much larger lowland species that seems to be restricted to altitudes below 500 metres. While there clearly is a vertical separation between these two species the armature of both sexes is almost identical and both are very similar in appearance at first glance. In contrast to the most other members of Haaniella this species has proven somewhat more difficult to rear in captivity. Stock from Mount Kinabalu has been introduced to Europe on several occasions, the first live ♀ having been imported by Jonathan Cocking (England) in 1984. However, no culture was established from this specimen and most later attempts to rear H. scabra have only been successful for a few generations. Stock introduced by the two first authors in 1996 was not very productive and could be maintained in culture for three generations by the second author. Stock introduced recently appears easier to rear and is meanwhile being successfully kept by numerous breeders in Europe. In the mid 1980’s H. scabra was included on the Phasmid Study Group culture list as culture No. 70. Alternative foodplants accepted in captivity in Europe include bramble (Rubus fruticosus, Rosaceae), raspberry (Rubus idaeus, Rosaceae), roses (Rosa spp., Rosaceae), pyracantha (Pyracantha coccinea, Rosaceae), hawthorn (Crataegus monogyna, Rosaceae), ivy (Hedera helix, Araliaceae), oaks (Quercus spp., Fagaceae) and salal (Gaultheria shallon, Ericaceae). Distribution (Fig. 496): Restricted to mountainous localities (above 800 m) in NE-Borneo. N-Sabah: Mount Kinabalu [NHMUK, NHMW, ETHZ, SMNS, ZMUH]; Mount Kinabalu, Lumu Lumu 5500 ft. [RMNH]; Mount Kinabalu, Kiau 3000 ft. [RMNH]; Mount Kinabalu, Kenokok 3300 ft. [RMNH]; Mount Kinabalu, near Park



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Headquarters 1580 m [NHMUK, coll. PEB]; Mount Kinabalu, Silau-Silau trail, 1500 m [NHMUK, coll. FH, coll. OC]; Mount Kinabalu, Mari-Parai [NHMUK]; Crocker Range National Park, Mount Trus Madi 4000–6000 ft. [NMSC]; Crocker Range National Park, 1200–1600 m [coll. FSC]. The record from Labuan Island by Rehn (1938: 369) was questioned by Bragg (1998; 2001) because all other records are from highly mountainous localities on Mount Kinabalu and Mount Trus Madi. This record can here be confirmed as wrong, since examination of the concerned specimens (1 ♂ and 2 ♀♀) in ANSP has shown these to be H. echinata (Redtenbacher, 1906). Specimens examined: 89 TABLE 18. Measurements [mm] of Haaniella scabra (Redtenbacher, 1906) ♂♂

♀♀

Body

45.5–57.0

59.3–79.5

Head

4.6–4.9

7.0–9.0

Pronotum

4.5–4.9

7.7–8.0

Mesonotum

5.2–6.0

8.2–11.2


7.2–7.6

9.0–11.0

Tegmina

13.0–13.2

16.3–19.8

Profemora

12.8–13.2

13.0–16.4

Protibiae

14.8–15.1

16.5–19.5

Mesofemora

10.2–10.5

11.3–14.8

Mesotibiae

12.1–12.3

13.0–17.0

Metafemora

15.4–15.8

17.6–21.6

Metatibiae

18.3–19.0

21.5–25.2

Antennae

41.5

31.5–43.0

4.2. Heteropteryx Gray, 1835 Type-species: Phasma dilatatum Parkinson, 1798: 190, pl. 18: 2–5 (♀ and egg), by monotypy. Heteropteryx Gray, 1835: 13, 32. Westwood, 1859: 81. (in part) Westwood, 1864: 16. Westwood, 1874: 173, pl. 32: 1. Lucas, 1882: 32. Brunner v. Wattenwyl, 1893: 98. (in part) Rehn, 1904: 89. (in part) Kirby, 1904b: 396. Dohrn, 1910: 403. (in part) Karny, 1923: 234. (in part) Bradley & Galil, 1977: 198. (in part) Brock, 1995: 96. Bragg, 1998: 28. Brock, 1999: 183. Bragg, 2001: 97, fig. 24 (egg). Seow-Choen, 2016: 411. Cyphocrania, Westwood, 1859: 111. (in part) Leocrates Stål, 1875: 48. Type-species: Phasma (Eurycantha ?) graciosa Westwood, 1848, by monotypy. [The type-species is a junior objective synonym of Phasma dilatatum Parkinson, 1798, synonymised by Kirby, 1904a: 443] Redtenbacher, 1906: 166. (in part) Giglio-Tos, 1910: 13.


HENNEMANN ET AL.

Phasma, Parkinson, 1798: 190, pl. 1: 2–5 (♀ and egg). Phasma (Eurycantha ?), Westwood, 1848: 16, pl. 7: 3 (♂).

Diagnosis (♂, ♀): Very large Heteropteryginae (body length of ♀♀ usually > 135.0 mm). General colouration of ♀♀ bright green or yellow. Head slightly longer than wide and higher than long, vertex strongly convex, conically elevated and projecting over anterior margin of pronotum. Antennae very long, with all except for the basal segments exttemely elongated and reaching to abdominal segment IV (♀♀) or almost to apex of abdomen (♂♂). Mesothorax of both sexes strongly widening towards the posterior and trapezoidal in dorsal aspect. Mesonotum of ♂♂ very short, equal in length to pronotum. ♂♂ macropterous; tegmina at least ¾ the length of alae. Alae of ♂♂ without a tympanal area (= stridulatory organ). Alae of ♀♀ plain translucent pink, anal region of alae in ♂♂ dull pink and reticulate, costal region brown. Abdomen of ♀♀ strongly dilated, with all tergites laterally deflexed. Abdominal tergites V–VII of ♀♀ with distinct dorsal spines. Abdominal tergites II–IX of ♂♂ with dentate lateral lobes. Ventral surface of meso- and metafemora smooth. Eggs: Large, barrel-shaped with anterior portion of capsule slightly narrowing. Surface of capsule with irregular distinct impressions. Polar-area and operculum flat. Differentiation: Distinguished from the sister-taxon Haaniella Kirby, 1904 by the larger size, distinctly conically elevated head, which is higher than long and distinctly widening mesothorax. Females furthermore differ by the bright green or yellow colour, presence of distinct dorsal spines on abdominal tergites V–VII and plain translucent pink alae (with dark veinature in Haaniella). Males are distinguished from those of Haaniella by the shorter mesothorax, which is roughly equal in length to the pronotum, lack of a tympanal area (= stridulatory organ) in the alae, having the tegmina at least ¾ the length of the alae (only ½ the length in macropterous taxa of Haaniella) and having the anal fan of the alae dull red. Eggs differ by the distinctly pitted capsule surface and having the polar-area and operculum never convex or conical as in Haaniella, but flattened. Distribution: Peninsular Malaysia, Singapore, Thailand, Sumatra and NW-Borneo.

Species included: 1. Heteropteryx dilatata (Parkinson, 1798: 190, pl. 18: 2–5). = Heteropteryx castelnaudii Westwood, 1859: 174, pl. 32: 2. = Phasma graciosa Westwood, 1848: 16, pl. 7: 3. = Heteropteryx hopei Westwood, 1864: 16. = Heteropteryx rollandi Lucas, 1882: 32.

Heteropteryx dilatata (Parkinson, 1798) (Figs. 84, 93, 323–340, 347, 350–354, 356–357, 377–378, 441–444, 447, 485–488, 497) [Jungle Nymph] Phasma dilatatum Parkinson, 1798: 190, pl. 18: 2–5 (♀ and egg). HT, ♀: Asia (ex. Leverian Muserum), E166 [MAMU]. Phasma dilatatum, Shaw, 1806: 348, pl. 347. Shaw, 1806: 45. Serville, 1838: 281. Diapherodes dilatata, Burmeister, 1838: 574. Heteropteryx dilatata, Gray, 1835: 32. Westwood, 1859: 82 & 181. Westwood, 1874: 173, pl. 32: 1. Kirby, 1904b: 397. Rehn, 1904: 89. Dohrn, 1910: 402. Günther, 1932: 314. Günther, 1935: 124. Günther, 1936: 124. Löser & Schulten, 1981: 23–27. Brock, 1989: 207–211. Brock, 1994: 33, pl. NN: 9 & 10 (Gynandromorph). Chan & Lee, 1994: 5, figs. of egg, ♂ and ♀.



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Seow-Choen, Brock & Seow-En, 1994: 7 (figure of ♂ and yellow ♀) Brock, 1995: 96. Schulten, 1995: 100, figs. 61 (♀), 62 (♀), 63 (♀ & egg). Seow-Choen, F., 1995: 49. (Gynandromorph) Seow-Choen, F., 1996: 122, fig. 1 (Gynandromorph). Brock, 1997: 109, fig. 1 (♀), pl. 97K : 5 & 6 (♀), 7 (♂ nymph), pl. 97L : 8 (♂). Bragg, 1998: 28 (figs. 37–38) Seow-Choen, 1998: 89, table 1. Seow-Choen, 2016: 411, figs. 1047-1049. Brock, 1999: 143 & 183, figs. 96 (♀), 97 (♂), pls. 28–30 (♂, ♀, gynandromorph) Lipinski et al., 1999: 98, fig,. 45. (REM-study of egg) Seow-Choen, 2000b: 40, pls. 105 (♂), 106 a–c (♀), d (egg), 107 (gynandromorphs) Bragg, 2001: 98, fig. 24 (egg). Otte & Bock, 2005: 160. Dräger, 2011: 40, figs. 3 (♀), 4 –5 (♂), 6 (♀ nymph), 7 (♂), 8–9 (♂ nymph), 37:1 (egg). Heteropteryx dilatatus, Miller, 1939: 48. Leocrates dilatatus, Redtenbacher, 1906: 166. (Description of ♂) Heteropteryx castelnaudii Westwood 1874: 174, pl. 32: 2 (♂). HT, ♂: Tringany [UMO, No. 527] (Synonymised by Günther, 1935: 124) Heteropteryx (?) Castelnaudi, Kirby, 1904b: 398. Leocrates Castelnaudi, Redtenbacher, 1906: 167. Phasma (Eurycantha ?) graciosa Westwood, 1848: 16, pl. 7: 3 (♂). HT, ♂: Prince of Wales Island [UMO, No. 526] (Synonymised by Kirby, 1904b: 397) Cyphocrania graciosa, Westwood, 1859: 111. Leocrates graciosus, Stal, 1875: 48. Heteropteryx hopei Westwood, 1864: 16. HT, ♀: without data [UMO, No. 488]. (Synonymised by Westwood, 1874: 173) Dohrn, 1910: 403. Günther, 1932: 314. Heteropteryx Rollandi Lucas, 1882: 32. HT, ♀: Malaysia [MNHN - not traced]. (Synonymised by Kirby, 1904b: 397)

Material examined (49 ♂♂, 47 ♀♀, 3 gynandromorphs, 18 nymphs, eggs): PENINSULAR MALAYSIA: 1 ♀, 1 ♀ (nymph): Malacca; Nr. 3202 [MNHU]; 1 ♀ (nymph): Malay Penin., Selangor, Bukit Kutu, 3000ft., 12.4.1926, M. Pendlebury [RMNH]; 1 ♀ (nymph): Pahang, F.M.S., Lubok Tamang, 3500ft., 16.3.1924, H.M. Pendlebury [RMNH]; 1 ♀ (nymph): without data; Heteropteryx dilatata (Parkinsin)det. Bragg, 1993 [RMNH]; 1 ♂ (nymph): Perak, F.M.S., Wrays Hill, e. ovo. Nr. Taiping, 1897 [RMNH]; 1 ♀ (nymph): Malay Penin., Pahang, East, 1921, W.H.D. Edwards [RMNH]; 1 ♂ (nymph): Perak, Malacca; Leocrates castelnaudii det., Coll. GiglioTos [MIZT]; 4 ♂♂, 2 ♀♀: Malaisie, Cameron Highlands, acheté Pfanner [MHNG]; 1 ♂: Selangor [MHNG]; 2 ♂♂ (penultimate instar): Cameron Highlands, Malaysia, 9.1979 [MHNG]; 1 ♀ (penultimate instar): Perak (Malacca), A. Grubauer leg., B. Jachau vend. 15.XII.1901 [ZMUH]; 4 ♀♀, 8 ♂♂: W-Malaysia, Perak, Tapah Hills 500 m, leg. M.K.P. Yeh, VIII. 1993 [coll. FH, No’s 0018-1 to 12]; 7 ♀♀: W-Malaysia, Perak, Tapah Hills 500 m, leg. M.K.P. Yeh 1995 [coll. FH, No’s 0018-22 to 28]; 6 ♂♂, 1 ♂ (penultimate instar), 1 ♀ (nymph n2), 1 ♂ (nymph n3), eggs: ex Zucht F. Hennemann, 1996, Herkunft: W-Malaysia [coll.FH, No’s 0018-13 to 21 & E1, E2]; 1 ♂ (nymph n3): W-Malaysia, Cameron Highlands, Tanah Rata, Jalan Tengkolog, 1500 m, leg. F. Hennemann & O. Conle, 21.–25.VI.1996 [coll. FH, No. 0018-29]; 2 ♂♂: ex Zucht F. Hennemann 2010, Herkunft: W-Malaysia [coll. FH, No’s 0018-30 to 31]; 1 Gynandromorph (mostly ♂): ex Zucht F. Hennemann 1996, Herkunft: W-Malaysia, Tapah Hills [coll. FH, No. 0018-37]; 1 right ♀ tegmen: ex Zucht F. Hennemann 1996, Herkunft: W-Malaysia, Tapah Hills [coll. FH, No. 0018-38]; 1 right ♀ ala: ex Zucht F. Hennemann 1996, Herkunft: W-Malaysia, Tapah Hills [coll. FH, No. 0018-39]; 5 ♀♀: West-Malaysia, Perak, Tapah Hills, ca. 500 m, 07.1996, via Wong Tet Fatt [coll. OC]; 1 ♀: West-Malaysia, Perak, Cameron Highlands, Kampong Raja, 02.2002, via Chaminade [coll. OC]; 8 ♀♀: West-Malaysia, Perak, Tapah Hills, ca. 500 m, local collector [coll. OC]; 1 ♀: West-Malaysia, Perak, Tapah Hills, ca. 500 m, 05.2003, via Wong Tet Fatt [coll. OC]; 3 ♂♂, 3 ♀♀: Ex Zucht O. Conle, 1995, Herkunft: WestMalaysia, Perak (PSG No. 18) [coll. OC]; 4 ♂♂, 1 ♀ (nymph): Ex Zucht O. Conle, 2001-03, Herkunft: WestMalaysia, Perak (PSG No. 18) [coll. OC]; 13 ♂♂: Ex Zucht O. Conle, 2005-08, Herkunft: West-Malaysia, Perak (PSG No. 18) [coll. OC]; 1 ♂, 1 ♀: Ex Zucht Kristien Rabaey 2006, Herkunft: West-Malaysia, Perak (PSG No. 18) [oll. OC]; 3 ♂♂, 3 ♀♀: W-Malaysia, Perak, Tapah Hills [coll. FSC]; 1 ♂: Tasek Chini, leg FSC [coll. FSC]; 2 gynadromorphs: W-Malaysia, Perak, Tapah Hills, leg Michael Yeh [coll. FSC].


HENNEMANN ET AL.

S-THAILAND: 1 ♂, 2 ♀♀, 1 ♀ (penultimate instar), 1 ♀ (nymph N5), eggs: S-Thailand, Phuket, leg Stobbe, 1997 (coll. FH, No’s 0018-32 to 36 & E3]; 2 ♂♂, 2 ♀♀: ex Zucht O. Conle, 1999–2000, S-Thailand [coll. OC]. SUMATRA: 1 ♀: Sumatra; Heteropteryx dilatata (Park.) det. Günther [RMNH]; 1 ♀: Deli, Sumatra [NHMM]. SINGAPORE: 1 ♀: Coll. Br. v. W., Singapore, B. Bader; 14.228 [NHMW, No. 279]. NO / UNPRECISE DATA: 1 ♀: Indoaustralisches Archipel, Veronique De Decker, I.G.: 2.8.271 [RBINS]; 2 ♀♀: bred by N. Mal (ex larva) [RBINS]; 1 ♂: bred by W. Potvin, 1994 (RBINS]; 1 ♀ (nymph): 1904; Heteropteryx dilatata (Parkinson) det. Bragg [RMNH]; 1 ♀: ex Zucht M. Heklau, Stuttgart, 1992 [SMNS]. Etymology: The name (dilato lat. = widened) refers to the strongly laterally dilated abdominal tergites and hence very broad, leaf-like abdomen of ♀♀. Description: ♀♀ (Figs. 323–325). Very large (body length 133.0–166.0 mm) and massive Heteropteryginae, body of leaf-like appearance with mesothorax strongly widening and trapezoidal and abdominal tergites II–VII strongly dilated and armed with prominent spines dorsally. General colour bright apple green dorsally and mid to dull green ventrally. Sometimes partly or fully yellow forms occur (Fig. 324). Legs bright green (or yellow), darker ventrally and the tibiae with a slight ochre to pale brown wash. Head and body armature bright red and all larger spines with the apex black; coronals and spines on the dorsal body surface ± blue to turqoise basally. Tegmina bright apple green, but may be green with yellow mottling in yellow specimens. Alae transparent pink with the apex green (Fig. 84). Posterior surface of pro- and mesocoxae and interior surface of metacoxae either pink (Fig. 335) or black (Fig. 336) basally. Head slightly longer than wide, higher than long and with vertex strongly convex, conically elevated and slightly projecting over anterior margin of pronotum (Fig. 339). Vertex with four very prominent outward pointing and roughly equally sized coronals, the anterior coronals somewhat more distant than the posterior pair (Fig. 337). Often with a pair of moderately distinct supra-occipitals and occasionally with a minute pair of supra-antennals, but the latter mostly obsolete. Lateral coronals considerably smaller than the anterior and posterior coronals. Four almost equally sized supa-orbitals present, which are ± similar in size to the pair of supra-occipitals. Antennae with 28 antennomeres. Pronotum trapezoidal with posterior margin considerably wider than anterior margin and with moderately sized anterior and posterior-pronotals, the inter-posterior pronotals of similar size but variable in length (Fig. 337). In front of the short V-shaped transverse median sulcus occasionally with a fairly small pair of premedian pronotals (Fig. 339); these often obsolete or sometimes only one spine present. Sensory area of profurcasternum weakly developed and covering almost entire anterior portion of segment. Mesothorax strongly, gradually widening towards the posterior and trapezoidal in shape; posterior portion about 2x wider than anterior portion (Fig. 337). Mesonotum roughly rectangular and some 1.4x longer than pronotum; lateral margins with four somewhat subequally sized spines, the disc usually with a pair of anterior mesonotals, a pair of post-median mesonotals and a somewhat smaller pair of less distant pre-median mesonotals (Figs. 337, 339). Between the bases of the tegmina with a fairly strong pair of posteromedial mesonotals (Figs. 337, 339). Tegminal moderately distinct. Mesopleurae with a fairly small supra-coxal and 8–10 somewhat unequally sized, sometimes slightly back-curving laterals along lower margin (fig. 337). Mesosternum with 6–8 small, paired spines and a small pair of inter-posterior mesosternals. Metasternum with 6–7 short spines on each side of the median line. Metapleurae strongly deflexed with the lower lateral margin distinctly rounded and armed with about eight somewhat unequally sized spines; supra-coxal small. Tegmina large, oval in shape and reaching ± half way along abdominal tergum II; radial vein raised and forming a prominent longitudinal keel. Abdominal tergites II–III widening, IV widest and V– X narrowing. II–VII strongly dilated, much wider than long with lateral margins strongly laterally deflexed and armed with five short spines. Lateral margins of tergum VIII only with 3–4 very minute teeth. Dorsal surface of II– IV unarmed, V–VIII with a very distant pair of spines near anterior margin and much smaller pair of median spines close to median line; these most prominent on V and gradually decreasing in size towards VIII. Often a further pair of much small spines close to posterior margin. V–X with a longitudinal median furrow, which is laterally bordered by a fine carina. Sternites with twofour pairs of short spines. Praeopercular organ on sternum VII formed by a distinct pair of spines at posterior margin and a curved carina inbetween (Fig. 331). Anal segment with posterior



99

margin rounded and gently notched medially (Fig. 330). Epiproct considerably longer than anal segment and distinctly projecting beyond apex of subgenital plate with the basal portion gradually narrowing and the apex deeply incised and bifid; median portion may be somewhat constricted (Figs. 329–330). Cerci small, conical and oval in cross-section. Subgenital plate strongly keeled with the apex broad, roundly truncate and ± indented (Fig. 331). Legs long and moderately slender, the metafemora laterally compressed and almost 2x higher than wide. Mesofemora longer than mesothorax, metafemora ± reaching to posterior margin of abdominal segment VI and metatibiae projecting considerably beyond apex of abdomen. All carinae of front and mid legs sparsely but distinctly dentate, the most apical tooth on the posterodorsal carina of the femora decidedly enlarged. Metafemora with all carinae densely spinose/dentate (dorsal carinae in particular). Two outer ventral carinae of metatibiae armed with about seven spines, which distinctly increase in size and become increasingly incurving towards apex of tibia; the most apical spine largest (Fig. 377). Ventral surface of femora smooth (Fig. 356). ♂♂ (Figs. 326–328). Fairly large (body length 81.5–98.0 mm), stocky Heteropteryginae with very long tegmina and alae and a very short mesothorax. General colour pale to dark, sometimes with a faint dark brown longitudinal median line on pro- and mesonotum. Abdominal tergum VI sometimes pale brown to dull cream. Head with a ± distinct black postocular stripe (Fig. 340). Cephalic and thoracic armature brown basally, then red and with the tips black. Tegmina pale to dark brown and with a ± distinct pale diagonal marking or stripe, sometimes with faint cream markings and occasionally with a white sub-median spot; anterior margin pale cream or yellow to white. Costal region of alae transparent brown with all veins dark brown, the anal fan dull pink to dull red with brown veins (Figs. 326–327). Shape and armature of head as in ♀♀ but all spines much longer and more slender (Figs. 338, 340). Antennae with 26 antennomeres. Pronotum somewhat more elongate and less trapezoidal than in ♀♀ with the median portion constricted; armature as in ♀♀. Sensory area of profurcasternum more distinctly raised than in ♀♀. Mesothorax short and only some 1.1–1.2x longer than pronotum, the anterior half roughly parallel-sided and the posterior half gradually widening with posterior portion some 1.8x wider than anterior margin. Mesonotum very slightly narrowing towards the posterior with the disc before the posteromedial spines and bases of the tegmina almost quadrate and the posterior portion strongly raised (Fig. 340). Lateral margins unarmed and disc with the six spines arranged as in ♀♀ but very minute to obsolete (Figs. 338, 340). The pair of posteromedial mesonotals considerably more pronounced than in ♀♀ but just slightly longer than the distinct tegminal spine (Fig. 340). Mesopleurae with six slender and distinct, almost equally sized laterals along lower margin and a fairly small supra-coxal spine (Fig. 338). Metapleurae as in ♀♀. Mesosternum with a very fine longitudinal median carina and 4–5 short spines on each side. Metapleurae set with 5–6 fairly distinct spines on each side of the median line. Tegmina very elongate, slender, roughly parallel-sided, more than ¾ the length of the alae and reaching to abdominal segment VII (Figs. 326–328); radial vein very prominent, raised and forming adistinct, tectiform longitudinal keel. Sub-basally with a prominent, laterally compressed to roundly conical hump. Alae very long and ± reaching to apex of abdomen (Figs. 326–328). Abdomen long and slender with segments II–VI very slightly narrowing, VII–VIII widening and IX widest segment; all roughly equal in length and almost as wide as long. Dorsal surface of all tergites smooth but VII–X with two very fine longitudinal median carinae. Lateral margins of II simple, of III–IX deflexed and protuded into a dentate, foliaceous lobe in posterior half, which is slender or indistinct in III, gradually increases in size towards VI, is small on VII and again very prominent in VIII and IX. Lobe of tergum IX roughly semicircular and extending laterally by almost half the width of segment (Fig. 333). All with about five teeth, four of which a restricted to the lobe and are more prominent than the anterior tooth. Sternites II–VII armed with eight paired spines, which are most prominent on II and gradually decrease in size towards VII. Anal segment a little wider than longwith lateral margin sub-basally expanded into a small, bi-denatate lobe (Fig. 334), the posterior margin with a shallow, rounded excavation (Fig. 333). Epiproct short and roundly angular to semicircular in shape (Fig. 333). Cerci fairly long and compressed laterally with the apex club-like (Fig. 404). Vomer broad with basal portion almost semicircular and the short and slender terminal hook directed dextrally (Fig. 404). Poculum large, very bulgy, somewhat narrowed towards the apex with the posterior margin labiate and roundly truncate (Fig. 334). Legs generally as in ♀♀ but the sub-apical tooth on the posterodorsal carina of the femora more pronounced and contrasting black. Armature as in ♀♀ but averaging more prominent. Mesofemora almost as long as pro- and mesonotum combined, metafemora reaching to abdominal segment VI and metatibiae projecting somewhat beyond apex of abdomen. Ventral spines of the metatibiae less curved than in ♀♀, almost straight and somewhat backward directed (Fig. 378). Ventral surface of all femora smooth (Fig. 357).


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Variability: Females are typically bright green but in the wild and also in captivity yellow specimens occur spontaneously (Brock, 1999; Bragg, 1998; Chan & Lee, 1994). The yellow colour of these ♀♀ appears to be caused by a recessive allele (Bragg, 2001: 101) and eggs from yellow specimens usually produce yellow and green offspring, but never in a ratio higher than one yellow to three green specimens. While the yellow colour-form is rarely encountered in Peninsular Malaysia it is much more common amongst specimens found in Phuket, South Thailand. From 36 live ♀♀ the first author received from that locality in 1997 as many as fifteen were yellow. Specimens of this Thailand colony are furthermore characteristic for having the inner surfaces of the meso- and metacoxae and ventral surface of the procoxae with a washed black marking, instead of pale pink as in specimens from Peninsular Malaysia or Borneo (Fig. 336). Morphologically there is little variability in ♀♀, which mainly is restricted to the size and development of the head and body armature. Males show somewhat more variability in both colouration, spination and relative length of the wings. The alae usually reach to or project over the apex of abdomen but sometimes may only reach to abdominal segment IX. The colouration ranges from pale to very dark brown, but specimens usually show a tendency to darken with rising age. The pale diagonal marking in the basal half of the tegmina may be faint to almost obsolete in some specimens and sometimes there is a washed pale marking apically. Nymphs show some variation in aspect of their colouration. While they are usually more or less plain brown, sometimes specimens occur which have pretty white markings all over the body or even a bold longitudinal white stripe along the entire dorsal body surface. These however disappear once the specimens reach maturity. Nymphs: The sex of nymphs can already be determined when they are newly hatched, ♀♀ having traces of the elongated epiproct while ♂♂ have not. The first four instars of ♀♀ are brown and the typical green colour is only achieved from 5th instar onwards. While ♀♀ already possess they typical body shape, ♂♂ look fairly unlike mature specimens, from 2nd instar onwards having all the abdominal tergites with broad and dentate lobes. These are also present in the adults but relatively smaller and folded downwards. Furthermore, the mesothorax is becoming increasingly shorter in ♂♂ during the nymphal development. Both sexes have the legs relatively shorter, more stocky and with the armature considerably stronger developed. Eggs (Figs. 441–444, 447): Large, barrel-shaped with polar-area rounded and capsule slightly narrowing towards the anterior; roughly 1.5x longer than wide. Capsule surface very rough and uneven with numerous irregular impressions and wart-like swellings; all over covered with very short hairy structures. Portion between micropylar plate and polar-area with two roughly parallel-sided, elongate swellings. Micropylar plate basically Xshaped with all four extensions fairly long and slender; anterior extensions at an angle of about 85° to each other with the apex gently incurving; posterior extensions with the upper portion of the rounded apex slightly expanded. Surface of plate similar to capsule but texture less decided and entire plate surrounded by a shallow bulge. Operculum almost circular and slightly convex, structured like capsule. Micropylar cup an almost circular impression with a small C-shaped carina posteriorly. General colouration plain dull grey, the outer margin of the micropylar plate dark brown. The shape of the micropylar plate shows some variability. Eggs at hand from S-Thailand (Phuket) however frequently differ from Malaysian stock by having the anterior extensions of the plate shorter and gradually narrowed towards the apex and the posterior extensions less expanded at the apex. Furthermore, the outer margin of the plate is less contrasting dark brown and hardly darker in colour than the capsule (Figs. 443–444). Measurements [mm]: length incl. operculum 8.1–9.8, length 7.9–9.2, width 5.1–5.2, height 5.6–5.8. Comments: This impressive species is the largest representative of the entire Heteropterygidae with females reaching body lengths of up to 166.0 mm. It is one of the most striking extant Phasmatodea known and with a maximum weight of up to 51.2 grams (Beccaloni, 2010) ♀♀ in full egg-production are perhaps the heaviest phasmids known and only exceeded by certain large Coleoptera, e.g. some rhinoceros beetles of the subfamily Dynastinae and goliath beetles of the subfamily Cetoniinae. H. dilatata is a very common and well known species in Peninsular Malaysia and frequently sold as live or preserved specimens by entomological suppliers, local souvenir outlets or Butterfly farms. It is also not unfamiliar to the local Orang Asli people, who call it “Duri”, meaning “Thorny” in the Malay language (Chan & Lee, 1994: 5). H. dilatata has been successfully reared in Europe since 1974 and because of its impressive size, beautiful colour and striking spination has become one of the most sought after phasmids ever to be cultured in Europe. Frequently, specimens are on display at entomological exhibitions or in zoological gardens around the world. Hence, numerous papers have been published mostly dealing with its biology, behaviour or breeding (e.g. Löser & Schulten, 1981; Herbert, 1986; Chan & Lee, 1994; Schulten, 1995). Brock (1999: 144) also provided information on historical facts.



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H. dilatata was originally recorded from “Asia” by Parkinson in 1789 and is a common species in Peninsular Malaysia even being extremely numerous in some localities, e.g. in the foothills of the Cameron Highlands (Chan & Lee, 1994: 5). It has additionally been recorded from Singapore, although it has not been found there recently. Rehn (1904: 89) recorded it from Borneo but from an unspecified area. Bragg (1998: 29; 2001: 99) however listed three specimens from NW-Borneo (Matang, Mt. Serapi and Entawa). The fourth author found two ♂♂ and two ♀♀ in Kubah National Park, hence the distribution in Borneo can be confirmed and H. dilatata is the only representative of Heteropteryginae that is found in Peninsular Malaysia and Borneo at the same time. It is however possible that H. dilatata was imported to Borneo by man, as it was often kept as pets by Chinese families who used the droppings or eggs as medicine against diarrhoea and gastroenteritis (Bragg, 1990b; Chan & Lee, 1994). More recently it has also been recorded from northern Thailand (Sorpongpaisal & Thanasinchayakul, 2006) and the two first authors collections contain specimens collected in Phuket, southern Thailand. The fourth author frequently encounters this species on Pulau Penang and Phuket. The original description of H. dilatata by Parkinson (1798) is remarkably detailed and includes colour illustrations of the ♀ holotype and egg. The egg of H. dilatata was the first ever to be described from the order. In 1806 the holotype was sold at auction to W. S. Macleay for £ 11.20s., a significant sum at that time; the specimen is deposited in the MAMU in Sydney, Australia. Due to the significant sexual dimorphism and the fact that nymphs look very unlike adult insects, several synonyms have been described all of which have been clarified by previous authors. The holotype of Heteropteryx castelnaudi Westwood, 1874 in UMO is an immature ♂, the holotype of Phasma (Eurycantha?) graciosa Westwood, 1848 in UMO is an adult ♂, and the holotypes of Heteropteryx hopei Westwood, 1864 in UMO and holotype of Heteropteryx rollandi Lucas, 1882 are both adult ♀♀ of H. dilatata. Gynandromorphism is more striking and more commonly found in H. dilatata than in any other species of the order. Gynandromorph specimens have been mentioned in the literature numerous times already (e.g. Brock, 1989; Brock, 1999; Seow-Choen, 2000b) and Seow-Choen (1996: 122) and Brock (1999, pl. 30) provided colour photographs. Seow-Choen (1996) described a mostly male gynandromorph specimen, out of whose abdomen several fully developed eggs were taken, that did not differ from those laid by normal ♀♀. Brock (1999: 144) and Seow-Choen (2000b: 40) provided lists of native foodplants which include Durio zibethinus (Bombacaceae), Eugenia aquea (Myrtaceae), Grewia acuminata (Tiliaceae), Uncaria sp. (Rubiaceae), Psidium guayava (Myrtaceae) and Rubus moluccanus (Rosaceae) (→ 7.6). In captivity H. dilatata feeds on a variety of plants including bramble (Rubus fruticosus, Rosaceae), raspberry (Rubus idaeus, Rosaceae), rose (Rosa spp., Rosaceae), oaks (Quercus spp., Fagaceae), salal (Gaultheria shallon, Ericaceae), hawthorn (Crataegus laevigata, Rosaceae), rowan (Sorbus spp., Rosaceae), cherry (Prunus spp., Rosaceae), apple (Prunus spp., Rosaceae), pear (Pyrus spp., Rosaceae), beech (Fagus sylvatica, Fagáceae), Japanese knotweed (Fallopia japonica, Polygonaceae), rhododendron (Rhododendron spp., Ericaceae), cotoneaster (Cotoneaster spp., Rosaceae), cherry laurel (Prunus laureocerasus, Rosaceae) and ivy (Hedera helix, Araliáceae). It is fairly easy to rear in large cages and at high humidity, which should include a daily spray with water, since the ♀♀ in particular need plenty of water for the egg-production. As for all members of the Heteropteryginae a substrate is necessary to allow ♀♀ to bury their eggs. Further information is provided below (→ 7.5). Distribution (Fig. 497): Peninsular Malaysia: Perak [MIZT, ZMUH]; Perak, Tapah Hills [coll. FH, coll. FSC, coll. OC]; Perak, Cameron Highlands, Tanah Rata 1500 m [MHNG, coll. FH]; Perak, Cameron Highlands, Parit waterfalls [Brock, 1999: 144]; Perak, Cameron Highlands, Perak, Gunong Jasar [Brock, 1999: 144]; Perak, Wrays Hill [RMNH]; Perak, Belum [Brock, 1999: 144]; Perak, Kuala Who [Brock, 1999: 144]; Perak, Pulau Penang, Butu Ferringhi [Brock, 1999: 144]; Perak, Pulau Penang [UMO, coll. FSC]; Perak, Kijang Falls [photo taken by the fourth author, Fig. 486]; Selangor [MHNG]; Selangor, Bukit Kutu 3000 ft. [RMNH]; Selangor, Bukit Lagong [Brock, 1999: 144]; Selangor, Kuala Lumpur [Brock, 1999: 144]; Selangor, Kuala Sleh [Brock, 1999: 144]; Pahang [RMNH]; Pahang, Lubok Tamang 3500 ft. [RMNH]; Pahang, Tanam Camp, 18 mile Railway track [BPBM]; Pahang, Tasek Chini [coll. FSC]; Kelantan [Brock, 1999: 144]; no precise data [MNHU, MHNG, coll. PEB]; Malacca [MNHN, NHMW, NHMB, ZMUH]; Johor, Endau Rompin [Chan & Lee, 1994: 5]. Singapore [NHMW]. S-Thailand: Trong [Brock, 1999: 144]; Phuket [coll. FH, coll. OC]. N-Thailand [Sorpongpaisal & Thansinchayakul, 2006: 22]. Sumatra: NE-Sumatra, Prov. Sumatera Utara, Medan (= Deli) [NHMM]; E-Sumatra, Prov. Riau, Indragiri Hulu, Indragiri River [NHMB]; “Sumatra” [NHMW, RMNH]. Borneo: NW-Sarawak: Mount Serapi 150 m [coll. PEB]; Matang [SMSM]; “Entawa” [SMSM]. Specimens examined: 122


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TABLE 19. Measurements [mm] of Heteropteryx dilatata (Parkinson, 1798) ♂♂

♀♀

Body

81.5–98.0

133.0–166.0

Head

6.7–7.8

12.5–16.5

Pronotum

6.8–7.7

10.4–14.5

Mesonotum

7.0–8.9

17.6–21.0

Metanotum + Median Segment

15.0–17.5

19.0–22.0

Tegmina

52.0–55.0

36.5–46.5

Alae

62.0–71.0

35.0–38.5

Profemora

20.0–23.0

21.5–31.0

Protibiae

22.0–24.8

30.0–35.0

Mesofemora

16.7–19.5

22.0–26.7

Mesotibiae

16.2–19.0

25.0–30.5

Metafemora

24.0–28.0

32.5–40.5

Metatibia

26.5–29.2

39.4–49.3

Antennae

76.0–85.0

78.5–101.0

5. Arguments for the inter- and intrageneric systematization of Heteropteryginae As discussed above (→ 3.4) and illustrated in Figure 97b there is good evidence that Heteropteryginae is the most derived subordinate clade of the family Heteropterygidae. This is supported by autapomorphies such as the secondary reduction of the paired sensory-areas on the prosternum, secondary gain of wings and frequent presence of a stridulatory organ in the basal portion of the alae of both sexes and straight profemora as well as the saltire or X-shaped micropylar plate and lack of a median line in the eggs. Synapomorphies shared with its sister-group, the subfamily Obriminae, are the beak-like secondary ovipositor of ♀♀, that is formed by an elongated epiproct and subgenital plate, and the presence of a medio-apical spine on the area apicalis of all six tibiae (Figs. 93–96, 97b). These results are in compilance with previous molecular and phylogenetic studies (e.g. Bradler, 2009, bradler et al., 2014, 2015). A full phylogenetic analysis of the Heteropteryginae still deserves knowledge of the unknown sexes and eggs of certain species and a more comprehensive treatment of its sister-group, the Obriminae. Furthermore, molecular data appear necessary to fully clarify all intrageneric relationships with confirmation. Nevertheless, a first attempt towards a phylogentic analysis seems warranted, and although somewhat speculative and hypothetical in some points, some hypotheses seem justified by their heuristic value and are meant to be a starting point for future studies (Fig. 355). The fact that the present day distributions of all three subfamilies of Heteropterygidae overlap on Borneo suggests the Bornean representatives of Heteropteryginae, i.e. the species here attributed to the grayii speciesgroup (→ 4.1.2) and echinata species-group (→ 4.1.3) of Haaniella Kirby, 1904, may be the basalmost taxa of the subfamily. This hypothesis is supported by several morphological characters of the insects and eggs. The most basal clade of Heteropterygidae is here revealed to be represented by the predominantly ground-dwelling Dataminae (→ 3.4). On the basis that Heteropteryginae evolved from mostly ground-dwelling phasmids and has a much smaller distributional range than the other two subfamilies (Fig. 342), that apart from a single species in the south of Vietnam is restricted to Sundaland (Fig. 345), some conclusions can be drawn on which characters might be plesiomorphic or apomorphic respectively. The monotypical Heteropteryx Gray, 1835 is the only representative of the subfamily that shows obvious adaptions to arboreal habits, hence seems to be the most derived taxon of Heteropteryginae. This hypothesis is supported by the bright green or yellow colouration of ♀♀, whereas all species of the second known genus, i. e. Haaniella, are exclusively various shades of brown. Besides the colouration, the strongly laterally deflexed and



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dilated abdominal tergites of ♀♀ of Heteropteryx give the insects a leaf-like general appearance and sufficiently provide for a good camouflage in foliage. Moreover ♂♂ of Heteropteryx have fully developed wings that cover the entire abdomen and make the insects capable of short active flight, which increases mobility and also supports arboreal habits. In fact, in its natural habitats Heteropteryx is known to live in shrubs or trees often several metres off the ground and ♀♀ will only descend from their host-plants for oviposition (→ 7.1). The peculiar morphological adaptions to arboreal habits, such as the green or yellow colouration and strongly dilated abdominal tergites of ♀♀ as well as the very long tegmina and alae of ♂♂ are autapomorphies of Heteropteryx. Further autapomorphies are the very elongate antennae of both sexes, the strongly shortened mesothorax of ♂♂, which is hardly longer than the prothorax (Figs. 338, 340), the presence of dentate lateral lobes on abdominal tergites II–IX in ♂♂ and the pink (♀♀) or dark red (♂♂) alae (Figs. 323, 326–327). The strongly conical vertex of both sexes in contrast seems to be plesiomorphic, since this character is also frequently found throughout various members of Dataminae and several genera of Obriminae. In contrast to the arboreal habits of Heteropteryx most species of Haaniella are mostly found on low vegetation but rarely high up in trees (→ 7.6). This is proven by numerous observations in the field carried out by the authors. In the strict sense however, Haaniella-species can not be regarded as true ground-dwelling Phasmatodea like Dataminae or supposedly basal Obriminae like the genera of the tribe Tisamenini n. trib. (→ 3.2.2). On the basis of external morphology of the insects and egg morphology and geographic distribution, Haaniella divides into three well marked species-groups (Table 1). A first attempt towards an intrageneric systematization of Haaniella is presented in the preliminary tree in Figure 355 but must in several aspects be regarded as speculative, since definite apomorphies are difficult to determine without a full phylogenetic treatment of the genus or extensive sets of molecular data also of the still unknown sexes and eggs of some species. The ten known species of the muelleri species-group are distributed throughout Sumatra and nearby islands, Singapore and Peninsular Malaysia and are more closely related to Heteropteryx than are the species of the other two other predominantly Bornean species-groups. The muelleri species-group is not only the most speciose group of Haaniella but also exhibits the greatest morphological diversity. It is the sister-group of the grayii species-group + echinata species-group (Fig. 355) and differs from these by a variety of characters. An autapomorphy of the eggs is the secondary reduction of setae, the capsule surface and operculum being granulose or rugulose and lacking any setae or hairy structures as in the other two species-groups of Haaniella or the genus Heteropteryx. Furthermore, eggs of the muelleri species-group are more or less lemon-shaped, having the operculum and often also the polararea conically raised to variable degrees (Figs. 412–419). Although this character may a priori appear to be an autapomorphy of the muelleri species-group, this distinctive shape of the eggs is also found in one species of the echinata species-group (i.e. H. saussurei Kirby, 1904; Figs. 431–432). Close relation to Heteropteryx is supported by the increasingly elongated antennae, which reach as far as to abdominal segment V in ♀♀ or are almost equal in length to the body in ♂♂, smooth ventral surface of the femora (Figs. 358–361), occasional presence of lateral lobes on abdominal segments VII–IX in ♂♂, laterally deflexed and lobed abdominal tergites IV–IV in ♀♀ of some species and the obsolete lateral invaginations of abdominal sternites II–IV. Moreover ♂♂ of some species resemble Heteropteryx by having more or less elongated wings or even full-sized alae, which cover most of or almost the entire abdomen, this is H. aculeata n. sp., H. glaber (Redtenbacher, 1906), H. macroptera n. sp. and H. mecheli (Redtenbacher, 1906). While the obsolete lateral invaginations of the abdominal sternites quite certainly is a symplesiomorphy of Heteropteryx + muelleri species-group, at least some of the aforementioned characters may be synapomorphies. The relationships within the muelleri species-group are difficult to clarify in a whole without a full phylogenetic study or sufficient molecular data, but some relationships or even sister-group relationships between certain species appear fairly obvious. A sister-group relationship is estimated between the two Sumatran H. glaber (Redtenbacher, 1906) and H. gintingi n. sp., which share several common characters that distinguish them from all other members of the muelleri species-group. The conspicuously enlarged composite anterior coronals and broad, triangular and contrasting black ventral teeth of the metatibiae of both sexes (Figs. 373–376) are only present in these two species and not seen elsewhere throughout the muelleri species-group or the entire genus. Both species furthermore share the reduced body armature, distinctly laterally deflexed abdominal tergites IV–V of ♀♀, elongate and slender mesothorax of ♂♂ and distinct triangular sub-apical tooth on the posterodorsal carina of the femora in both sexes. Another sister-group relationship seems obvious between the Malayan H. erringtoniae (Redtenbacher, 1906) and H. macroptera n. sp. from Singapore and southern Peninsular Malaysia, these two species merely differing by genitals structures of both sexes, minor differences in the thoracic armature


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and the egg morphology. Interestingly, in both these presumed sister-group relationships one species has micropterous ♂♂ while ♂♂ of the adelphotaxon are macropterous. The two Sumatran species H. muelleri (Haan, 1842) and H. rosenbergii (Kaup, 1871) have micropterous ♂♂ and are obviously very closely related, hence are also possible sister-taxa. Close relation of these two species to H. erringtoniae (Redtenbacher, 1906) and H. macroptera n. sp. is obvious in serveral aspects. The only representative of the genus from Simeulue Island some 150 km off the west coast of Sumatra, namely H. jacobsoni Günther, 1944, and the fairly small but distinctly spinose H. kerincia n. sp. from western Sumatra share the strongly deflexed and lobed abdominal tergites IV–VI of ♀♀ and presence of large lateral lobes in abdominal tergites VII–IX of ♂♂, which readily distinguish them from all other members of the genus. Hence, these two species appear to be closely related as well and are likely to be sister-taxa. While H. jacobsoni is characteristic for having very slender, not overalapping wings and a distinct pair of spines between those on the metanotum and median segment (Figs. 153–156), H. kerincia n. sp. is peculiar for its extremely long cephalic and thoracic spines and having the pre-median pronotals and posteromedial mesonotals dichotomic, composite or antler-like (Figs. 164–165). Both features are unique within the entire genus and certainly represent autapomorphies of these two species. The systematic position of the two Sumatran H. aculeata n. sp. and H. mecheli (Redtenbacher, 1906) is difficult to decide without knowledge of the still unknown ♀♀. Males of both species are macropterous and appear to be close to H. macroptera n. sp. and H. erringtoniae (Redtenbacher, 1906). The three known species of the echinata species-group, H. echinata (Redtenbacher, 1906), H. saussurei Kirby, 1904 and H. scabra (Redtenbacher, 1906), are endemic to Borneo and more closely related to the partly Bornean grayii species-group than to the muelleri species-group, which is not at all represented in Borneo. This is supported by common characters such as the presence of a median row of spines on the ventral surface of the femora (Figs. 364–365), distinct lateral invaginations of abdominal sternites II–IV (Figs. 85, 87) and mostly specialized apex of the epiproct of ♀♀ (Fig. 355). All three characters seem to be apomorphic states. Furthermore, the densely setose eggs readily distinguish these two species-groups from the muelleri species-group, but this must be regarded as a plesiomorphic since hairy structures in the eggs are also present in the subfamily Dataminae (→ 3.1 and Fig. 97b). Females of the echinata species-group differ from those of the grayii speciesgroup by the remarkably shortened epiproct, which does not reach the tip of the subgenital plate and has the apex angular and multi-denatate. The subgenital plate bears a blunt angular projectiong medio-basally, has the apex rounded to slightly pointed and never notched as in members of the grayii species-group. Males readily differ from those of the grayii species-group by lacking a central pair of spines on abdominal tergites II–V and having the lateral margins of the anal segment sub-basally protruded into a triangular tooth. Furthermore, there are never distinct pre-median mesonotals as in members of the grayii species-group. A sister-group relationship is obvious between H. echinata (Redtenbacher, 1906) and H. scabra (Redtenbacher, 1908), which share the presence of interposterior pronotals and enlarged antero-lateral mesonotals in both sexes as well as the gradually tapered abdominal segments II–V and small, rounded epiproct of ♂♂. Eggs of both species are barrel-shaped with the polar-area rounded and the operculum just very slightly conical (Figs. 433–440). While H. scabra is geographically restricted to the Mount Kinabalu massive and surrounding mountains in Northern Sabah and altitudes above 800 metres (Fig. 496), H. echinata is widely distributed in lower elevations throughout northern and eastern Borneo and has also been recorded from Labuan Island off the northern coast of Sabah (Fig. 495). The sister-group of H. echinata + H. scabra is H. saussurei Kirby, 1904, which differs from both species by the lack of inter-posterior pronotals and not enlarged antero-lateral mesonotals of both sexes, as well as the almost parallel-sided abdominal tergites II–VII and strongly elongated and slender epiproct of ♂♂, which distinctly projects over the apex of the abdomen (Figs. 296– 298). At least the elongated epiproct may be regarded as an autapomorpyh of H. saussurei, but is also seen in some species of the muelleri species-group, hence seems to be homoplastic. Furthermore, the eggs of H. saussurei readily differ from those of H. echinata and H. scabra by being distinctly lemon-shaped with the polar-area and operculum strongly conical and having the micropylar plate of a very distinctive shape (Figs. 431–432). The shape of the eggs resembles that seen in the muelleri species-group. The distribution of H. saussurei appears to be restricted to coastal or coastal near regions in the very northwestern portion of the island (Fig. 496). The intraspecific relationships within the grayii species-group are much less obvious than in the echinata species-group. Haaniella dehaanii (Westwood, 1859) and H. grayii (Westwood, 1859), are endemic to Borneo, H. parva Günther, 1944 occurs only in northern Sumatra and H. gorochovi n. sp. has a remarkably disjunct distribution being the only representative of the entire genus found in southern Vietnam (Fig. 493). The species of the grayii species-group are more closely related to those the echinata species-group rather than those of the



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muelleri species-group, which is supported by common characters such as the presence of a median row of spines on the ventral surface of the femora (Figs. 362–363), distinct lateral invaginations of abdominal sternites II–IV (Figs. 86, 88) and mostly specialized apex of the epiproct of ♀♀ (Fig. 355). Males of all four species are characteristic for having a prominent pair of central spines on abdominal tergites II–V, which readily distinguishes them from all other species of Haaniella. The anal segment lacks the sub-basal lateral tooth seen in ♂♂ of the echinata species-group. In contrast to the echinata species-group the epiproct of ♀♀ is elongate and strongly projects over the apex of the subgenital, the latter of which is variable in shape and ranges from rounded over angular to having a distinct notch medially. While ♀♀ of the two Bornean species have the apex of the epiproct multi-dentate, it is simple and bifid in H. parva Günther, 1944 and H. gorochovi n. sp.. Hence, the epiproct of the latter two species resembles that seen throughout members of the muelleri species-group. The eggs of all four species are more or less distinctly setose and barrel-shaped with the operculum flat to gently convex but never conical as in members of the echinata species-group (Figs. 420–427). The distribution in Borneo, presence of distinct pre-median mesonotals in both sexes and the specialized epiproct of ♀♀ suggest a sister-group relationship between H. grayii (Westwood, 1859) and H. dehaanii (Westwood, 1859). The lack of pre-median mesonotals and simple bifid apex of the epiproct of ♀♀ at the other hand support close relation between the Vietnamese H. gorochovi n. sp. and the Sumatran H. parva Günther, 1944, which however deserves further evaluation, since biogeographic arguments rather support close relation to the Bornean taxa for H. gorochovi n. sp. (→ 6.3). It is hoped that in future phylogenetic studies and molecular data can add to our knowledge on the intrageneric relationships of Haaniella and reveal all sister-group relationships, some of which have been suggested above but may still be somewhat speculative. Certainly, knowledge of the still unknown ♀♀ of H. aculeata n. sp. and H. mecheli (Redtenbacher, 1906) as well as the so far unknown eggs of H. aculeata n. sp., H. glaber (Redtenbacher, 1906), H. jacobsoni Günther, 1944, H. mecheli (Redtenbacher, 1906) and H. muelleri (Haan, 1842) will help in completing our current knowledge on the intrageneric systematization of Haaniella. In due course expeditions throughout Sumatra seem much warranted.

6. Biogeography The subfamily Heteropteryginae, a subordinate taxon of the Heteropterygidae, is a typical faunistic element of the Oriental region (→ see definition below) and includes some of the most striking, spiniest and heaviest representatives of the phasmid fauna of that region, representing some of the world’s most massive and heavy phasmids. For instance ♀♀ of Heteropteryx dilatata (Parkinson, 1798) can reach maximum weights of 51,2 grams when in full egg production. All of the known taxa are distributed throughout Sundaland or Neomalaya to be more precise (Peninsular Malaysia, Singapore, Sumatra, and Borneo) but are absent on Java and the Philippines. Interestingly, the genus Haaniella Kirby, 1904 has one species in Vietnam, a disjunct but not unusual distribution that is discussed below. In addition to the distribution of the subfamily Heteropteryginae, information is also provided on the distributional patterns of the other two subfamilies of Heteropterygidae, namely Dataminae and Obriminae. Hence, some background information on the geological history and biogeography of Australasia and Southeastern Asia appears warranted at this stage to better understand the distributional patterns of the Heteropterygidae and its subordinate taxa. The Oriental region comprises continental Asia south of the Palearctic Region, the Large Sunda Islands (Sumatra, Java and Borneo as well as their satellite islands), the Philippines, Sulawesi and the Lesser Sunda Islands as far east as Timor. The eastern portion of the Oriental region, which includes Sulawesi and the Lesser Sunda Islands except Bali, biogeographically belong to a zone of transition to the Australian region, the Wallacea whose fauna shows influences from both these regions (Dickerson et al., 1928). The Oriental region and Papuan region (or in a strict biographic sense the Papuan subgregion of the Australian region) are mostly considered to be separated by Weber’s line, which roughly marks the western boundary of the Sahul Shelf (see Briggs, 1987) and runs between the Philippines, Sulawesi and the Lesser Sunda Islands in the west and the Maluku Islands (also called Moluccas) in the east. All islands and land masses east of Weber’s line are attributed to the Papuan region (Gressitt, 1982). Mayr (1944) regarded Weber’s line as a line of faunal balance, east of which the Papuan faunistic elements have a numerical superiority over Oriental elements. Recent studies have revealed that also the Philippines, which are part of the Oriental region, need to be considered as part of Wallacea (e.g. Vallejo, 2011). The northern regions


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of continental Asia as well as Japan are included in the Paleartic region, which comprises all portions north of a line running along the Himalaya, through central China and between Japan and the Ryukyu Islands. On the basis of its geological history and physiogeography the Oriental region is split into several geographical subregions or units some of which are still controversely defined (see Inger, 1999). The subregions helpful for understanding the distributional pattern of Heteropterygidae can be summarized as follows (see Inger, 1999): a) the Himalaya and northeastern montane regions comprising Nepal, Bhutan, Sikkim, Darjeeling, Pradesh, Ladakh, Kashmir, Manipur, Nagaland and the mountainous regions of northern Myanmar, Thailand and Vietnam as well as soutern China; b) the Southeast Asian lowlands which comprise the central valleys of Myanmar, Thailand, Cambodia as well as southern and coastal Vietnam; c) Sundaland; d) Sulawesi; e) the Philippine Islands excluding the Palawan chain; f) the Lesser Sunda Islands from Lombok on the west to Timor on the east. Sundaland (Fig. 341) or former “Malaya” (see Mell, 1930; Johnson, 1964; Voris, 2000; Sathiamurthy & Voris, 2006), the main distributional area of the Heteropterygidae, is one of those subregions which are still controversially defined by zoologists, mainly due to Peninsular Malaysia showing strong Burmese elements among its fauna (see Eliot, 1992). Accordingly, most zoologists regard Peninsular Malaysia as part of Sundaland while others regard it as a separate subregion, which additionally comprises the peninsular regions of Myanmar and Thailand. Herein, Sundaland is defined to cover the zoogeographical unit of the land masses on the Sunda shelf, that is: Peninsular Malaysia south of the Isthmus of Krah (or south of Latitude 7–10°N depending on authors), the coastal areas of Cambodia, parts of the eastern and southern coasts of Thailand and Vietnam, including the islands Sumatra, Java, Borneo, Bali, Balabac, the Palawan chain, the Calamaines and Cuyos Islands, all the smaller intervening islands, as well as the islands off the west coast of Sumatra (Simeulue, Nias, the Batu Islands, Kepulauan, the Mentawei Islands and Engano) and the Christmas and Cocos-Keeling Islands (see Eliot, 1992; Nässig & Treadaway, 1998: 231; Sathiamurthy & Voris, 2006). Within the defined area the sea is quite shallow but depth increases rapidly outside. Sundaland is located on the Sunda Shelf, which forms a large submerged extension of the present day land masses. This is believed to have been exposed during the Last Glacial Maximum (LGM) in the late Pleistocene and early Holocene when sea leves are estimated to have been almost 120 m lower than modern mean sea levels (Haneburth et al., 2000; Sathiamurthy & Voris, 2006). The exposed shelf formed a large land mass, that connected the islands of Borneo, Java and Sumatra, and most certianly also the islands of Palawan and Balabac with continental Asia during the LGM and from time to time also in the more distant past. The land or island chain bridges between the Asian continent and the Large Sunda Islands, allowed biota to disperse throughout present Sundaland and the Southeast Asian lowlands including Thailand, Cambodia and Vietnam (see Briggs, 1987; Voris, 2000; Sathiamurthy & Voris, 2006). The present fragmentation was brought about by the conditions prevailing in the last Ice Age (= last glacial period) some 110.00–12.000 BP, since the world-wide post-glacial increase of the sea level large areas were submerged which caused an isolation of the islands. It is however believed that until a – 50 m contour sea level some 12.000 BP, the Greater Sunda Islands were more or less directly connected to the mainlands of Asia (Peninsular Malaysia, Cambodia, Thailand and Vietnam) by land or island chain bridges (Voris, 2000; Sathiamurthy & Voris, 2006). During the LGM the vegetation of the exposed portions of Sundaland is estimated to have consisted mainly of tropical rainforest or grassland, as well as monsoon and dry forests (Voris, 2000). Java was the first major island to be isolated from the rest of Sundaland and exhibits a less rich but stronger differentiated fauna from that of Peninsular Malaysia, Sumatra or Borneo, which together are also termed Neomalaya (see Moulton, 1915; Nässig & Treadaway, 1998). Confirmation for this hypothesis is represented by the distribution of the subfamily Heteropteryginae, whose taxa are distributed throughout entire Neomalaya but absent on Java. The present study has shown the few historical records of Heteropteryginae-members from Java to be erroneous. The small islands off the west coast of Sumatra (Simeuluë, Nias and Engano) are not on the Sunda Shelf and the Batu and Mentawei Islands are separated from Sumatra by deep channels and possess quite some degree of endemicity. Accordingly, they are often termed Paramalaya (e.g. Toxopeus, 1926) but in fact these islands should be included in Sundaland (Nässig & Treadaway, 1998). This is here supported by the distribution of the genus Haaniella, which has one species on Simeuluë Island but is otherwise restricted to Neomalaya. The eastern boundary of Sundaland is formed by Wallace’s line (named by Huxley in 1868), which runs along the Lombok Strait between Bali in the west and Lombok in the east as well as the Makassar Strait between Borneo in the west and Sulawesi in the east. Wallace’s line marks the eastern boundary of the Sunda Shelf and deep ocean straits, which separate shallow waters in Sundaland from deep waters througout Wallacea. Due to the lands east of Wallace’s line, namely the Lesser Sunda Islands from Lombok to Timor, Sulawesi and the Maluku Islands as well



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as the Philippines (except Palawan and the very eastern islands of the Sulu Archipelago) are not situated on the Sunda shelf these islands do not belong to Sundaland (see Johnson, 1964; Vane-Wright, 1990) but are included in the Wallacea. The Philippines is an archipelago of approximately 7000 islands in the Western Pacific ocean northeast of Sundaland and regarded as part of Wallacea (e.g. Dickerson et al., 1928; Vallejo, 2011). Physically, the Philippines can be divided into three groupings, this is a) Luzon, which is the largest island of the Philippines with an area of 104.688km² and positioned in the northern portion of the archipelago, b) Mindanao, which is the second largest (97.530 km²) and southeasternmost island, and c) the Visayas, which are the westernmost islands of the archipelago and are composed of the major islands of Panay, Negros, Masbate, Cebu, Samar and Leyte (Vallejo, 2011: 27). While the archipelago has predominantly biotic components from Sundaland there are also significant Wallacean, New Guinean and Australian elements in the biota. The southeasternmost major island Mindanao in particular shows evolutionary relationships with that of Sulawesi and has a significant Australian component in the eastern portion called the “Eastern Mindanao Corridor”, while the western section of the island has stronger biogeographical affinities to Borneo (Vallejo, 2011). The strong Gondwanan or Australian element in the eastern and southern Philippines, but Mindanao in particular, suggest a land bridge connection via volcano islands with Sulawesi and the rest of Wallacea at different times (Dickerson et al., 1928). This theory is for instance supported by the distribution of certain pigs and parrots with a Gondwanan origin, that occur on Mindanao but also throughout Wallacea, New Guinea and in Australia (Vallejo, 2011: 36). Hypothetical panbiogeographical tracklines via estimated island chain bridges, that allowed interchange between the faunas of Sundaland in the west and Sulawesi in the south with that of the Philippines and important for explaining the present day distributional patterns of the Heteropterygidae subfamilies Dataminae and Obriminae, ran from Borneo via the Sulu archipelago towards the Zamboanga Peninsula of western Mindanao and from central and southern Mindanao via the Talaud Islands towards northern Sulawesi (Heaney, 1986; Vallejo, 2011: 38, fig. 6). Further interchange between the biota of Borneo and the Philippines may have taken place via island chain bridges that included Palawan and in addition to earlier times were present during the LGM in the late Pleistocene and early Holocene when sea leves were almost 120 m lower than today (Heaney, 1986). Other hypothetical land connections appear to predate the Pleistocene and may date back as far as to the Pliocene some 5.33–2.58 Ma, when there were areas of shallow sea between Mindanao and Sulawesi, a corridor from northern and eastern Sulawesi including the Sula islands and Buru towards Seram and between Sulawesi and the southern Maluku Islands and the Lesser Sunda Islands (Hall, 1996; Vallejo, 2011: 30, fig. 3c). The following chapters discuss the distributional patterns of the three subfamilies of Heteropterygidae, this is Dataminae, Obriminae and Heteropteryginae, and provide additional biogeographical arguments for the phylogeny and systematic arrangement proposed herein. The family Heteropterygidae is distributed throughout the entire Oriental region with one species as far north as Taiwan and the Ryukyu Islands south of Japan and single representatives as far east as the southern Maluku Islands (Province Maluku) and as far south as some of the Lesser Sunda Islands (Fig. 342). The latter islands belong to Wallacea with the more eastern islands regarded as part of the Papuan region (see Gressitt, 1982). One species is found on Palau and represents the easternmost distribution of the family. No members of Heteropterygidae are found in the northern Maluku Islands (Province Maluku Utara), which predominantly have Papuan and Australian components in their biota. Borneo is the only island, which inherits numerous representatives of all three subfamilies of Heteropterygidae. Hence, Borneo must be regarded as the centre of the present day distribution of the family. The wide distribution of the Heteropterygidae indicates a high evolutionary age, which suggests a Gondwanan origin for this clade.

6.1. Distribution of Dataminae (Fig. 343) Based on morphological characters the Dataminae are here interpreted as the basalmost clade of the family Heteropterygidae (→ 3.1 and Fig. 97). This is supported by the very widespread distribution, which covers the entire distributional ranges of the other two subfamilies, namely Obriminae (→ 6.2) and Heteropteryginae (→ 6.3), except for the Philippines (Fig. 342). The largest number of Dataminae taxa is found in Borneo and throughout


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Sundaland, with the number of taxa gradually decreasing towards the more outer regions of the subfamilies present day distributional range. Five of the seven known genera are present in Borneo and two of these are Bornean endemics (Epidares Redtenbacher, 1906 and Spinodares Bragg, 1998). One genus (Dares Stål, 1875) is predominently Bornean with only one species found on Palawan (D. philippinensis Bragg, 1998). Orestes Redtenbacher, 1906 and Hainanphasma Ho, 2013 are the only genera, that are not found in Borneo and only two of the genera represented on Borneo have dispersed towards the north and northwest (Planispectrum Rehn & Rehn, 1939) or into all directions except Palwan (Pylaemenes Stål, 1875). Dataminae have settled various biotopes from swamp and coastal near lowland forests to highly mountainous habitats up to altitudes of 2000 metres. All members are exceptionally ground dwelling phasmatodeans and are found in low vegetation usually no more than 1 metre above the ground. Due to the lack of a beak-like ovipositor in ♀♀, the egg-laying strategy differs from that observed in Obriminae or Heteropteryginae and certainly represents the more basal stade, with eggs either being simply dropped to the ground or laid into low pits that are dug with the front legs. The Dataminae have dispersed more northward than than the either the Obriminae or Heteropteryginae and have settled great parts of the Southeast Asian lowlands including the very southern regions of China (Provinces Yunnan and Guangxi), the eastern and southeastern Chinese lowlands and coastal regions (Provinces Hainan, Guangdong, Jiangxi and Fujian) and even have one species (Pylaemenes shirakii Ho & Brock, 2013) as far north and northwest as Taiwan and the Ryukyu Islands south of Japan (Fig. 343). The latter dispersal might by explained by a glacial period in the late Pliocene and early Pleistocene or the LGM, when there supposedly were land connections between Taiwan and Japan and the Asian mainland (e.g. Hall, 1996). The presence of Dataminae on Java, the Lesser Sunda Islands, Sulawesi and the southern Maluku Islands but absence in the Philippines and northern Maluku Islands can only be explained by a dispersal via hypothetical tracklines which once covered island chain bridges that existed between Java and the Lesser Sunda Islands in the south towards Sulawesi and the southern Maluku Islands in the north perhaps as early as during the early Pliocene some 5 Ma, when there were areas of shallow sea in these regions (see Hall, 1996; Vallejo, 2011: 30, fig. 3c). There seem to have been no such land or island chain connections, that would support the described dispersal, during the LGM (see Sathiamurthy & Voris, 2006). One species, namely Pylaemenes coronatus (Haan, 1842), is in addition to Sulawesi and the southern Maluku Islands also found on the Talaud Islands north of Sulawesi. These islands must have been settled from northern Sulawesi via volcanic islands that formed island chain bridges, a theory which corresponds to a hypothetical trackline running between Sulawesi and Mindanao suggested by Vallejo (2001: 38, fig. 6). However, Dataminae are not present on Mindanao and no representatives have yet been recorded from this island or anywhere else in the Philippine archipelago. The lack of Dataminae in the Philippines may be explained by two different hypotheses. Firstly members of the subfamily have never reached Mindanao and the Philippines because dispersal into other directions was more successful. Secondly, perhaps Dataminae have become extinct on Mindanao and other islands of the Philippine archipelago, because their ecological niche was settled by the predominantly ground dwelling members of the Obriminae tribe Tisamenini n. trib., which has radiated strongly in the Philippines and has developed a good number of different species throughout almost the entire archipelago (→ 6.2). The historic record for P. coronatus (Haan, 1842) from New Guinea (see Kaup, 1871: 31; Redtenbacher, 1906: 48) is questionable and deserves further evaluation, since there are no recent records from New Guinea and this distribution is not supported by biogeographical arguments. The dispersal mechanisms and hypothetical tracklines for the distribution suggested for Dataminae above are indicated by arrows in Fig. 343.

6.2. Distribution of Obriminae (Fig. 344) The Obriminae are the most speciose and diverse subfamily of the Heteropterygidae and have predominantly radiated in the Philippine archipelago with only few representatives known from outside the Philippines. Hence, Obriminae must be regarded as a principally Wallacean clade. The mostly ground dwelling members of the tribe Tisamenini n. trib. settle this particular ecological niche peculiar to the subfamily Dataminae in all other regions (→ 6.1). Only four of the 15 known genera are not represented in the Philippines. Hoploclonia Stål, 1875 (tribe Tisamenini n. trib.) is endemic to Borneo and here postulated to be the most derived taxon of this tribe. Aretaon



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Rehn & Rehn, 1939 has one species in northeastern Borneo and an as yet undescribed species on Palawan and is the sister-taxon of the exceptionally Philippine Trachyaretaon Rehn & Rehn, 1939. The monotypical Heterocopus Redtenbacher, 1906 (tribe Obrimini) is endemic to the Palau Islands and represents the easternmost distribution of the subfamily. The only winged and here interpreted as the perhaps most derived genus of Obriminae, the monotypical Miroceramia Günther, 1934 (tribe Miroceramiini), is distributed throughout Wallacea and only found in northern Sulawesi and the island of Seram in the southern Maluku Islands. It represents the southernmost distribution of the subfamily. The Obriminae have settled a wide range of different habitats ranging from coastal or coastal near dry forests, humid lowland forests and montane forests up to altitudes of 1500 metres. Most taxa are found in ground near vegetation but single species (e.g. Mearnsiana bullosa Rehn & Rehn, 1939) are arboreal (pers. comm. with Joachim Bresseel) and the three genera of the tribe Tisamenini n. trib. are predominantly ground dwelling. All share the same derived egg-laying strategy, which is in common with the Heteropteryginae (→ 7.1). ♀♀ descend from their host plants during the night and use their beak-like ovipositor to bury the eggs in a suitable substrate. Ten of the 15 known genera are exceptionally Philippine taxa and only one predominatly Philippine genus (Theramenes Stål, 1875) has a single species on the Talaud Islands. Hence, the Philippine archipelago is the centre of radiation for Obriminae. All distribution outside the archieplago is estimated to have been reached by emigration from the Philippines and fairly late dispersal towards these regions must be estimated perhaps as late as the LGM in the late Pleistocene and early Holocene when sea leves are believed to have been almost 120 m lower than modern mean sea levels (Haneburth et al., 2000; Sathiamurthy & Voris, 2006). During that time land or island chain connections between the Philippines and Borneo in the west as well as the Talaud islands in the southeast are believed to have existed. The origin of the Obriminae however appears to be based on imigration to the Philippines possibly from Borneo via the Sulu islands towards Mindanao as early as the the early Pliocene some 5 Ma (see Hall, 1996; Vallejo, 2011: 30, fig. 3c). This hypothesis is based on the fact that the here suggested basalmost clade of Heteropterygidae, the subfamily Dataminae, is not present in the Philippines and might have become extinct subsequently, supposedly due to their ecological niche was settled by the more successful ground dwelling Tisamenini n. trib., which also have a fairly derived genus (Hoploclonia Stål, 1875) on Borneo (→ 6.1). Accordingly, Hoploclonia must be estimated to have a Philippine origin which is also undisputably the case for the two representatives of Aretaon Rehn & Rehn, 1939 (tribe Obrimini) on Palawan and in northeastern Borneo. While an emigration via the Sulu archipelago is more likely for Hoploclonia, the presence of an as yet undescribed species of Aretaon on Palawan rather suggests a dispersal to Borneo via this more northern trackline for the latter genus (Fig. 343). Due to the complex geography and geology of the Philippine archipelago, which consists of more than 7000 islands (Vallejo, 2011), the radiation of Obriminae within this large archipelago is apparently complicated and possible tracklines difficult to reconstruct. In general however a northward and northwestward dispersal can be estimated (Fig. 343), which started in western Mindanao and continued to Luzon and to the numerous smaller western islands referred to as the Visayas lateron (see Vallejo, 2011: 27). Any more comprehensive discussion on the dispersal of the Obriminae throughout the Philippine archipelago however warrants much detailed information on the tectonic history and geology of the Philippines and a more intense study of the Obriminae, which would go far beyond the aims of the present study. This shall be subject of future taxonomic or phylogenetic studies dealing with the numerous Philippine members of the subfamily and their relationships. Hypotheses for the Obriminae-taxa occuring outside the Philippine archipelago however, are discussed below. The genus Theramenes Stål, 1875 has three species in the southern Philippines islands of Mindanao and Panay but also one species on the Talaud islands situated between Mindanao in the north and Sulawesi in the south. The presence of Th. dromedarius (Westwood, 1859) on these islands may be explained by an accretion of terranes and volcano islands perhaps during the early Pliocene, which created a hypothetical panbiogeographic trackline formed by island chain bridges between Mindanao and northern Sulawesi (Vallejo, 2011: 38, fig. 6). The only winged member of Obriminae, the genus Miroceramia Günther, 1934 (tribe Miroceramiini), is found in northern Sulawesi and the island of Seram in the southern Maluku islands. It is not surprising that the supposedly closest relatives of this fairly derived genus are represented by members belonging in the here defined Theramenes-group of the tribe Obrimini found on Mindanao in the southern Philippines (e.g. Mearnsiana Rehn & Rehn, 1939) and the aforementioned Theramenes, which also has a species on the Talaud islands. The distribution of Miroceramia might be explained by a dispersal from Sulawesi via the Sula islands and Buru towards Seram perhaps as early as


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the Pliocene some 5.33–2.58 Ma when there were areas of shallow sea in this particular fairly narrow corridor between the mentuoned islands (see Vallejo, 2011: 30, fig. 3c). During this period, when the eastern Mindano terranes emerged, it is hypothesized there has also been a narrow area of shallow sea towards the Palau islands some 900 kilometres east of Mindanao (see Vallejo, 2011: 30, fig. 3c). This might explain the remarkable and disjunct distribution of the monotypical genus Heterocopus Redtenbacher, 1906, which is endemic to these islands but just as Miroceramia has its closest relatives represented by genera of the Theramenes-group of the tribe Obrimini on the Philippine island of Mindanao. The possible dispersal mechanisms and hypothetical tracklines explaining the present distribution of Obriminae above are indicated by arrows in Fig. 344.

6.3. Distribution of Heteropteryginae (Fig. 345) The Heteropteryginae are here interpreted as the most derived clade of the family Heteropterygidae, which is in addition to the morphological features discussed above (→ 3.4 and Fig. 97) supported by the much more restricted distribution compared to either Dataminae (→ 6.1.) or Obriminae (→ 6.2). The two genera contained, this is Haaniella Kirby, 1904 and Heteropteryx Gray, 1835, are peculiar to Sundaland with a single species represented in southern Vietnam (Haaniella gorochovi n. sp.). However, no members have so far been recorded from Palawan and the subfamily is apparently not present on Java, which geologically is the first island that became isolated from the rest of Sundaland during to the post-glacial increase of the sea levels after the last Ice Age some 12.000 BP. The monotypical Heteropteryx is exclusively Neomalayan and quite widely distributed, being most common throughout the Malayan Peninsula and southern Thailand, but is also found on Sumatra and with, so far, two records also known from northwestern Borneo. The records include various habitats and range from lowland and coastal forests (e.g. Phuket in southern Thailand and Pulau Penang) to mountainous regions up to 1500 m e.g. in the Cameron Highlands, Peninsular Malaysia. The insects predominantly live in low vegetation but sometimes may be found as high as 3 to 4 metres off the ground. The derived egg-laying strategy is shared with the Obriminae (→ 6.2), ♀♀ descending from the vegetation at night to bury the eggs with the help of their beak-like ovipositor (→ 7.1). The 17 known species of Haaniella are distributed throughout entire Sundaland except for Palawan, which includes both Neomalaya and Paramalaya. With eight known species Sumatra inherits the largest number of representatives of the genus, which are distributed throughout the entire island and have settled in all kinds of biotopes ranging from lowland to highly mountainous areas. While morphological observation shows seven of the Sumatran species to be closely related to each other and closely related to the two known congenerics in Peninsular Malaysia and Singapore (→ muelleri species-group), one species (H. parva Günther, 1944) is nested within a group of predominantly Bornean species (→ grayii species-group, see below). The only Paramalayan species of the genus (H. jacobsoni Günther, 1944) is endemic to Simaluë Island, the northernmost of the Batu Islands and some 150 kilometres off the Sumatra west coast, and most closely related to certain species found on Sumatra, hence belongs in the here defined muelleri species-group of Haaniella (→ 4.1.1). This supports the opinion of Nässig & Treadaway (1998) and various other authors, who suggest that, although not on the Sunda Shelf, the Batu and Mentawei Islands should biogeographically be regarded as part of Sundaland. The two species known from Peninsular Malaysia and Singapore, this is H. erringtoniae Redtenbacher, 1906 and H. macroptera n. sp., have their closest relatives in Sumatra and consequently belong to the muelleri species-group. Borneo has five species, all of which are more closely related to each other than to the Sumatran and Malayan species of the muelleri species-group, but morphologically divide into two groups of very closely related species. The three known species of the here defined echinata species-group are exceptionally Bornean and are morphologically the most distant representatives from the species here aggregated in the muelleri species-group (Fig. 355). While H. echinata (Redtenbacher, 1906) is widely distributed throughout almost the entire island and is also recorded from Labuan Island off the northern coast of Borneo (Fig. 495), H. saussurei Kirby, 1904 appears to be restricted to coastal or coastal near regions in the very northwestern portion of the island (Fig. 496). Both species are only found throughout lowland regions below altitudes of 800 metres. The third species of the echinata species-group, H. scabra (Redtenbacher, 1906), is peculiar to mountainous habitats above 800 metres in northeastern Borneo. The



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grayii species-group exhibits the widest distribution amongst the two other species-groups of Haaniella and is the only group that has species in and outside Borneo, hence is very interesting in biogeographical aspects. The two Bornean species H. grayii (Westwood, 1859) and H. dehaanii (Westwood, 1859) a very closely related and here interpreted as sister-taxa. While H. grayii is widely distributed throughout almost the entire island and fairly commonly encountered (Fig. 493), the rarer H. dehaanii is restricted to northwestern Borneo (Fig. 494). One remarkably small and specialized species (H. parva Günther, 1944) is restricted to northern Sumatra and is more closely related to H. dehaanii rather than H. grayii. The Vietnamese representative of the grayii species-group, H. gorochovi n. sp., is very interesting in a biogeographical aspect and combines morphological characters of both, the grayii species-group and the non-Bornean muelleri species-group. The here suggested close relation to the grayii species-group suggests a dispersal of Haaniella directly from Borneo towards Vietnam (Fig. 345) during the LGM in the late Pleistocene and early Holocene when sea levels are estimated to have been almost 120 m lower than modern mean sea levels. However, on the basis of morphology H. gorochovi n. sp. appears to be more close to the small Sumatran H. parva rather than the two Bornean species of that species-group. It is estimated that direct land connections existed from Borneo towards southern Thailand and Vietnam until as late as 10.880 BP when sea levels were no more than 40 metres lower than today (see Voris, 2000; Sathiamurthy & Voris, 2006). Accordingly, the disjunct distribution of H. gorochovi n. sp. supports close relation between the biota of Sundaland and the southern Indo-Chinese subregion, which has already been observed by Sterling et al. (2006) and various other authors. Another example for a similar disjunct distribution within Phasmatodea is represented by the genus Dajaca Brunner v. Wattenwyl, 1893 (Aschiphasmatidae: Aschiphasmatinae), which has several species in Borneo, one species in Vietnam (D. napolovi Brock, 2000) and one species in Myanmar (D. nigrolineata Hennemann et al., 1996) but is not present in either Sumatra or Peninsular Malaysia (see Bragg, 2001: 345). In general, since the supposedly most basal forms of Heteropteryginae are exceptionally Bornean and the clade is restricted to Sundaland, with the exception of a single species in southern Vietnam, a dispersal throughout present Sundaland must be estimated to have taken place rather late. Perhaps the main radiation had not occurred earlier than during the last period of low sea levels in the LGM in the late Pleistocene and early Holocene some 110.000–12.000 BP and was most intense on Sumatra with single taxa only having dispersed further northward and reached the Malay Peninsular or southern Vietnam. The dispersal mechanisms and hypothetical tracklines for the distribution here estimated for Heteropteryginae are indicated by arrows in Fig. 345.

7. Biology & Life history Members of the subfamily Heteropteryginae, e.g. Heteropteryx dilatata (Parkinson, 1798), represent some of the world’s most massive and heavy phasmids. Females of H. dilatata can reach maximum weights of 60 grams when in full egg production which is only exceeded by certain large Coleoptera, including some rhinoceros beetles of the subfamily Dynastinae and goliath beetles of the subfamily Cetoniinae. Also, some species are amongst the most spiny Phasmatodea known with e.g. the ♂♂ of Haaniella grayii (Westwood, 1859) or Haaniella jacobsoni Günther, 1944, bearing numerous long and very pointed spines on the head thorax and abdomen. The large size (14 cm +) of ♀♀ of some species stocky and massive body and remarkable spination makes the Heteropteryginae one of the most striking subfamilies of the Oriental fauna. With a maximum length of up to 12 mm and weights of up to 0.3 g the eggs of Haaniella echinata (Redtenbacher, 1906) are the largest and heaviest eggs of the entire Phasmatodea. Other facts, like the spectacular active defensive behaviour, which includes pinching with the spiny hind legs (Figs. 346–349) and producing loud stridulating noises from the wing bases, as well as interesting colour patterns make them some of the very fascinating representatives of the order. Hence, much attention has since been given to the species of this particular family with numerous papers published. Some of the most interesting facts are summarised below.

7.1. Behaviour In addition to their striking habitus, all species exhibit a very interesting and impressive active defensive behaviour. When disturbed or attacked, the insects will arch their abdomen forward with one or both hind legs widely splayed, which possess long, pointed or in-curving spines on the ventral carinae. If further disturbed, the insects will


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repeatedly pinch with their hind legs and some of the larger species are capable of inflicting painful wounds by this action. More rarely specimens will even try to bite and sometimes small amounts of a secretion are sprayed from defensive glands positioned in the anterior corners of the pronotum. At the same time a loud hissing or rattling stridulating sound is produced, by rubbing the tegmina and alae against each other and on the upper surface of the abdomen. The stridulating has been described numerous times in the literature already (e.g. Löser & Schulten, 1981; Dräger, 2011) and is produced by a stridulatory organ of the wings. This is formed by a number of fused wing-cells in the basal portion of the alae positioned between the radial and cubital vein (Fig. 84). Furthermore, the dorsal surface of the radial vein is set with numerous minute teeth which resembles a file and is rubbed against the ventral surface of the tegmina (Fig. 84). Males of Heteropteryx dilatata (Parkinson, 1798) are fully winged and lack the stridulatory organ found in all brachypterous species, hence are not able to produce a hissing sound. These insects however may open their large wings to display the bright red or pink colour of the alae and make them appear much larger than they actually are. Löser & Schulten (1981) reported H. dilatata to follow a large enough object that is moved round the insects when these are in their defensive position, which shows they must be fairly well able to see their direct environment. This active defensive reaction is supported by brightly and contrastingly coloured coxal trochanters of the metacoxae and intersegmental membranes between the basal abdominal sternites in most species of Haaniella (Figs. 389–396). These are often bright blue, green or red and readily exposed when the abdomen is arched forward and the hind legs widely splayed (346–349). Furthermore, the exposed interior surfaces of the metafemora of some species are bright red which is also meant to warn potential predators. ♀♀ of all Heteropteryginae have a beak-like ovipositor that is formed by an elongated epiproct and subgenital plate. This is used to bury the eggs by bending the abdomen downward and pushing the ovipositor into a substrate. Often as much as up to half of the abdomen may be pushed into the substrate, so that the eggs are released several centimetres below the surface. Usually eggs are laid individually but often several eggs are laid during a single night. Oviposition mostly takes place at night and particularly after rain falls and in Peninsular Malaysia H. dilatata (Parkinson, 1798) is particularly plentiful on the forest floor following rain falls (personal communication Michael K.P. Yeh). To lay their eggs ♀♀ will leave their daytime hiding places or the host-plants in the early evening and are often seen to meticulously search for a suitable place or substrate. The stocky and sturdy hatchlings are easily capable to grub themselves out and will immediately after emerging from the egg and substrate search for vegetation to climb on.

7.2. Life cycle Mating often takes place immediately after the ♀ has undergone the final ecdysis. During mating, the ♂ rests on the back of the ♀, the abdomen is bent downward with the anal segment covering the sternum VII of the ♀. The mostly triangular and movable vomer of the ♂ is then used as an anchorage on the praeopercular organ of the ♀, which is mostly represented by a raised central area near the posterior margin of sternum VII and set with two or three spines. Fertilization is brought about by transfer of spermatophores; these are spherical, white or pale cream and about 2–3 mm in diameter. Mating usually takes places at night and may last for several hours. Adult ♀♀ start laying eggs a few weeks after the first copulation and use their beak-like ovipositor for burying the eggs several centimetres deep in a substrate. Eggs take fairly long to hatch compared to most other phasmids, with hatching times ranging from 6–18 months depending on the species. Hatchlings emerge from the eggs by popping of the operculum of the eggs, grub out of the substrate and immediately search for vegetation to ascend. Nymphs undergo a minimum of five skin sheds and are quite slowly growing, in some species taking up to one year to reach maturity. Adults are very long-lived and usually live for at least one year. An adult ♀ of Haaniella scabra (Redtenbacher, 1906) that was collected by the two first authors on Mount Kinabalu in August 1996 survived in captivity in Europe until early 2001, which makes a total age of almost five years. Hence, Heteropteryginae are amongst the most long-living Phasmatodea, similar ages are only achieved by certain members of the related subfamily Dataminae.



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7.3. Parasitism Bragg (1993) reported all species of Haaniella to prone to parasitic mites (ectoparasites) to the extent that it is very rare to find a specimen without mites. In a survey of four species in 1991, the author found 63 of the 64 specimens examined to exhibit mites, the exception being a first instar nymph. The mites are reddish-brown and usually found in the abdominal pits but also on the antennae, legs and dorsal surface of the body. A ♀ of Haaniella grayii (Westwood, 1859) collected at Bengoh, Sarawak was found to be affected by more than 20 mites (Bragg, 2001: 27). Parasitism by endoparasites has also been reported by Bragg (1993: 40) for a ♀ of Haaniella echinata (Redtenbacher, 1906) which contained a 54 cm long Mermithid larva. At least in culture conditions eggs show a strong tendency to infection with fungus and regularly become mouldy.

7.4. Gynandromorphism (Figs. 350–354) The Jungle Nymph Heteropteryx dilatata (Parkinson, 1798) is well known to produce gynandromorph specimens, both in the field and in captivity. Several gynandromorphs of this species have already been mentioned and illustrated in the literature (e.g. Brock, 1989: 207) and due to the striking sexual dimorphism in H. dilatata these are particularly spectacular. The term gynandromorph is based on the Greek gyne (= female) and andro (= male) and describes the presence of female and male characteristics in a single specimen. Mostly the cause of this phenomenon is an event in miosis during the early development, when one of the dividing cells does not split its sex chromosomes typically, hence is a chromosomal disorder. Both known kinds of gynandromorphy i.e. bilateral gynandromorphs and mosaic gynandromorphs have been reported for H. dilatata. Bilateral gynandromorphs appear to be more common and are specimens that are male one one side and female on the other. Mosaic gynandromorphs have the male and female portions not bilaterally separated but less clearly defined and irregularly scattered throughout the entire morphology. The specimen shown in Figures 350–352 is a mosaic gynandromorph. The specimen is mostly male and at first glance looks like a usual ♂, having both wings fully developed and and average body length of 92.0 mm excluding the epiproct. Several portions of the head, thorax and abdomen however were bright green when alive, as typical for ♀♀. The legs typically male in morphology and colouration, but on the mesonotum there is a distinct anterolateral spine on the right and a similarly distinct pre-medial spine on the left, and abdominal tergum V bears a single fairly prominent median spine (Fig. 351). Apart from these features and the unusual green coloured portions of the head and body, the gynandromorphism of this specimen is most obviously seen in the genitalia. While generally male and having a fairly normal vomer, there is a large and well developed but somewhat misshaped epiproct, similar to that seen in ♀♀ (Figs. 351–352). Figures 353 and 354 show two further mosaic gynandromorphs, which both have more female characteristics than the previous specimen. The specimen in Figure 353 has male proportions and genitalia but many portions of the head, body, legs and even the tegmina show bright green colour typical for ♀♀. The specimen in Figure 354 is roughly half male and half female, having fully developed wings but a considerably broadened meso- and metathorax typical for ♀♀. The abdomen of this specimen is strongly deformed and the genitalia are similar to that of the insect shown in Figures 350–352 except for that the poculum is strongly reduced and only represented by a slender, lanceolate structure and the vomer is strongly reduced. The legs of this specimen also rather resemble that of ♀♀ than that of a typical ♂.

7.5. Local beliefs & medicinal For a long time species of this subfamily have been of importance or interest to native people in Southeast Asia, either as a source of food or medicinal properties. In Borneo the large members of Haaniella were, at least until recently, widely used as a source of food by the indigenous tribes and are still eaten in the region around Kapit, Sarawak (Bragg, 2001: 75). The indigenous people (Bidayuh Dayaks) at Bengo, Sarawak eat the eggs of Haaniella grayii (Westwood, 1859) after boiling in water for 30 seconds. Then the shell is removed and the content of the


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eggs eaten (Bragg, 1998b). Traditional Chinese families in Peninsular Malaysia rear Heteropteryx dilatata (Parkinson, 1798) for the healing powers of the droppings and keep them in cages on guava (Psidium guajava, Martaceae) in their gardens. The dried excreta are collected, mixed with herbs and are said to cure a number of ailments, such as asthma, stomach pains and muscular pains. Chan & Lee (1994) reported the excreta to be predominantly used as a medicine to treat diarrhoea and gastroenteritis by local people in Penang. The excreta have recently been found to be a concentrate of Vitamin E which is assumed to result from the guava the insects feed on (Brock, 1999: 145). Rich men in Kelantan, Peninsular Malaysia kept H. dilatata to secure the eggs, which they set in rings like jewels, and considered them a powerful charm against evil spirits (Annandale, 1900).

7.6. Habitats & host plants Most members of Heteropteryginae prefer warm and particularly humid habitats, ranging from lowland rainforests, over coastal forests to swamp forests. Haaniella scabra (Redtenbacher, 1906), H. parva Günther, 1944 and H. gintingi n. sp. are peculiar to mountainous habitats and only recorded from altitudes between 800 and 1700 metres, where average temperatures are comparatively lower than in all habitats of the other known species. Only Heteropteryx dilatata (Parkinson, 1798) is found in both highland and lowland forests being found as well up to 1500 metres in the Cameron Highlands, Peninsular Malaysia. In most cases, specimens of lowland Heteropteryginae are found within 100 metres of running water or swamps. In contrast, highland species do not seem to need water on the forest floor as part of their habitat as all montane forests are mist or cloud covered especially during the night and provide sufficient humidity and water on the foliage of the hostplants that the insects can drink (females generally need large amounts of water when in full egg-production, → 7.7). Haaniella echinata (Redtenbacher, 1906) for example is very common at Batu Niah (NE-Sarawak), an extremely hot and damp lowland dipterocarp forest, where temperatures usually range between 26–36°C and the relative humidity is generally above 85% the whole year round. During the rainy season, some areas are completely flooded, hence walkways have been constructed to allow tourists to reach the well known caves in the National Park area. Specimens can be found on the forest floor, low vegetation or even up to six metres in small trees and feed on a variety of native jungle plants. ♀♀ seem to stay in the vicinity of favourite foodplants for a considerable amount of their life-time and are generally less mobile and active than ♂♂. Abercrombie (1993: 14) reported that a ♀ of Haaniella grayii (Westwood, 1859), which was found feeding on Rubus sp. (Rosaceae) on Mount Serapi (NWSarawak), was still there when returning to this collecting locality four weeks later. While ♂♂ actively search for ♀♀ to mate with, ♀♀ seem to restrict their mobility mostly to descend from their foodplants to bury their eggs in the forest floor and other suitable substrates. Haan (1842: 108) reported that Haaniella muelleri (Haan, 1842) live under dead and dehydrated leaves in the jungle. All members of Heteropteryginae appear to be polyphagous and are not at all specialized feeders (e.g. Blüthgen et al., 2006; Eilmus, 2009). Below are list of the known natural foodplants of some species. Haaniella erringtoniae (Redtenbacher, 1906): Rubus moluccanus (Rosaceae), Psidium guajava (Myrtaceae), Mangifera indica (Anarcadiaceae) (Seow-Choen, 2000b: 40). Haaniella echinata (Redtenbacher, 1906): Saurauia ferox (Actinidiaceae), Fissiostigma sp. (Annonaceae), unidentified liana (Combretaceae), Costus sp. (Costaceae), Dryobalanops lanceolata (Dipterocarpaceae), Diospyros sp. (Ebenaceae), Baccaurea stipulata (Euphorbiaceae), Glochidion sp. (Euphorbiaceae), Mallotus wrayi and Mallotus sp. (Euphorbiaceae), Bauhinia sp. (Fabaceae), Spatholobus spp. (Fabaceae), Fordia sp. (Fabaceae), Ryparosa hullettii (Flacourtiaceae), Leea sp. cf. indica (Leeaceae), Algaia sp. (Meliaceae), Chisocheton sp. (Meliaceae), Knema sp. (Myristicaceae), Urophyllum sp. (Rubiaceae), Selaginella sp. (Selaginellaceae), Pterospermum sp. (Sterculiaceae), Brownlowia peltata (Tiliaceae), Tetrastigma sp. (Vitaceae), Globba pendula (Zingiberaceae) (Blüthgen et al, 2006). Haaniella grayii (Westwood, 1859): Nepenthes ampullaria (Nepenthaceae), ferns of Dicranopteris linearis (Gleicheniaceae), unidentified Zingiberaceae (Bragg, 2001: 13), Dillenia suffruticosa (Dilleniaceae). Rubus sp. (Rosaceae) (Abercrombie, 1993: 14). Haaniella macroptera n. sp.: Rubus moluccanus (Rosaceae) and Dillenia suffruticosa (Dilleniaceae). Heteropteryx dilatata (Parkinson, 1798): Rubus moluccanus (Rosaceae), Grewia acuminata (Tiliaceae), Eugenia aquea (Myrtaceae), Uncaria sp. (Rubiaceae) (Brock, 1999: 144). Durio zibethinus (Bombacaceae),



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Manihot esculenta (Euphorbiaceae), Psidium guajava (Myrtaceae) (Seow-Choen, 2000b: 40). Mangifera indica (Anarcadiaceae) (Seow-Choen, 1997: 128).

7.7. Rearing in captivity Most Heteropteryinae can be easily reared in captivity and feed on a variety of alternative foodplants, hence have become favourite pets by breeders and phasmid enthusiast worldwide. The larger species such as the well-known “Green Jungle Nymph” Heteropteryx dilatata (Parkinson, 1798) in particular are often the main attractions at entomological exhibitions or in zoological gardens. Numerous papers have since dealt with the biology, life cycle and breeding of certain species (e.g. Cocking, 1985 & 1986; D’Hulster, 1990; Abercrombie, 1993; Schulten, 1995; Dräger, 2011). Species that have so far been successfully cultured in Europe are Haaniella dehaanii (Westwood, 1859), H. echinata (Redtenbacher, 1906), H. erringtoniae erringtoniae (Redtenbacher, 1906), H. gintingi n. sp., H. gorochovi n. sp., H. grayii (Westwood, 1859), H. saussurei Kirby, 1904, H. scabra (Redtenbacher, 1906) and Heteropteryx dilatata (Parkinson, 1798) and it is hoped in future that further species will be introduced to culture. Alternative foodplants used in captivity in Europe include numerous Rosaceae like bramble (Rubus fruticosus), raspberry (Rubus idaeus), rose (Rosa spp.), pyracantha (Pyracantha coccinea), hawthorn (Crataegus monogyna), cherry (Prunus spp.), apple (Malus spp.) and cherry laurel (Prunus laureocerasus) but also plants from various other families, including oak (Quercus spp., Fagaceae), guava (Psidium guayava, Myrtaceae), salal (Gaultheria shallon, Ericaceae) and ivy (Hedera helix, Araliaceae). Most species are very polyphagous (→ 7.6) many of these and also other plants are accepted by most members of the subfamily that have been reared so far. In general, there are quite standard conditions for breeding Heteropteryginae successfully in culture, since most species originate from similar natural habitats (→ 7.4) and consequently deserve similar climatic conditions. Medium-sized cages or glass terrariums are suitable for most species, although larger cages with a minimum height of 60 cm are necessary for the large Heteropteryx dilatata (Parkinson, 1798). All Heteropteryginae prefer high humidity and only little ventilation is needed. All species are very fund of water, which should be sprayed into the cage at least once a day. The massive ♀♀ in particular need large amounts of water for producing the huge eggs, and will drink every day if given the possibility. While a spray of water is plenty for nymphs which generally seem to get enough liquid from the foodplants, a flat plastic plant pot saucer filled with water that is placed on the cagefloor appears essential for the wellbeing of the insects. The insects will drink every day, often leaving their daytime hiding places or descend from the foodplants to submerge their whole head and soak up the water (Abercrombie, 1993: 15). Females should be given a plastic container filled with vermiculite, peat or sand for oviposition or the cage-floor should be covered with a several centimetre thick layer of substrate. Due to the large size of the eggs ♀♀ are not very productive and at best produce one egg per day. Species of Haaniella in particular appreciate a hollow log or large pieces of cork bark to hide during the daytime unless the vegetation in the cage or terrarium is very thick and dense. Nymphs will also stay in the vegetation during the day if they can crawl into withered brown leaves (Abercrombie, 1993: 14). Nymphs and adults of Heteropteryx dilatata in contrast mostly rest in the vegetation during the daytime and nymphs often form aggregations. Since eggs show a tendency to easily go mouldy the best hatching ratios are achieved by removing the eggs from the cage and incubating them in damp vermiculite or sand with only the operculum showing. This helps to prevent other eggs being infected by fungus. As nymphs and adults of Heteropteryginae are stocky and very robust, they can easily be kept together in the same cage, but to avoid confusion it is preferable to keep different species separated.

8. Summary The areolate Oriental family Heteropterygidae has been re-arranged and a detailed revision of the subfamily Heteropteryginae at the species-level has been provided in the presented paper. Former treatments of the family and the intra-familiar arrangement of Heteropterygidae proposed by Zompro (2004) have proven unsatisfactory and have also received little support by phylogenetic and molecular results in Whiting et al. (2003) and Bradler (2009). The arrangement and relationships established herein widely corresponds to the aforementioned phylogenetic and molecular studies and is also supported by biogeography, which was not discussed by Zompro (2004). While the subdivision into the three subfamilies Dataminae, Obriminae and Heteropteryginae suggested by Zompro (2004:


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194) is supported, the relationships between these subfamilies and their subordinate taxa however are here interpreted differently and have resulted in a decidedly more satisfactory re-arrangement (Fig. 97b). Several of the characters stated to be plesiomorphic by Zompro (2004) are here shown to be in fact apomorphic character states (→ 3. & 3.4). Examples include the secondary development of a beak-like ovipositor and correlating derived strategy of egg depostion in Obriminae and Heteropteryginae (see Bradler, 2009; Goldberg et al., 2015), the secondary development of wings and even further specialization of these organs in Heteropteryginae and two genera of Obriminae (Miroceramia Günther, 1934 and Pterobrimus Redtenbacher, 1906) as well as the secondary development of a medio-apical spine on the area apicalis of the tibiae in Heteropteryginae and Obriminae. The apomorphy of these characters consequently places the predominantly ground-dwelling Dataminae as the basalmost subordinate clade of Heteropterygidae and not as the most derived one as postulated by Zompro (2004: 191, Fig. 119). This is supported by biogeographical facts, Dataminae having a much wider distributional pattern than either Obriminae or Heteropteryginae. It extends from Japan and Taiwan over entire Sundaland to as far southeast as Wallacea (→ 6.1) and hence almost completely incorporates the present-day geographical distributions of the other two subfamilies (Fig. 342). Obriminae are restricted to the Philippines, Borneo, certain islands of Wallacea and the Palau Islands (→ 6.2; Fig. 344) and Heteropteryginae are entirely restricted to Sundaland (with the exception of Java) and have a single species in southern Vietnam (→ 6.3; Fig. 345). The ground-pattern of Heteropterygidae includes small, stocky and robust, wingless, ground-dwelling phasmids with moderate sexual dimorphism, a short mesothorax, which is hardly longer than the prothorax, short and robust antennae, which are shorter than the fore legs, an area apicalis on all six tibiae, basally compressed profemora, a well developed vomer in ♂♂ and no ovipositor in ♀♀, most certainly with a Gondwanan origin. The basalmost taxon of Dataminae is Planispectrum Rehn & Rehn, 1939 (→ 3.1), which has very short and robust, perlamorph antennae with no more than 21 but sometimes only 13 antennomeres, that are distinctly shorter than the fore legs. This leads to the hypothesis that the common ancestor might have been a phasmid with even more considerably shortened antennae. The presence of rough sensory-areas on the prosternum and profurcasternum might possibly be an autapomorphy of Heteropterygidae, but wether these have already been present in the common ancestor or have been evolved de novo within Heteropterygidae is still to be clarified. The secondary elongation of the antennae, secondary development of wings and secondary development of a beak-like ovipositor in ♀♀ with the correlating derived strategy of egg deposition are undoubtedly apomorphic character states within Heteropterygidae, as are the reduction of the paired sensory-areas of the prosternum in Heteropteryginae and reduction of the central sensory-area on the profurcasternum in Obriminae. The arrangement of the Obriminae suggested by Zompro (2004) as well has not reveived much support by the present study and is shown to be arbitrary and unsatisfactory in several aspects. Taking the aforementioned groundpattern of Heteropterygidae and biogeographical facts into account, this study questions the three tribes introduced by Zompro (2004), leading to a fairly different intra-subfamiliar arrangement and revealing a principally Wallacean origin of the clade. Three tribes are now recognized within Obriminae, the Obrimini, the Miroceramiini and the Tisamenini n. trib. (→ 3.2). Autapomorphies of Obriminae are the secondary reduction of the central sensory-area on the profurcasternum and increasingly elongated basitarsus, which is decidedly longer than the second tarsomere (Fig. 97b). Tisamenini n. trib. is the sister-group of Obrimini + Miroceramiini and shown to contain the basalmost taxa, which resemble members of the Dataminae in several aspects, i.e. on average small size, stocky body, moderate sexual dimorphism, mostly ground-dwelling habits, short and robust antennae, which are hardly equal in length to the fore legs, indistinct medio-apical spine of the area apicalis of the tibiae and hairy structures of the eggs (→ 3.2.2). Three of the four genera contained are geographically restricted to the Philippine Islands, and Hoploclonia Stål, 1875 is endemic in Borneo. A secondary beak-like ovipositor in ♀♀ is present but it is less developed with the epiproct and subgenital plate comparatively shorter than in either Obrimini or Miroceramiini. In Hoploclonia, the only Bornean representative, the ovipositor is formed by the subgenital plate and and elongated anal segment, which is an autapomorphy of this genus and unique amongst the entire Heteropterygidae. Furthermore, Hoploclonia is the only genus of the entire Heteropteryginae which has a gula. The eggs of all four genera of Tisamenini n. trib. bear hairy structures on the capsule and operculum, which are secondarily reduced in Obrimini and Miroceramiini, and are considerably less elongate than those of either tribe. The composite or dichotomic spines of the pronotum and raised triangular area on the mesonotum are autapomorphies of Tisamenini n. trib. (Fig. 97). The composite posterior spines of the meso- and metanotum and the secondarily reduced hairy structures of the eggs are synapomorphies of Obrimini + Miroceramiini (Fig. 97b), as are the increasingly



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elongated antennae and antennomeres. The latter tribe, which is restricted to Wallacea, is characterized by the secondary development of wings which is an autapomorphy but also a homoplasy shared with Heteropteryginae. Autapomorphies of Obrimini are still difficult to define and deserve further research. One undoubtedly apomorphic character however is represented by the metasternal foramina of some genera, which are pits, holes or invaginations near the lateral margin of the metasternum (→ 3.2.1, Figs. 38–40). Three well marked generic groups are herein recognized within Obrimini, the Obrimus-group, the Stenobrimus-group and the Theramenes-group. The Heteropteryginae is the sister-group of the Obriminae, with which it shares synapomorphies such as the medio-apical spine on the area apicalis of the tibiae (Figs. 93–96) and the beak-like ovipositor of ♀♀ (Fig. 97) as well as the elongated antennae and antennomeres. Autapomorphies of Heteropteryginae are the secondary reduction of the paired sensory-areas on the prosternum, straight profemora, secondary development of wings in both sexes which have a tympanal region (= stridulatory organ) in the basal portion of the alae of brachypterous taxa as well as the X-shaped micropylar plate, which has two anterolateral extensions (Fig. 97b) and secondary lack of a median line (Figs. 428–430, 445–447). It is the most derived clade of Heteropterygidae (→ 3.4) and one of the two genera contained, i.e. Heteropteryx Gray, 1835, has evolved several characters that support increasingly aboreal habits. These include the bright green or yellow colour and strongly dilated abdominal tergites of ♀♀, which give the insects a leaf-like appearance and support camouflage amongst the foliage of its hostplants, as well as the fully developed wings of ♂♂, which enables the insects of active short-distance flights (→ 5). A detailed revision of Heteropteryginae at the species-level is provided in the present study and five new species are described (→ 4). The second genus of Heteropteryginae, namely Haaniella Kirby, 1904, is divided into three well marked species-groups, the muelleri species-group (→ 4.1.1), the grayii species-group (→ 4.1.2) and the echinata speciesgroup (→ 4.1.3). The subfamily has the most restricted geographical distribution amongst the clades of Heteropterygidae and is peculiar to Sundaland, being only found on Borneo, Sumatra, the Mentawei Islands, Singapore, Peninsular Malaysia and Thailand but with a single species in southern Vietnam (Fig. 345). More detailed information on the inter- and intra-generic relationships of Heteropteryginae are presented above (→ 5) and shown in Figure 355. While the relationships between the three subfamilies of Heteropterygidae could mostly be revealed and explained in more detail (→ 3.4; Fig. 97b) and an attempt of an intra-generic systematization of the Heteropteryginae has been presented herein (→ 5; Fig. 355), the inter-generic relationships of the Obriminae in particular deserve more extensive research based on phylogenetic and molecular studies and the attempt of an intergeneric systematization within Dataminae presented by Bragg (1998: 113, fig 113) and Bragg (2001: 128, fig. 34) warrants further evaluation. It is much hoped that future research will be executed to complete our knowledge on the phylogeny of this fascinating group of Oriental Phasmatodea, which contains some of the most striking extant insects.

8.1. List of taxonomic changes 1. Eubulidini Zompro, 2004: 205 (Type-genus: Eubulides Stål, 1877: 68), synonymised with the tribe Obrimini Brunner v. Wattenwyl, 1893 n. syn. [→ 3.2.1] 2. Haaniella erringtoniae novae guineae Günther, 1930: 737, here removed from synonymy with Haaniella muelleri (de Haan, 1842) and synonymised with Haaniella rosenbergii (Kaup, 1871) n. syn. [Valid name = Haaniella rosenbergii (Kaup, 1871), → 4.1] 3. Haaniella mülleri simplex Günther, 1944: 72, here removed from synonymy with Haaniella muelleri (de Haan, 1842) and synonymised with Haaniella glaber (Redtenbacher, 1906) n. syn. [Valid name = Haaniella glaber (Redtenbacher, 1906), → 4.1] 4. Haaniella parva Günther, 1944: 73, here removed from Miniopteryx Zompro, 2004: 198 and retransferred to Haaniella Kirby, 1904 rev. stat. [Valid name = Haaniella parva Günther, 1944, → 4.1.] 5. Hennobrimus Conle, 2006: 44. (Type-species: Hennobrimus hennemanni Conle, 2006: 45), here synonymised with Mearnsiana Rehn & Rehn, 1939 n. syn. [→ 3.2.1] 6. Hennobrimus hennemanni Conle, 2006: 45, here synonymised with Mearnsiana bullosa Rehn & Rehn, 1939 n. syn. [Valid name = Mearnsiana bullosa Rehn & Rehn, 1939, → 3.2.1] 7. Heteropteryx erringtoniae Redtenbacher, 1906: 169, here removed from synonymy with Haaniella muelleri (de Haan, 1842) and re-established as a valid species (rev. stat.) [Valid name = Haaniella erringtoniae erringtoniae (Redtenbacher, 1906), → 4.1]


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8. Heteropteryx Rosenbergii Kaup, 1871a: 35, here removed from synonymy with Haaniella muelleri (de Haan, 1842) and reestablished as a valid species (rev. stat.) [Valid name = Haaniella rosenbergii (Kaup, 1871), → 4.1] 9. Hoploclonia Stål, 1875b: 8 (Type-species: Acanthoderus gecko Westwood, 1859: 52), here transferred to the tribe Tisamenini n. trib. [→ 3.2.2] 10. Ilocano Rehn & Rehn, 1939: 460 (Type-species: Ilocano hebardi Rehn & Rehn, 1939: 461), here transferred to the tribe Tisamenini n. trib. [→ 3.2.2] 11. Leocrates glaber Redtenbacher, 1906: 167, here removed from synonymy with Haaniella muelleri (de Haan, 1842) and reestablished as a valid species (rev. stat.) [Valid name = Haaniella glaber (Redtenbacher, 1906), → 4.1] 12. Leocrates mecheli Redtenbacher, 1906: 167, here removed from synonymy with Haaniella muelleri (de Haan, 1842) and re-established as a valid species (rev. stat.) [Valid name = Haaniella mecheli (Redtenbacher,1906), → 4.1] 13. Mearnsiana Rehn & Rehn, 1939: 458 (Type-species: Mearnsiana bullosa Rehn & Rehn, 1939: 459), here removed from Miroceramiini Zompro, 2004 and transferred to the tribe Obrimini Brunner v. Wattenwyl, 1893 [→ 3.2.1] 14. Miniopteryx Zompro, 2004: 198 (Type-species: Haaniella parva Günther, 1944: 73), here synonymised with Haaniella Kirby, 1904 n. syn. [→ 4.1.] 15. Phasma (Heteropteryx) mülleri var. b de Haan, 1842: 108, here synonymised with Haaniella rosenbergii (Kaup, 1871) n. syn. [Valid name = Haaniella rosenbergii (Kaup, 1871), → 4.1] 16. Pterobrimus Redtenbacher, 1906: 43 (Type-species: Pterobrimus depressus Redtenbacher, 1906: 43), here transferred to the tribe Tisamenini n. trib. [→ 3.2.2] 17. Theramenes dromedarius Stål, 1877: 68, here removed from synonymy with Theramenes olivaceus (Westwood, 1859) and re-established as a valid species (rev. stat.) [Valid name = Theramenes dromedarius Stål, 1877, → 3.2.1] 18. Tisamenus Stål, 1875a: 50, 92 (Type-species: Tisamenus serratorius Stål, 1875a: 92), here transferred to the tribe Tisamenini n. trib. [→ 3.2.2] 19. Trachyaretaon manobo Lit & Eusebio, 2005: 80, here synonymised with Mearnsiana bullosa Rehn & Rehn, 1939 n. syn. [Valid name = Mearnsiana bullosa Rehn & Rehn, 1939, → 3.2.1]

9. Acknowledgements The authors would like to express their thanks to the curators and staff of the following museums and institutions for access to their collections, the loan of specimens for examination or providing photographs and information on specimens in the corresponding collections: Dr. Andrej Gorochov (ZIN, St. Peterburg); Dr. Cristiane Amedegnato, Dr. Toni Robillard, Simon Paulain and Emmanuel Delfosse (MNHN, Paris); Dr. Ulrike Aspöck and Dr. Susanne Randolf (NHMW, Vienna); Dr. Martin Brancucci, Dr. Daniel Burckhardt and Dr. Matthias Borer (NHMB, Basel); Dr. Jacques Cools, Jérôme Constant and Joachim Bresseel (RBINS, Bruxelles); Dr. George McGavin (OXUM, Oxford); Prof. Volker Mahnert und Dr. Peter Schwendinger (MHNG, Geneve); Dr. George Beccaloni and Mrs Judith Marshall (NHMUK, London); Dr. M. Müller (ETHZ, Zurich); Dr. Michael Ohl and Mrs Ulrike Dorandt (MNHU, Berlin); Dr. Roberto Poggi (MCSN, Genova); Dr. Wolfgang Schawaller (SMNS, Stuttgart); Dr. Wolfgang Schneider (HLMD, Darmstadt); Prof. Hans Strümpel, Dr. Kai Schütte and Mrs Ilona Rehmann (ZMUH, Hamburg); Dr. Roland Emmerich (SMTD, Dresden); Dr. Jan van Tol and Dr. Luc Willemse (RMNH, Leiden); Dr. Jason D. Weintraub and Dr. John Gelhaus (ANSP); Prof Klaus Schönitzer (ZSMC, Munich); Dr. Dimitri Luganov (MMUM, Manchester). The following colleagues are thanked for the loan of specimens from their collections, providing information, culture material or photos and helpful correspondence: Bruno Kneubühler (Zürich, Switzerland), Ian Abercrombie (Kent, England), Kristien Rabaey & Rob Simoens (Veurne, Belgium), Rob Krijns (Maastricht, Netherlands), Holger Dräger (Schwerin, Germany), Dr. Philip E. Bragg (Nottinghamshire, England), Mel Herbert (Hütschenhausen, Germany), Peter Heusi (Uster, Switzerland), Kim D’Hulster (Belgium), Wim Potvin (St-Pieters Leeuw, Belgium), Dr. John T.C. Sellick (Kettering, England) and Michael K.P. Yeh (Ipoh, Malaysia). Paul Bertner (UK) generously forwarded pictures of Haaniella-species he took on several occasions throughout Borneo and Jeremy Holden (Vietnam) kindly forwarded pictures of live males of Haaniella kerincia n. sp. taken on Gunung Tujuh, Kerinci-Seblat NP, Sumatra. Further thanks are due to Mel Herbert (Hütschenhausen) for his help and guidance during night collecting and accomodation for the first author in Brunei.



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The Insect Diversity Association at the Academy of natural Sciences, Philadelphia. Parkinson, J. (1798) Description of the Phasma dilatatum. Transactions of the Linnean Society, London, 4 (1), 190–192, pl. 18. http://dx.doi.org/10.1111/j.1096-3642.1798.tb00527.x Redtenbacher, J. (1906) Die Insektenfamilie der Phasmiden. I. Phasmidae Areolatae. Leipzig. pp. 1–180, pls. 1–6. Rehn, J.A.G. (1904) Studies in the Orthopterous family Phasmidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 56, 38–107. Rehn, J.W.H. (1938) Notes on the genus Haaniella with the description of a new species (Orthoptera: Phasmatidae). Transactions of the American Entomological Society, 63, 85–87, pl. 7. Sathiamurthy, E. & Voris, H.K. (2006) Maps of Holocene sea level transgression and submerged lakes on the Sunda Shelf. The Natural History Journal of Chulalongkorn University, Supplement 2, 1–43. Saussure, H. (1869) Mélanges Orthoptèrologiques. III. Phasmides.- Mémoires de la Socieété de Physique etd´Histoire naturelle de Genève, 20 (1), 227–326, pls. 2–3. Schulten, D. (1995) Wandelnde Blätter, Stab- und Gespenstschrecken. Entomologische Mitteilungen aus dem Löbbecke Museum &Aquazoo, 3, 1–132. Sellick, J.T.C. (1997) Descriptive terminology of the phasmid egg capsule, with an extended key to the genera based on eggstructure. Systematic Entomology, 22, 97–122. http://dx.doi.org/10.1046/j.1365-3113.1997.d01-30.x Sellick, J.T.C. (1998) The micropylar plate of the eggs of Phasmida, with a survey of the range of plate form within the order. Systematic Entomology, 23, 203–228. http://dx.doi.org/10.1046/j.1365-3113.1998.00056.x Seow-Choen, F. (1995) Two more gynandromorphs of the Malayan jungle nymph, Heteropteryx dilatata (Phasmida) with notes on captive behaviour. Bulletin of the Amateur entomologist´s Society, 54 (399), 49–51, colour fig. 1. Seow-Choen, F. (1996) Eggs found in a gynandromorph of the Malayan jungle nymph, Heteropteryxdilatata (Phasmida). Bulletin of the Amateur entomologist´s Society, 55 (406), 122. Seow-Choen, F. & Brock, P.D. (1994) An introduction to the Stick and Leaf-insects of Singapore. Malayan Naturalist, 48, 7– 11. Seow-Choen, F. (1998) List of stick-insects (Phasmida) in the collection of the centre for insect systematics Universiti Kebangsaan Malaysia. Serengga, 3 (1), 87–92.


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Seow-Choen, F. (2000a) Illustrated guide and key to the Haaniella (Phasmida, Bacillidae: Heteropteryginae) species of Malaysia. Serangga, 5, 149–164. Seow-Choen, F. (2000b) An illustrated guide to the Stick and Leaf Insects of Peninsular Malaysia and Singapore. Natural History Publications (Borneo), Kota Kinabalu, 173 pp. Seow-Choen, F. (2016) A taxonomic guide to the stick insects of Borneo including new genera and species. Natural History Publications (Borneo) Sdn. Bhd., Kota Kinabalu, 454 pp. [ISBN 978-983-812-169-9] Sorpongpaisal, W. & Thanasinchayakul, S. (2006) Identification of some Stick and Leaf Insects, (Order Phasmida) in Thailand. Kamphaengsean Academy Journal, 4 (3), 10–32. Shaw, G. (1806) Generale Zoology or systematic natural history. Insects. Volume 6, part 1. London. Shiraki, T. (1935) Orthoptera of the Japanese Empire (Part IV) Phasmidae. Memoirs of the Faculty of Science and Agriculture, Taihoku lmperial University, Formosa. 14 (3), 23–88, pl. 4–10. Stål, C. (1875a) Recensio Orthopterorum III. Revue critique des Orthoptères déscrits par Linné, de Geer et Thunberg. P.A. Norstedt & Söner, Stockholm. Stål, C. (1875b) Recherches sur le Système des Phasmides. Bihang till Kongliga Svenska Vetenskaps-Akademiens Handlingar. 2 (17), 1–19. Stål, C. (1877) Orthoptera nova ex Insulis Philippinis descripsit. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar. 34 (10), 33–58. Sterling, E.J, Hurley M.M. & Minh, L.D. (2010) Vietnam: a natural history. Yale University Press, New Haven, 423 pp. Tilgner, E.H. (2001) Fossil Record of Phasmida. Insect Systematics and Evolution, 31, 473–480. http://dx.doi.org/10.1163/187631200X00507 Tilgner, E.H. (2002) Systematics of Phasmida. Dissertation. University of Georgia, USA. Toxopeus, L.J. (1926) [not title, in the proceedings of the Society of the Nederlansche Entomologische Vereeniging]. Tijdschrift vor Entomologie, 69, Versla: Ixx-IxxxI. Uvarov, B.P. (1940) Twenty four new generic names in Orthoptera. Annals and Magazine of Natural History, London. 11 (6), 112–117. http://dx.doi.org/10.1080/03745481.1940.9723661 Vallejo, B. (2011) The Philippines in Wallacea. In: Telnov, D.M. (Ed.), Biodiversity, Biogeography and Nature Conservation In Wallacea and New Guinea, Vol. 1. Latvian Society of Entomology, Riga, Latvia. pp. 27–42. Van-Wright, R.I. (1990) Chapter 2: The Philippines—Key to the biogeography of Wallacea? In: Knight, W.J. & Holloway, J.D. (Eds.), Insects and the rain forest of South East Asia (Wallacea). Royal Entomological Society, London. pp 19–34. Voris, H.K. (2000) Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems and time durations. Journal of Biogeography, 27, 1153–1167. http://dx.doi.org/10.1046/j.1365-2699.2000.00489.x Westwood, J.O. (1848) Cabinet of Oriental Entomology. London. Westwood, J.O. (1859) Catalogue of Orthopterous insects in the collection of the British Museum. Part 1: Phasmidae.British Museum, London, 196 pp., pls. 1–40. Westwood, J.O. (1864) Heteropteryx hopei, Westw. Transactions of the Entomological Society of London, 2 (3), 16–17. Westwood, J.O. (1873–1874) Thesaurus Entomologicus Oxoniensis; or, illustrations of new, rare, and interesting Insects, for the most part contained in the collections presented to the University of Oxford by the Rev. F.W. Hope. Oxford [produced in four parts: Part 1, pp.1–56 in 1873; parts II–IV in 1874] Whiting. M.F., Bradler, S. & Maxwell, T. (2003) Loss and recovery of wings in stick insects. Nature, 421 (6920), 264–267. http://dx.doi.org/10.1038/nature01313 Zompro, O. (1996) Bemerkungen über philippinische Obriminen, mit einer Neubeschreibung (Phasmatodea: Heteropterygidae: Obriminae). Entomologische Zeitschrift. 106 (11), 450–456. Zompro, O. (1998) Eine verkannte Planispectrum-Art aus Zentral-Borneo (Phasmatodea). Entomologische Zeitschrift, 108 (2), 83–84. Zompro, O. (2000) Die Phasmidensammlung des Übersee-Museums Bremen. TenDenZen Supplement 1999, pp. 61–70. Zompro, O. (2004) Revision of the genera of the Areolatae, including the status of Timema and Agathemera (Insecta, Phasmatodea). Abhandlungen des Naturwissenschaftlichen Vereins in Hamburg, (NF) 37, 327 pp, 161 figs. Zompro, O. (2005a) Inter- and intra-ordinal relationships of the Mantophasmatodea, with comments on the phylogeny of polyneopteran orders (Insecta: Polyneoptera). Mitteilungen des Geologisch-Paläontologischen Instituts der Universität Hamburg, 89, 85–116. Zompro, O. (2005b) Catalogue of type-material of the insect order Phasmatodea, housed in the Museum für Naturkunde der Humboldt-Universität zu Berlin, Germany and in the Institut der Martin-Luther-Universität in Halle (Saale), Germany. Mitteilungen aus dem Museum für Naturkunde Berlin, Deutsche entomologische Zeitschrift, 52, 251–290. http://dx.doi.org/10.1002/mmnd.20041018 Zompro, O. & Eusebio, O.L. (2002) A new species of stick insect (Phasmatodea: Heteropterygidae: Obriminae: Obrimini) from Cebu Island, philippines. Philippine Entomologist, 15 (1), 23–26.



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FIGURE 1. Acanthotaxy of Heteropteryginae


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FIGURES 2–10. Subfamily Dataminae Rehn & Rehn, 1939 2. Dares verrucosus Redtenbacher, 1906 ♀ (45.2 mm): W-Sabah, Kinabatangan District, Sepilok [coll. FH, No. 0074-20]; 3. Planispectrum bengalensis (Redtenbacher, 1906) ♀ (27.5 mm): Peninsular Malaysia [coll. FH, No. 0171-1]; 4. Pylaemenes shirakii Ho & Brock, 2013 ♀ (47.0 mm): Japan, Okinawa, Kunigasira [coll. FH, No. 0879-1]; 5. Dares verrucosus Redtenbacher, 1906 ♂ (38.5 mm): Captive reared from Sabah [coll. FH, No. 0074-2]; 6. Dares ulula (Westwood, 1859) ♀ (52.5 mm): N-Sarawak, Niah N.P. [coll. FH, No. 0090-1]; 7. Epidares nolimetangere (Haan, 1842) ♀ (41.0 mm): NW-Sarawak, Mount Serapi [coll. FH, No. 0098-31]; 8. Pylaemenes coronatus (Haan, 1842) ♀ (59.0 mm): Indonesia, Prov. Maluka, Seram [coll. FH, No. 0150-4]; 9. Pylaemenes coronatus (Haan, 1842) ♂ (51.5 mm): Indonesia, Prov. Maluka, Seram [coll. FH, No. 0150-11]; 10. Prosternum and profurcasternum of ♀ Pylaemenes coronatus (Haan, 1842) showing the sensory-areas marked by arrows [coll. FH, No. 0150-4].



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FIGURES 11–20. Eggs of Dataminae Rehn & Rehn, 1939 (a = dorsal view, b = lateral view, c = caudal view) 11 a–c. Dares validispinus Stål, 1875; 12 a–c. Dares verrucosus Redtenbacher, 1906; 13 a–c. Dares ziegleri Zompro & Fritzsche, 1999; 14 a–c. Dares ulula (Westwood, 1859); 15 a–c. Epidares nolimetangere (Haan, 1842); 16 a–c. Orestes mouhotii (Bates, 1865); 17 a–c. Pylaemenes mitratus (Redtenbacher, 1906); 18 a–c. Pylaemenes borneensis borneensis Bragg, 1998; 19 a–c. Planispectrum hongkongense Zompro, 2004 (enlarged); 20 a–c. Spinodares jenningsi Bragg, 1998.


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FIGURES 21–26. Obriminae: Obrimini Brunner v. Wattenwyl, 1907 (Obrimus-group) 21. Obrimus bicolanus Rehn & Rehn, 1939 ♀ (93.0 mm): Captive reared from Philippines, N-Luzon, Prov. Bicol, Sorsogon, Mount Pulog [coll. FH, No. 0730-5]; 22. Obrimus bicolanus Rehn & Rehn, 1939 ♂ (65.8 mm): Captive reared from Philippines, N-Luzon, Prov. Bicol, Sorsogon, Mount Pulog [coll. FH, No. 0730-24]; 23. Brasidas samarensis Rehn & Rehn, 1939 ♀ (95.0 mm): Captive reared from Philippines, Samar Id. [coll. FH, No. 0496-6]; 24. Brasidas samarensis Rehn & Rehn, 1939 ♂ (60.0 mm): Captive reared from Philippines, Samar Id. [coll. FH, No. 0496-20]; 25. Trachyaretaon sp. ♀ (77.5 mm): Captive reared from Philippines, N-Negros Id., Mount Kanlaon [coll. FH, No. 0716-2]; 26. Trachyaretaon sp. ♂ (61.0 mm): Philippones, Panay Id., Mount Nangtud [coll. FH, No. 0717-9].



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FIGURES 27–30. Obriminae: Obrimini Brunner v. Wattenwyl, 1907 (Obrimus-group) 27. Aretaon asperrimus (Redtenbacher, 1906) ♀ (77.0 mm): N-Sabah, Mount Kinabalu, Poring Hot Springs 480 m [coll. FH, No. 0118-13]; 28. Aretaon asperrimus (Redtenbacher, 1906) ♂ (53.5 mm): N-Sabah, Mount Kinabalu, Poring Hot Springs 480 m [coll. FH, No. 0118-20]; 29. Mearnsiana bullosa Rehn & Rehn, 1939 ♀ (95.0 mm): Philippines, Mindanao, South Cotabato Prov., Mount Apo [coll. FH, No. 0600-3]; 30. Mearnsiana bullosa Rehn & Rehn, 1939 ♂ (51.5 mm): Captive reared from Philippines, Mindanao, South Cotobato Prov. [coll. FH, No. 0600-11].


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FIGURES 31–33. Obriminae: Obrimini Brunner v. Wattenwyl, 1907 (Stenobrimus-group) 31. Stenobrimus bolivari Redtenbacher, 1906 ♀ (73.6 mm): Captive reared from Philippines, Luzon, Prov. Quezon, Quezon N.P. [coll. FH, No. 0680-4]; 32. Stenobrimus bolivari Redtenbacher, 1906 ♂ (57.8 mm): Captive reared from Philippines, Luzon, Prov. Quezon, Quezon N.P. [coll. FH, No. 0680-13]; 33. Stenobrimus bolivari Redtenbacher, 1906 ♀, apex of abdomen in dorsal view showing the multi-dentate apex of the subgenital plate: Philippines, Luzon, Prov. Quezon, Quezon N.P. [coll. FH, No. 0680-4].



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FIGURES 34–37. Obriminae: Obrimini Brunner v. Wattenwyl, 1907 (Theramenes-group) 34. Theramenes exiguus Hennemann & Conle, 2003 ♀ PT (82.5 mm): Philippines, Panay Id., Mount Nangtud [coll. FH, No. 0236-1]; 35. Theramenes exiguus Hennemann & Conle, 2003 ♂ PT (54.0 mm): Philippines, Panay Id., Mount Nangtud [coll. FH, No. 0236-7]; 36. Eubulides igorrote Rehn & Rehn, 1939 ♀ (75.0 mm): Captive reared from Philippines, Luzon, Prov. Quezon, Quezon N.P. [coll. FH, No. 0685-1]; 37. Eubulides igorrote Rehn & Rehn, 1939 ♂ (55.5 mm): Captive reared from Philippines, Luzon, Prov. Quezon, Quezon N.P. [coll. FH, No. 0685-5].


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FIGURES 38–46. Obriminae: Obrimini Brunner v. Wattenwyl, 1907 38. Metasternum of ♀ Euobrimus lacerta Redtenbacher, 1906 showing the metasternal foramina: Captive reared from Philippines, N-Luzon, Prov. Bicol, Sorsogon, Mount Pulog [coll. FH, species No. 0768]; 39. Metasternum of ♀Obrimus bicolanus Rehn & Rehn, 1939 showing the metasternal foramina: Captive reared from Philippines, N-Luzon, Prov. Bicol, Sorsogon, Mount Pulog [coll. FH, species No. 0730]; 40. Metasternum of ♀ Brasidas foveolatus Rehn & Rehn, 1939 showing the metasternal foramina: Captive reared from Philippines, Mindanao, South Cotobato Prov., Mount Apo [coll. FH, species No. 0316]; 41. Prosternum and profurcasternum of ♀ Brasidas samarensis Rehn & Rehn, 1939 showing the two prosternal sensoryareas marked by arrows [coll. FH, species No. 0496]; 42. Prosternum and profurcasternum of ♀ Mearnsiana bullosa Rehn & Rehn, 1939 showing the two prosternal sensory-areas marked by arrows [coll. FH, species No. 0600]; 43. Aretaon sp. ♀, apex of abdomen in lateral view: Captive reared from Palawan, Brooke´s Point, Mount Gantung [coll. FH, species No. 0731]; 44. Aretaon sp. ♀, apex of abdomen in dorsal view: Captive reared from Palawan, Brooke´s Point, Mount Gantung [coll. FH, species No. 0731]; 45. Obrimus bicolanus Rehn & Rehn, 1939 ♀, apex of abdomen in lateral view: Captive reared from Palawan, Brooke´s Point, Mount Gantung [coll. FH, species No. 0730]; 46 Obrimus bicolanus Rehn & Rehn, 1939 ♀, apex of abdomen in dorsal view: Captive reared from Palawan, Brooke´s Point, Mount Gantung [coll. FH, species No. 0730].



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FIGURES 47–56. Eggs of Obriminae: Obrimini Brunner v. Wattenwyl, 1907 (a = dorsal view, b = lateral view) 47 a–b. Aretaon asperrimus (Redtenbacher, 1906); 48 a–b. Trachyaretaon sp., Philippines, N-Luzon [coll. FH, No. 0497]; 49 a–b. Trachyaretaon sp., Philippines, Panay Id. [coll. FH, No. 0717]; 50 a–b. Brasidas samarensis Rehn & Rehn, 1939; 51 a–b. Eubulides igorrote Rehn & Rehn, 1939; 52 a–b. Heterocopus leprosus Redtenbacher, 1906; 53 a–b. Sungaya inexpectata Zompro, 1996; 54 a–b. Stenobrimus tagalog Rehn & Rehn, 1939; 55 a–b. Theramenes olivaceus (Westwood, 1859); 56 a–b. Theramenes exiguus Hennemann & Conle, 2003.


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FIGURES 57–66. Obriminae: Tisamenini n. trib. 57. Hoploclonia abercrombiei Bragg, 1998 ♀ (52.8 mm): Captive reared from N-Sarawak, Niah N.P. [coll. FH, No. 0238-4]; 58. Hoploclonia gecko (Westwood, 1859) ♀ (47.0 mm): Captive reared from NW-Sarawak, Bako N.P. [coll. FH, No. 0117-12]; 59. Hoploclonia gecko (Westwood, 1859) ♂ (33.5 mm): Captive reared from NW-Sarawak, Bako N.P. [coll. FH, No. 0117-19]; 60. Ilocano hebardi Rehn & Rehn, 1939 ♀ HT (30.0 mm): Philippines, Luzon, Baguio, Benguet [ANSP]; 61. Pterobrimus depressus Redtenbacher, 1906 ♂ ST (25.0 mm): “Fiji” [ZMUH]; 62. Tisamenus spadix (Rehn & Rehn, 1939) ♀ (61.0 mm): Philippines, Panay Id., Mount Nangtud [coll. FH, No. 0271-1]; 63. Tisamenus spadix (Rehn & Rehn, 1939) ♂ (42.7 mm): Philippines, Panay Id., Mount Nangtud [coll. FH, No. 0271-2]; 64. Prosternum and profurcasternum of ♀ Tisamenus serratorius Stål, 1875 showing the two prosternal sensory-areas marked by arrows [coll. FH, species No. 0727]; 65. Tisamenus serratorius Stål, 1875 ♀, apex of abdomen in lateral view: Captive reared from Philippines, Luzon, Prov. Quezon, Quezon N.P. [coll. FH, No. 0727-12]; 66. Hoploclonia gecko (Westwood, 1859) ♀, apex of abdomen in lateral view showing the fused tergum X and epiproct: Captive reared from from NW-Sarawak, Bako N.P. [coll. FH, species No. 0117-5].



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FIGURES 67–71. Eggs of Obriminae: Tisamenini n. trib. (a = dorsal view, b = lateral view, c = caudal view) 67 a–c. Hoploclonia gecko (Westwood, 1859); 68 a–c. Hoploclonia cuspidata Redtenbacher, 1906; 69 a–c. Hoploclonia abercrombiei Bragg, 1998; 70 a–c. Tisamenus hystrix (Rehn & Rehn, 1939); 71 a–c. Tisamenus spadix (Rehn & Rehn, 1939).


HENNEMANN ET AL.

FIGURES 72–77. Obriminae: Miroceramiini Zompro, 2004 72. Miroceramia westwoodii (Bates, 1865) ♂ dorsal view (47.5 mm): Indonesia, Maluka, NC-Seram, Manusela N.P., Wae Mual Plain [NHMUK]; 73. Miroceramia westwoodii (Bates, 1865) ♂ lateral view (47.5 mm): Indonesia, Maluka, NC-Seram, Manusela N.P., Wae Mual Plain [NHMUK]; 74. Miroceramia westwoodii (Bates, 1865) ♀ (66.0 mm): Indonesia, Maluka, Central Seram, Manacia 2500 ft. [NHMUK]; 75. Prosternum and profurcasternum of ♀ Miroceramia westwoodii (Bates, 1865) showing the two prosternal sensory-areas marked by arrows [NHMUK]; 76. Miroceramia westwoodii (Bates, 1865) egg in dorsal view (ex ovipositor HT): Indonesia, N-Sulawesi, Menado [UMO]; 77. Miroceramia westwoodii (Bates, 1865) egg in lateral view (ex ovipositor HT): Indonesia, N-Sulawesi, Menado [UMO].



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FIGURES 78–83. Sensory-area of profurcasternum of Heteropteryginae Kirby, 1896 [scale = 1mm] 78. Haaniella echinata (Redtenbacher, 1906) ♀ [coll. FH, No. 0230-16]; 79. Haaniella glaber (Redtenbacher, 1906) ♀ [RMNH]; 80. Haaniella macroptera n. sp. ♀ PT [coll. FH, No. 0869-2]; 81. Haaniella parva Günther, 1944 ♀ [coll. FH, No. 0867-1]; 82. Haaniella rosenbergii (Kaup, 1871) ♀ [coll. FH, No. 0114-1]; 83. Haaniella saussurei Kirby, 1904 ♀ [coll. FH, No. 0231-2].


HENNEMANN ET AL.

FIGURE 84. Right ala of ♀ Heteropteryx dilatata (Parkinson, 1789) showing the venation and tympanal area. The enlarged inset shows a portion of the first anal vein (A1) which is densely set with minute spine-like structures and part of the stridulatory organ. Legend: TA = tympanal area; C = Costa; Sc = Subcosta; R = Radial; M = Media; Cu = Cubital; P = Plical; E = Empusal; A = Anal; Ic = Intercalary.



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FIGURES 85–88. Ventral view of basal abdominal sternites of ♀♀ of Haaniella spp. (Heteropteryginae) showing the lateral invaginations. 85. Haaniella saussurei Kirby, 1904 [coll. FH, No. 0231-10]; 86. Haaniella grayii (Westwood, 1859) [coll. FH, No. 0116-38]; 87. Haaniella scabra (Redtenbacher, 1906) [coll. FH, No. 0328-12]; 88. Haaniella parva Günther, 1944 [coll. FH, No. 0867-1].


HENNEMANN ET AL.

FIGURES 89–92. Ventral view of basal abdominal sternites of ♂♂ of Haaniella spp. (Heteropteryginae) showing the lateral invaginations. 89. Haaniella grayii (Westwood, 1859) [coll. FH, No. 0116-39]; 90. Haaniella echinata (Redtenbacher, 1906) [coll. FH, No. 0230-28]; 91. Haaniella scabra (Redtenbacher, 1906) [coll. FH, No. 0328-16]; 92. Haaniella parva Günther, 1944 [coll. FH, No. 0867-3].

FIGURES 93–96. Apex of metatibia of Heteropteryginae showing the medio-apical spine of the area apicalis 93. Heteropteryx dilatata (Parkinson, 1798); 94. Haaniella erringtoniae (Redtenbacher, 1906); 95. Haaniella grayii (Westwood, 1859); 96. Haaniella scabra (Redtenbacher, 1906).



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FIGURE 97. Relationships within the family Heteropterygidae. (a) according to Bradler (2009), modified; (b) according to the present analysis (*/** not true for all taxa contained; *** homoplasy). All characters mentioned are apomorphies.


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FIGURES 98–104. Haaniella aculeata n. sp. 98. ♂, HT dorsal view (70.5 mm): W-Sumatra, Prov. Sumatera Barat, Balun [RMNH]; 99. ♂, HT lateral view (70.5 mm): WSumatra, Prov. Sumatera Barat, Balun [RMNH]; 100. ♂, HT: apex of abdomen in lateral view [RMNH]; 101. ♂, HT: apex of abdomen in dorsal view [RMNH]; 102. ♂, HT: apex of abdomen in ventral view [RMNH]; 103. ♂, HT: head, pro- and mesothorax in lateral view [RMNH]; 104. ♂, HT: head, pro- and mesothorax in dorsal view [RMNH].



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FIGURES 105–108. Haaniella erringtoniae (Redtenbacher, 1906) 105. ♀ dorsal view (106.0 mm): Captive reared from Peninsular Malaysia, Selangor, Templer Park [coll. FH, No. 0385-14]; 106. ♀ dorsolateral view (105.0 mm): Captive reared from Peninsular Malaysia, Selangor, Templer Park [coll. FH, No. 038515]; 107. ♂ dorsal view (75.5 mm): Peninsular Malaysia, Pahang, Genting Highlands 1500-1600 m [coll. FH, No. 0385-22]; 108. ♂ dorsal view (77.0 mm): Captive reared from Peninsular Malaysia, Selangor, Templer Park [coll. FH, No. 0385-5].


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FIGURES 109–114. Haaniella erringtoniae (Redtenbacher, 1906) 109. ♀ head, pro- and mesothorax in lateral view [coll. FH, No. 0385-15]; 110. ♀ head, pro- and mesothorax in dorsal view [coll. FH, No. 0385-15]; 111. ♀ head, pro- and mesothorax in ventral view [coll. FH, No. 0385-15]; 112. ♂ head, pro- and mesothorax in lateral view [coll. FH, No. 0385-19]; 113. ♂ head, pro- and mesothorax in dorsal view [coll. FH, No. 0385-19]; 114. ♂ head, pro- and mesothorax in ventral view [coll. FH, No. 0385-19].



143

FIGURES 115–120. Haaniella erringtoniae (Redtenbacher, 1906) 115. ♀ apex of abdomen in lateral view [coll. FH, No. 0385-16]; 116. ♀ apex of abdomen in dorsal view [coll. FH, No. 038516]; 117. ♀ apex of abdomen in ventral view [coll. FH, No. 0385-16]; 118. ♂ apex of abdomen in lateral view [coll. FH, No. 0385-18]; 119. ♂ apex of abdomen in dorsal view [coll. FH, No. 0385-18]; 120. ♂ apex of abdomen in ventral view [coll. FH, No. 0385-18].


HENNEMANN ET AL.

FIGURES 121–125. Haaniella gintingi n. sp. 121. ♀, HT dorsal view (98.4 mm): N-Sumatra: Prov. Aceh, Gunung Leuser NP, Ketambe, Atjeh, Tenggara 350 m [RMNH]; 122. ♀, PT dorsal view (92.0 mm): Captive reared from NW-Sumatra: Prov. Sumatera Utara, Mount Sibayak 1400–1600 m [coll. FH, No. 0753-6]; 123 ♂, PT dorsal view (61.0 mm): Captive reared from NW-Sumatra: Prov. Sumatera Utara, Mount Sibayak 1400–1600 m [coll. FH, No. 0753-8]; 124. ♀, PT lateral view (91.0 mm): Captive reared from NW-Sumatra: Prov. Sumatera Utara, Mount Sibayak 1400–1600 m [coll. FH, No. 0753-5]; 125. ♂, PT lateral view (61.0 mm): Captive reared from NW-Sumatra: Prov. Sumatera Utara, Mount Sibayak 1400-1600 m [coll. FH, No. 0753-8].



145

FIGURES 126–131. Haaniella gintingi n. sp. 126. ♀, PT head, pro- and mesothorax in lateral view [coll. FH, No. 0753-7]; 127. ♀, PT head, pro- and mesothorax in dorsal view [coll. FH, No. 0753-5]; 128. ♀, PT head, pro- and mesothorax in ventral view [coll. FH, No. 0753-7]; 129. ♂, PT head, pro- and mesothorax in lateral view [coll. FH, No. 0753-8]; 130. ♂, PT head, pro- and mesothorax in dorsal view [coll. FH, No. 0753-10]; 131. ♂, PT head, pro- and mesothorax in ventral view [coll. FH, No. 0753-10].


HENNEMANN ET AL.

FIGURES 132–137. Haaniella gintingi n. sp. 132. ♀, PT apex of abdomen in lateral view [coll. FH, No. 0753-5]; 133. ♀, PT apex of abdomen in dorsal view [coll. FH, No. 0753-5]; 134. ♀, PT apex of abdomen in ventral view [coll. FH, No. 0753-5]; 135. ♂, PT apex of abdomen in lateral view [coll. FH, No. 0753-8]; 136. ♂, PT apex of abdomen in dorsal view [coll. FH, No. 0753-8]; 137. ♂, PT apex of abdomen in ventral view [coll. FH, No. 0753-8].



147

FIGURES 138–140. Haaniella glaber (Redtenbacher, 1906) 138. ♀, HT dorsal view (86.0 mm): E-Sumatra, Prov. Riau, Lakat [MHNG]; 139. ♀ dorsal view (86.5 mm): S-Sumatra, Prov. Lampung, Mount Tanggamus, Giesting 750 m [RMNH]; 140. ♀ dorsolateral view (86.5 mm): S-Sumatra, Prov. Lampung, Mount Tanggamus, Giesting 750 m [RMNH].


HENNEMANN ET AL.

FIGURES 141–143. Haaniella glaber (Redtenbacher, 1906) 141. ♂ dorsal view (68.0 mm): W-Sumatra, Prov. Sumatera Utara, Tapanuli Distr., Kapanoek River [RMNH]; 142. ♂ dorsolateral view (68.0 mm): W-Sumatra, Prov. Sumatera Utara, Tapanuli Distr., Kapanoek River [RMNH]; 143. ♂ lateral view (69.8 mm): SW-Sumatra, Prov. Benkulu, Lebong Tandai [MNHN].



149

FIGURES 144–152. Haaniella glaber (Redtenbacher, 1906) 144. ♀ head, pro- and mesothorax in dorsolateral view [RMNH]; 145. ♂ head, pro- and mesothorax in dorsal view [RMNH]; 146. ♂ head, pro- and mesothorax in lateral view [RMNH]; 147. ♀, HT apex of abdomen in lateral view [MHNG]; 148. ♀ apex of abdomen in dorsal view [RMNH]; 149. ♀ apex of abdomen in ventral view [RMNH]; 150. ♂ apex of abdomen in lateral view [MHNG]; 151. ♂ apex of abdomen in dorsal view [RMNH]; 152. ♂ apex of abdomen in ventral view [RMNH].


HENNEMANN ET AL.

FIGURES 153–156. Haaniella jacobsoni Günther, 1944 153. ♀, AT dorsolateral view (90.4 mm): Simeulue Island, Sinabang [RMNH]; 154. ♀, AT dorsal view (90.4 mm): Simeulue Island, Sinabang [RMNH]; 155. ♂, HT dorsal view (68.8 mm): Simeulue Island, Sinabang [RMNH]; 156. ♂, HT lateral view (68.8 mm): Simeulue Island, Sinabang [RMNH].



151

FIGURES 157–160. Haaniella jacobsoni Günther, 1944 157. ♀, AT apex of abdomen in lateral view [RMNH]; 158. ♀, AT apex of abdomen in dorsal view [RMNH]; 159. ♂, PT apex of abdomen in lateral view [SMTD]; 160. ♂, PT apex of abdomen in dorsal view [SMTD].


HENNEMANN ET AL.

FIGURES 161–165. Haaniella kerincia n. sp. 161. ♀, PT dorsal view (56.0 mm): W-Sumatra, Prov. Jambi, Kerinci Seblat N.P., Mount Kerinci 1500–2000 m [ZIN]; 162. ♀, PT dorsal view (57.0 mm): W-Sumatra, Prov. Jambi, Kerinci Seblat N.P., Mount Kerinci 1500–2000 m [ZIN]; 163. ♂, PT dorsal view (40.0 mm): W-Sumatra, Prov. Jambi, Kerinci Seblat N.P., Mount Kerinci 1500–2000 m [ZIN]; 164. ♀, PT lateral view (57.0 mm): W-Sumatra, Prov. Jambi, Kerinci Seblat N.P., Mount Kerinci 1500–2000 m [ZIN]; 165. ♂, PT dorsolateral view (40.0 mm): W-Sumatra, Prov. Jambi, Kerinci Seblat N.P., Mount Kerinci 1500–2000 m [ZIN].



153

FIGURES 166–170. Haaniella kerincia n. sp. 166. ♀, PT apex of abdomen in lateral view [ZIN]; 167. ♀, PT apex of abdomen in dorsal view [ZIN]; 168. ♂, HT apex of abdomen in lateral view [ZIN]; 169. ♂, HT apex of abdomen in dorsal view [ZIN]; 170. ♀, PT head, pro- and mesothorax in dorsolateral view [ZIN].


HENNEMANN ET AL.

FIGURES 171–174. Haaniella macroptera n. sp. 171. ♀, PT dorsal view (122.0 mm): Singapore, Nee Soon swamp forest [coll. FH, No. 0860-2]; 172. ♀, PT dorsolateral view (122.0 mm): Singapore, Nee Soon swamp forest [coll. FH, No. 0860-2]; 173. ♂, PT dorsal view (83.0 mm): Singapore, Nee Soon swamp forest [coll. FH, No. 0860-1]; 174. ♂, PT dorsolateral view (83.0 mm): Singapore, Nee Soon swamp forest [coll. FH, No. 0860-1].



155

FIGURES 175–181. Haaniella macroptera n. sp. 175. ♀, PT apex of abdomen in lateral view [coll. FH, No. 0860-2]; 176. ♀, PT apex of abdomen in dorsal view [coll. FH, No. 0860-2]; 177. ♀, PT apex of abdomen in ventral view [coll. FH, No. 0860-2]; 178. ♂, PT apex of abdomen in lateral view [coll. FH, No. 0860-1]; 179. ♂, PT apex of abdomen in dorsal view [coll. FH, No. 0860-1]; 180. ♂, PT apex of abdomen in ventral view [coll. FH, No. 0860-1]; 181. ♀, PT head, pro- and mesothorax in dorsolateral view [coll. FH, No. 0860-2].


HENNEMANN ET AL.

FIGURES 182–185. Haaniella mecheli (Redtenbacher, 1906) 138. ♂, HT dorsal view (74.1 mm): E-Sumatra, Prov. Riau, Indragiri Hulu [NHMB]; 183. ♂, HT apex of abdomen in lateral view [NHMB]; 184. ♂, HT apex of abdomen in dorsal view [NHMB]; 185. ♂, HT apex of abdomen in ventral view [NHMB].



157

FIGURES 186–188. Haaniella muelleri (Haan, 1842) 186. ♀, PLT dorsal view (84.0 mm): W-Sumatra, Prov. Sumatera Utara, Batang Singgalang [RMNH]; 187. ♀ lateral view (96.0 mm): N-Sumatra, Prov. Benkulu, Lebong Tandai [MNHN]; 188. ♂, LT dorsolateral view (60.0 mm): W-Sumatra, Prov. Sumatera Utara, Batang Singgalang [RMNH].


HENNEMANN ET AL.

FIGURES 189–196. Haaniella muelleri (Haan, 1842) 189. ♂, LT head, pro- and mesothorax in dorsal view [RMNH]; 190. ♀ head, pro- and mesothorax in lateral view [ZMUH]; 191. ♀ head, pro- and mesothorax in dorsal view [ZMUH]; 192. ♀ apex of abdomen in lateral view [ZMUH]; 193. ♀ apex of abdomen in dorsal view [ZMUH]; 194. ♀ apex of abdomen in ventral view [ZMUH]; 195. ♂, LT dorsal view of abdominal tergum X and epiproct [RMNH]; 196. ♂, LT apex of abdomen in lateral view [RMNH].



159

FIGURES 197–199. Haaniella rosenbergii (Kaup, 1871) 197. ♀ dorsal view (84.5 mm): NW-Sumatra, Prov. Sumatera Barat, Padang [coll. FH, No. 0114-2]; 198. ♀ dorsolateral view (84.5 mm): NW-Sumatra, Prov. Sumatera Barat, Padang [coll. FH, No. 0114-2]; 199. ♀ (HT of synonym H. muelleri novaeguineae Günther, 1930) dorsolateral view (86.5 mm) [ZMUH].


HENNEMANN ET AL.

FIGURES 200–202. Haaniella rosenbergii (Kaup, 1871) 200. ♂ dorsal view (64.8 mm): NW-Sumatra, Prov. Aceh, Subulussalam [coll. FH, No. 0114-5]; 201. ♂ (AT of synonym H. muelleri novaeguineae Günther, 1930) dorsolateral view (67.0 mm) [ZMUH]; 202. ♂ var. dorsolateral view (51.0 mm): NWSumatra, Prov. Sumatera Barat, Padang [coll. FH, No. 0114-5].



161

FIGURES 203–213. Haaniella rosenbergii (Kaup, 1871) 203. ♀ apex of abdomen in lateral view [coll. FH, No. 0114-4]; 204. ♀ apex of abdomen in dorsal view [coll. FH, No. 0114-4]; 205. ♀ apex of abdomen in ventral view [coll. FH, No. 0114-4]; 206. ♂ apex of abdomen in lateral view [coll. FH, No. 0114-5]; 207. ♂ apex of abdomen in dorsal view [coll. FH, No. 0114-5]; 208. ♂ apex of abdomen in ventral view [coll. FH, No. 0114-5]; 209. ♀ head, pro- and mesothorax in lateral view [coll. FH, No. 0114-2]; 210. ♀ head, pro- and mesothorax in dorsal view [coll. FH, No. 0114-2]; 211. ♂ head, pro- and mesothorax in lateral view [coll. FH, No. 0114-5]; 212. ♂ head, pro- and mesothorax in dorsal view [coll. FH, No. 0114-5]; 213. ♂ (AT of synonym H. muelleri novaeguineae Günther, 1930) head, pro- and mesothorax in dorsal view [ZMUH].


HENNEMANN ET AL.

FIGURES 214–216. Haaniella dehaanii (Westwood, 1859) 214. ♀ dorsal view (95.0 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0115-18]; 215. ♀ dorsolateral view (95.0 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0115-18]; 216. ♀ dorsolateral view (89.0 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0115-23].



163

FIGURES 217–219. Haaniella dehaanii (Westwood, 1859) 217. ♂ dorsal view (71.5 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0115-7]; 218. ♂ dorsolateral view (71.5 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0115-7]; 219. ♂ dorsolateral view (68.5 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0115-29].


HENNEMANN ET AL.

FIGURES 220–229. Haaniella dehaanii (Westwood, 1859) 220. ♀ apex of abdomen in lateral view [coll. FH, No. 0115-18]; 221. ♀ apex of abdomen in dorsal view [coll. FH, No. 011518]; 222. ♀ apex of abdomen in ventral view [coll. FH, No. 0115-18]; 223. ♂ apex of abdomen in lateral view [coll. FH, No. 0115-28]; 224. ♂ apex of abdomen in dorsal view [coll. FH, No. 0115-28]; 225. ♂ apex of abdomen in ventral view [coll. FH, No. 0115-28]; 226. ♂ head, pro- and mesothorax in dorsal view [coll. FH, No. 0115-30]; 227. ♂ head, pro- and mesothorax in lateral view [coll. FH, No. 0115-5]; 228. ♀ head, pro- and mesothorax in dorsal view [coll. FH, No. 0115-19]; 229. ♀ head, proand mesothorax in lateral view [coll. FH, No. 0115-19].



165

FIGURES 230–233. Haaniella gorochovi n. sp. 230. ♀, PT dorsal view (101.0 mm): S-Vietnam, Prov. Gia Lai, Buôn Luoi [ZIN]; 231. ♀, PT dorsal view (96.0 mm): SVietnam, Prov. Gia Lai, Buôn Luoi [ZIN]; 232. ♂, HT dorsal view (61.0 mm): S-Vietnam, Prov. Gia Lai, Buôn Luoi [ZIN]; 233. ♂, PT dorsal view (59.0 mm): S-Vietnam, Prov. Gia Lai, Tram Lap [ZIN].


HENNEMANN ET AL.

FIGURES 234–242. Haaniella gorochovi n. sp. 234. ♀, PT head, pro- and mesothorax in dorsal view [RBINS]; 235. ♀, PT head, pro- and mesothorax in lateral view [RBINS]; 236. ♀, PT head, pro- and mesothorax in dorsolateral view [ZIN]; 237. ♀, HT head, pro- and mesothorax in lateral view [ZIN]; 238. ♀, PT apex of abdomen in lateral view [ZIN]; 239. ♀, PT apex of abdomen in dorsal view [ZIN]; 240. ♀, PT apex of abdomen in ventral view [ZIN]; 241. ♂, HT apex of abdomen in lateral view [ZIN]; 242. ♂, HT apex of abdomen in dorsal view [ZIN].



167

FIGURES 243–245. Haaniella grayii (Westwood, 1859) 243. ♀ dorsal view (124.0 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0116-6]; 244. ♀ dorsolateral view (128.5 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0116-3]; 245. ♀ dorsolateral view (128.0 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0116-39].


HENNEMANN ET AL.

FIGURES 246–253. Haaniella grayii (Westwood, 1859) 246. ♂ dorsal view (93.0 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0116-40]; 247. ♂ dorsolateral view (93.0 mm): Captive reared from NW-Sarawak, Mount Serapi [coll. FH, No. 0116-40]; 248. ♂ apex of abdomen in lateral view [coll. FH, No. 0116-40]; 249. ♂ apex of abdomen in dorsal view [coll. FH, No. 0116-40]; 250. ♂ apex of abdomen in ventral view [coll. FH, No. 0116-40]; 251. ♂ head, pro- and mesothorax in lateral view [coll. FH, No. 0116-40]; 252. ♂ head, pro- and mesothorax in dorsal view [coll. FH, No. 0116-40]; 253. ♂ head, pro- and mesothorax in ventral view [coll. FH, No. 0116-40].



169

FIGURES 254–259. Haaniella grayii (Westwood, 1859) 254. ♀ apex of abdomen in lateral view [coll. FH, No. 0116-38]; 255. ♀ apex of abdomen in dorsal view [coll. FH, No. 011638]; 256. ♀ apex of abdomen in ventral view [coll. FH, No. 0116-37]; 257. ♀ head, pro- and mesothorax in lateral view [coll. FH, No. 0116-39]; 258. ♀ head, pro- and mesothorax in dorsal view [coll. FH, No. 0116-39]; 259. ♀ head, pro- and mesothorax in ventral view [coll. FH, No. 0116-39].


HENNEMANN ET AL.

FIGURES 260–263. Haaniella parva Günther, 1944 260. ♀ dorsal view (52.0 mm): NW-Sumatra, Prov. Aceh, Subulussalam [coll. FH, No. 0867-1]; 261. ♀ dorsolateral view (52.0 mm): NW-Sumatra, Prov. Aceh, Subulussalam [coll. FH, No. 0867-1]; 262. ♀ dorsal view (35.7 mm): NW-Sumatra, Prov. Aceh, Subulussalam [coll. FH, No. 0867-3]; 263. ♀ dorsolateral view (35.7 mm): NW-Sumatra, Prov. Aceh, Subulussalam [coll. FH, No. 0867-3].



171

FIGURES 264–273. Haaniella parva Günther, 1944 264. ♀ apex of abdomen in lateral view [coll. FH, No. 0867-1]; 265. ♀ apex of abdomen in dorsal view [coll. FH, No. 0867-1]; 266. ♀ apex of abdomen in ventral view [coll. FH, No. 0867-1]; 267. ♂ apex of abdomen in lateral view [coll. FH, No. 0867-3]; 268. ♂ apex of abdomen in dorsal view [coll. FH, No. 0867-3]; 269. ♂ apex of abdomen in ventral view [coll. FH, No. 0867-3]; 270. ♀ head, pro- and mesothorax in lateral view [coll. FH, No. 0867-1]; 271. ♀ head, pro- and mesothorax in dorsal view [coll. FH, No. 0867-1]; 273. ♂ head, pro- and mesothorax in lateral view [coll. FH, No. 0867-3]; 273. ♂ head, pro- and mesothorax in dorsal view [coll. FH, No. 0867-3].


HENNEMANN ET AL.

FIGURES 274–277. Haaniella echinata (Redtenbacher, 1906) 274. ♀ dorsolateral view (115.0 mm): “Borneo” [NHMW]; 275. ♀ dorsolateral view (117.0 mm): N-Sarawak, Niah N.P. [coll. FH, No. 0230-2]; 276. ♀ dorsolateral view (112.0 mm): Captive reared from Brunei, Kuala Belalong, Badas [coll. FH, No. 0230-14]; 277. ♀ dorsal view (113.0 mm): Captive reared from Brunei, Kuala Belalong, Badas [coll. FH, No. 0230-16].



173

FIGURES 278–281. Haaniella echinata (Redtenbacher, 1906) 278. ♂ dorsolateral view (83.8 mm): Captive reared from Brunei, Kuala Belalong, Badas [coll. FH, No. 0230-20]; 279. ♂ dorsal view (83.8 mm): Captive reared from Brunei, Kuala Belalong, Badas [coll. FH, No. 0230-20]; 280. ♂ dorsolateral view (81.5 mm): Captive reared from Brunei, Kuala Belalong, Badas [coll. FH, No. 0230-21]; 281. ♂ dorsal view (81.5 mm): Captive reared from Brunei, Kuala Belalong, Badas [coll. FH, No. 0230-21].


HENNEMANN ET AL.

FIGURES 282–290. Haaniella echinata (Redtenbacher, 1906) 282. ♀ head, pro- and mesothorax in dorsal view [coll. OC]; 283. ♀ head, pro- and mesothorax in lateral view [coll. FH, No. 0230-6]; 284. ♂ head, pro- and mesothorax in lateral view [coll. FH, No. 0230-4]; 285. ♀ apex of abdomen in lateral view [coll. OC]; 286. ♀ apex of abdomen in dorsal view [coll. FH, No. 0230-14]; 287. ♀ apex of abdomen in ventral view [coll. FH, No. 0230-14]; 288. ♂ apex of abdomen in lateral view [coll. FH, No. 0230-4]; 289. ♂ apex of abdomen in dorsal view [coll. FH, No. 0230-20]; 290. ♂ apex of abdomen in ventral view [coll. FH, No. 0230-20].



175

FIGURES 291–293. Haaniella saussurei Kirby, 1904 291. ♀ dorsal view (130.0 mm): Captive reared from NW-Sarawak, Tarum nr. Debak [coll. FH, No. 0231-9]; 292. ♀ dorsolateral view (130.0 mm): Captive reared from NW-Sarawak, Tarum nr. Debak [coll. FH, No. 0231-9]; 293. ♀ dorsal view (118.5 mm): Captive reared from NW-Sarawak, Tarum nr. Debak [coll. FH, No. 0231-10].


HENNEMANN ET AL.

FIGURES 294–300. Haaniella saussurei Kirby, 1904 294. ♂ dorsal view (82.5 mm): Captive reared from NW-Sarawak, Bako N.P. [coll. FH, No. 0231-4]; 295. ♂ dorsolateral view (82.5 mm): Captive reared from NW-Sarawak, Bako N.P. [coll. FH, No. 0231-4]; 296. ♂ apex of abdomen in lateral view [coll. FH, No. 0231-5]; 297. ♂ apex of abdomen in dorsal view [coll. FH, No. 0231-4]; 298. ♂ apex of abdomen in ventral view [coll. FH, No. 0231-5]; 299. ♂ head, pro- and mesothorax in lateral view [coll. FH, No. 0231-5]; 300. ♂ head, pro- and mesothorax in dorsal view [coll. FH, No. 0231-5].



177

FIGURES 301–305. Haaniella saussurei Kirby, 1904 301. ♀ apex of abdomen in lateral view [coll. FH, No. 0231-#]; 302. ♀ apex of abdomen in dorsal view [coll. FH, No. 0231-#]; 303. ♀ apex of abdomen in ventral view [coll. FH, No. 0231-9]; 304. ♀ head, pro- and mesothorax in lateral view [coll. FH, No. 0231-10]; 305. ♀ head, pro- and mesothorax in dorsal view [coll. FH, No. 0231-10].


HENNEMANN ET AL.

FIGURES 306–308. Haaniella scabra (Redtenbacher, 1906) 306. ♀ dorsal view (79.5 mm): Captive reared from Sabah, Mount Kinabalu [coll. FH, No. 0328-12]; 307. ♂ dorsal view (57.0 mm): Captive reared from Sabah, Mount Kinabalu [coll. FH, No. 0328-16]; 308. ♀ dorsal view (62.0 mm): Captive reared from Sabah, Mount Kinabalu [coll. FH, No. 0328-14].



179

FIGURES 309–312. Haaniella scabra (Redtenbacher, 1906) 309. ♀ dorsolateral view (63.5mm): Sabah, Mount Kinabalu N.P. near headquarters, Silau-Silau trial 1550 m [coll. FH, No. 0328-1]; 310. ♀ lateral view (61.5mm): Sabah, Mount Kinabalu N.P. near headquarters, Silau-Silau trial 1550 m [coll. FH, No. 0328-6]; 311. ♂ dorsolateal view (57.0 mm): Captive reared from Sabah, Mount Kinabalu [coll. FH, No. 0328-16]; 312. ♂ dorsolateral view (45.5mm): Sabah, Mount Kinabalu N.P. near headquarters, Silau-Silau trial 1550 m [coll. FH, No. 0328-3].


HENNEMANN ET AL.

FIGURES 313–322. Haaniella scabra (Redtenbacher, 1906) 313. ♀ head, pro- and mesothorax in lateral view [coll. FH, No. 0328-12]; 314. ♀ head, pro- and mesothorax in dorsal view [coll. FH, No. 0328-12]; 315. ♂ head, pro- and mesothorax in lateral view [coll. FH, No. 0328-17]; 316. ♂ head, pro- and mesothorax in dorsal view [coll. FH, No. 0328-17]; 317. ♀ apex of abdomen in lateral view [coll. FH, No. 0328-12]; 318. ♀ apex of abdomen in dorsal view [coll. FH, No. 0328-12]; 319. ♀ apex of abdomen in ventral view [coll. FH, No. 0328-12]; 320. ♂ apex of abdomen in lateral view [coll. FH, No. 0328-17]; 321. ♂ apex of abdomen in dorsal view [coll. FH, No. 0328-17]; 322. ♂ apex of abdomen in ventral view [coll. FH, No. 0328-17].



181

FIGURES 323–325. Heteropteryx dilatata (Parkinson, 1798) 323. ♀ with spread wings, dorsal view (156.0 mm): Peninsular Malaysia, Perak, Tapah Hills ca. 500 m [coll. FH, No. 0018-3]; 324. ♀ yellow variety, dorsal view (150.0 mm): Peninsular Malaysia, Perak, Tapah Hills ca. 500 m [coll. FH, No. 0018-4]; 325. ♀ dorsolateral view (159.0 mm): Peninsular Malaysia, Perak, Tapah Hills ca. 500 m [coll. FH, No. 0018-27].


HENNEMANN ET AL.

FIGURES 326–327. Heteropteryx dilatata (Parkinson, 1798) 326. ♂ dorsal view (92.0 mm): Peninsular Malaysia, Perak, Tapah Hills ca. 500 m [coll. FH, No. 0018-7]; 327. ♂ dorsolateral view (93.0 mm): Peninsular Malaysia, Perak, Tapah Hills ca. 500 m [coll. FH, No. 0018-6].



183

FIGURES 328–336. Heteropteryx dilatata (Parkinson, 1798) 328. ♂ dorsolateral view (98.0 mm): Captive reared from Peninsular Malaysia, Perak, Tapah Hills [coll. FH, No. 0018-16]; 329. ♀ apex of abdomen in lateral view [coll. FH, No. 0018-26]; 330. ♀ apex of abdomen in dorsal view [coll. FH, No. 001826]; 331. ♀ apex of abdomen in ventral view [coll. FH, No. 0018-26]; 332. ♂ apex of abdomen in lateral view [coll. FH, No. 0018-10]; 333. ♂ apex of abdomen in dorsal view [coll. FH, No. 0018-10]; 334. ♂ apex of abdomen in ventral view [coll. FH, No. 0018-10]; 335. Right mesocoxa of ♀ from Peninsular Malaysia, showing the usual pale pink colouration of the inner surface [coll. FH, No. 0018-27]; 336. Right mesocoxa of ♀ from Phuket in S-Thailand, showing the black colouration of the inner surface typical for specimens from this locality [coll. FH, No. 0018-#].


HENNEMANN ET AL.

FIGURES 337–340. Heteropteryx dilatata (Parkinson, 1798) 337. ♀ head, pro- and mesothorax in dorsal view [coll. FH, No. 0018-#]; 338. ♂ head, pro- and mesothorax in dorsal view [coll. FH, No. 0018-5]; 339. ♀ head, pro- and mesothorax in lateral view [coll. FH, No. 0018-#]; 340. ♂ head, pro- and mesothorax in lateral view [coll. FH, No. 0018-5].



185

FIGURE 341. Map of Southeast Asia showing the prinicipal islands and the Sundaland and Wallacea biogeographical subregions.


HENNEMANN ET AL.

FIGURE 342. Map showing the distributions of the three subfamilies of Heteropterygidae.



187

FIGURE 343. Distribution of Dataminae Rehn & Rehn, 1939. Arrows indicate the hypothetical tracklines.


HENNEMANN ET AL.

FIGURE 344. Distribution of Obriminae Brunner v. Wattenwyl, 1893. Arrows indicate the hypothetical tracklines.



189

FIGURE 345. Distribution of Heteropteryginae Kirby, 1896. Arrows indicate the hypothetical tracklines.


HENNEMANN ET AL.

FIGURES 346–349. Defensive reactions of species of Heteropteryginae 346. Haaniella dehaanii (Westwood, 1859) ♀; 347. Heteropteryx dilatata (Parkinson, 1798) ♂; 348. Haaniella echinata (Redtenbacher, 1906) ♀; 349. Haaniella echinata (Redtenbacher, 1906) ♂.



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FIGURES 350–354. Gynandromorphism in Heteropteryx dilatata (Parkinson, 1798) 350. Predominantly male mosaic gynandromorph with full-length wings (97.5 mm): Captive reared from Peninsular Malaysia, Perak, Tapah Hills [coll. FH, No. 0018-27]; 351. Apex of abdomen of above specimen in dorsal view. Arrow indicates a single green posteromedian spine on abdominal tergum VI; 352. Apex of abdomen of above specimen in ventral view; 353. Live mosaic gynandromorph (Copyright F. Seow-Choen); 354. Live mainly mmosaic gynandromorph (Copyright F. Seow-Choen). Ep = epiproct, ThP = thorn pads, Vo = vomer, Po = poculum.


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FIGURE 355. Phenogram showing relationships within Heteropteryginae. The preliminary tree does not restrict to using autapomorphies or synapomorphies but also uses key-features that distinguish the genera and species-groups of Haaniella Kirby, 1904 defined herein, hence is no phylogenetic cladogram. (*) not true for all species contained.



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PLATE 1. Metafemora in ventral view 356. Heteropteryx dilatata (Parkinson, 1798) ♀; 357. Heteropteryx dilatata (Parkinson, 1798) ♂; 358. Haaniella glaber (Redtenbacher, 1906) ♀; 359. Haaniella gintingi n. sp. ♀, PT; 360. Haaniella erringtoniae (Redtenbacher, 1906) ♀; 361. Haaniella aculeata n. sp. ♂, HT; 362. Haaniella dehaanii (Westwood, 1859) ♀; 363. Haaniella gorochovi n. sp. ♀, PT; 364. Haaniella saussurei Kirby, 1904 ♀; 365. Haaniella scabra (Redtenbacher, 1906) ♀.


HENNEMANN ET AL.

PLATE 2. Right metatibiae of Haaniella (muelleri species-group) and Heteropteryx in interior view 366. Haaniella aculeata n. sp. ♂, HT; 367. Haaniella macroptera n. sp. ♀, PT; 368. Haaniella macroptera n. sp. ♂, PT; 369. Haaniella erringtoniae (Redtenbacher, 1906) ♀; 370. Haaniella erringtoniae (Redtenbacher, 1906) ♂; 371. Haaniella rosenbergii (Kaup, 1871) ♀; 372. Haaniella rosenbergii (Kaup, 1871) ♂; 373. Haaniella gintingi n. sp. ♀, PT; 374. Haaniella gintingi n. sp. ♂, PT; 375. Haaniella glaber (Redtenbacher, 1906) ♀; 376. Haaniella glaber (Redtenbacher, 1906) ♂; 377. Heteropteryx dilatata (Parkinson, 1798) ♀; 378. Heteropteryx dilatata (Parkinson, 1798). ♂.



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PLATE 3. Right metatibiae of Haaniella (echinata species-group & grayii species-group) in interior view 379. Haaniella echinata (Redtenbacher, 1906) ♀; 380. Haaniella echinata (Redtenbacher, 1906) ♂; 381. Haaniella scabra (Redtenbacher, 1906) ♀; 382. Haaniella scabra (Redtenbacher, 1906) ♂; 383. Haaniella saussurei Kirby, 1904 ♀; 384. Haaniella saussurei Kirby, 1904 ♂; 385. Haaniella dehaanii (Westwood, 1859) ♀; 386. Haaniella dehaanii (Westwood, 1859) ♂; 387. Haaniella grayii (Westwood, 1859) ♀; 388. Haaniella grayii (Westwood, 1859) ♂.


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PLATE 4. Metacoxae of Haaniella spp. showing the distinctive colouration of the coxal trochanters and membranes between the basal abdominal sternites 389. Haaniella dehaanii (Westwood, 1859) ♀; 390. Haaniella gorochovi n. sp. ♀, PT; 391. Haaniella grayii (Westwood, 1859) ♀; 392. Haaniella echinata (Redtenbacher, 1906) ♂; 393. Haaniella saussurei Kirby, 1904 ♀; 394. Haaniella saussurei Kirby, 1904 ♂; 395. Haaniella scabra (Redtenbacher, 1906) ♀; 396. Haaniella scabra (Redtenbacher, 1906) ♂.



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PLATE 5. Ventral view of abdominal tergum X of ♂♂ showing the vomer 397. Haaniella erringtoniae (Redtenbacher, 1906); 398. Haaniella rosenbergii (Kaup, 1871); 399. Haaniella gintingi n. sp., PT; 400. Haaniella mecheli (Redtenbacher, 1906) HT; 401. Haaniella macroptera n. sp., PT; 402. Haaniella glaber (Redtenbacher, 1906); 403. Haaniella aculeata n. sp., HT; 404. Heteropteryx dilatata (Parkinson, 1798); 405. Haaniella gorochovi n. sp., HT.


HENNEMANN ET AL.

PLATE 6. Ventral view of abdominal tergum X of ♂♂ showing the vomer 406. Haaniella dehaanii (Westwood, 1859); 407. Haaniella grayii (Westwood, 1859); 408. Haaniella scabra (Redtenbacher, 1906); 409. Haaniella echinata (Redtenbacher, 1906) from Brunei, Badas; 410. Haaniella echinata (Redtenbacher, 1906) from Sabah, Mount Kinabalu, Poring Hot Springs 480 m; 411. Haaniella saussurei Kirby, 1904.



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PLATE 7. Eggs and internal micropylar plates of Haaniella (muelleri species-group and grayii species-group) 412. Haaniella erringtoniae (Redtenbacher, 1906), dorsal view; 413. Haaniella erringtoniae (Redtenbacher, 1906), lateral view; 414. Haaniella macroptera n. sp., dorsal view (PT); 415. Haaniella macroptera n. sp., lateral view (PT); 416. Haaniella gintingi n. sp., dorsal view (PT); 417. Haaniella gintingi n. sp., lateral view (PT); 418. Haaniella rosenbergii (Kaup, 1871), dorsal view; 419. Haaniella rosenbergii (Kaup, 1871), lateral view; 420. Haaniella gorochovi n. sp. , dorsal view (PT); 421. Haaniella gorochovi n. sp. , lateral view (PT); 422. Haaniella parva Günther, 1944, dorsal view; 423. Haaniella parva Günther, 1944, lateral view; 424. Haaniella dehaanii (Westwood, 1859), dorsal view; 425. Haaniella dehaanii (Westwood, 1859), lateral view; 426. Haaniella grayii (Westwood, 1859), dorsal view; 427. Haaniella grayii (Westwood, 1859), lateral view; 428. Internal micropylar plate of Haaniella erringtoniae (Redtenbacher, 1906), enlarged; 429. Internal micropylar plate of Haaniella grayii (Westwood, 1859), enlarged; 430. Internal micropylar plate of Haaniella dehaanii (Westwood, 1859), enlarged.


HENNEMANN ET AL.

PLATE 8. Eggs and internal micropylar plates of Haaniella (echinata species-group) and Heteropteryx 431. Haaniella saussurei Kirby, 1904, dorsal view; 432. Haaniella saussurei Kirby, 1904, lateral view; 433. Haaniella scabra (Redtenbacher, 1906), dorsal view (from Sabah, Mount Kinabalu N.P., near headquarters, Silau-Silau trail 1550 m); 434. Haaniella scabra (Redtenbacher, 1906), lateral view (from Sabah, Mount Kinabalu N.P., near headquarters, Silau-Silau trail 1550 m); 435. Haaniella scabra (Redtenbacher, 1906), dorsal view (Captive reared from Sabah Mount Kinabalu); 436. Haaniella scabra (Redtenbacher, 1906), lateral view (Captive reared from Sabah Mount Kinabalu); 437. Haaniella echinata (Redtenbacher, 1906), dorsal view (from Sabah, Mount Kinabalu, Poring Hot Springs 480 m); 438. Haaniella echinata (Redtenbacher, 1906), lateral view (from Sabah, Mount Kinabalu, Poring Hot Springs 480 m); 439. Haaniella echinata (Redtenbacher, 1906), dorsal view (from Brunei, Kuala Belalong, Badas); 440. Haaniella echinata (Redtenbacher, 1906), lateral view (from Brunei, Kuala Belalong, Badas); 441. Heteropteryx dilatata (Parkinson, 1789), dorsal view (from Peninsular Malaysia, Perak, Tapah Hills); 442. Heteropteryx dilatata (Parkinson, 1789), lateral view (from Peninsular Malaysia, Perak, Tapah Hills); 443. Heteropteryx dilatata (Parkinson, 1789), dorsal view (from S-Thailand, Phuket); 444. Heteropteryx dilatata (Parkinson, 1789), lateral view (from S-Thailand, Phuket); 445. Internal micropylar plate of Haaniella echinata (Redtenbacher, 1906), enlarged (from Brunei, Kuala Belalong, Badas); 446. Internal micropylar plate of Haaniella echinata (Redtenbacher, 1906), enlarged (from Sabah, Mount Kinabalu, Poring Hot Springs 480 m); 447. Internal micropylar plate of Heteropteryx dilatata (Parkinson, 1798), enlarged (from Peninsular Malaysia, Perak, Tapah Hills).



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PLATE 9. Live insects 448. Haaniella kerinci n. sp. ♂: W-Sumatra, Prov. Jambi, Kerinci Seblat N.P., Mount Tujuh, I.2014 [@ Jeremy Holden]; 449. Haaniella kerinci n. sp. ♂: W-Sumatra, Prov. Jambi, Kerinci Seblat N.P., Mount Tujuh, I.2014 [@ Jeremy Holden]; 450. Haaniella erringtoniae (Redtenbacher, 1906) ♂ and ♀: Peninsular Malaysia, Pahang, Genting Highlands 1500-1600 m, VIII.2009 [photo by Francis Seow-Choen].


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PLATE 10. Live insects 451. Haaniella erringtoniae (Redtenbacher, 1906) ♀ with a dark longitudinal median stripe on thorax and unusual pale markimgs on thorax and tegmina: Peninsular Malaysia, Pahang, Genting Highlands 1500-1600 m, VIII.2009 [photo by Francis Seow-Choen]; 452. Haaniella erringtoniae (Redtenbacher, 1906) ♀: Peninsular Malaysia, Pahang, Genting Highlands 15001600 m, VIII.2009 [photo by Francis Seow-Choen].



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PLATE 11. Live insects 453. Haaniella macroptera n. sp. mating couple (Paratypes): Singapore, Nee Soon swamp forest, V.2013 [photo by Francis Seow-Choen]; 454. Haaniella macroptera n. sp. ♂ (Paratype) with an unusual white stripe along anterior margin of tegmina: Singapore, Nee Soon swamp forest, V.2013 [photo by Francis Seow-Choen]; 455. Haaniella macroptera n. sp. ♀ (Paratypes) with a bold white basal marking on tegmina: Singapore, Nee Soon swamp forest, V.2013 [photo by Francis Seow-Choen].


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PLATE 12. Live insects 456. Haaniella rosenbergii (Kaup, 1871) ♀: NW-Sumatra, Prov. Aceh, Subulussalam, IV.2013 [photo by Francis SeowChoen]; 457. Haaniella rosenbergii (Kaup, 1871) mating couple: NW-Sumatra, Prov. Aceh, Subulussalam, IV.2013 [photo by Francis Seow-Choen].



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PLATE 13. Live insects 458. Haaniella gintingi n. sp. couple: NW-Sumatra, Prov. Aceh, alongside road leading from Berastagi to Danau Toba, 12001400 m, IV.2013 [photo by Francis Seow-Choen]; 459. Haaniella gintingi n. sp. ♀ with unusual colouration: NW-Sumatra, Prov. Aceh, alongside road leading from Berastagi to Danau Toba, 1200-1400 m, IV.2013 [photo by Francis Seow-Choen].


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PLATE 14. Live insects 460. Haaniella dehaanii (Westwood, 1859) mating couple: NW-Sarawak, Gunung Gading N.P. [photo by Francis SeowChoen]; 461. Haaniella dehaanii (Westwood, 1859) ♀ with a n unusually small pale central marking on mesonotum: NWSarawak, Gunung Gading N.P. [photo by Francis Seow-Choen].



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PLATE 15. Live insects 462. Haaniella dehaanii (Westwood, 1859) ♂: NW-Sarawak, Gunung Gading N.P. [photo by Francis Seow-Choen]; 463. Haaniella dehaanii (Westwood, 1859) ♂: Captive reared from NW-Sarawak, Mount Serapi, V.2010 [photo by Frank Hennemann].


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PLATE 16. Live insects 464. Haaniella gorochovi n. sp. ♀, PT: S-Vietnam, Prov. Lâm Dông, Bidoup-Nùi Bá N.P. [@ Joachim Bresseel]; 465. Haaniella gorochovi n. sp. ♀, PT: S-Vietnam, Prov. Lâm Dông, Bidoup-Nùi Bá N.P. [@ Joachim Bresseel]; 466. Haaniella grayii (Westwood, 1859) plain reddish brown ♀: NW-Sarawak, Gunung Gading N.P., 2008 [@ Paul Bertner].



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PLATE 17. Live insects 467. Haaniella grayii (Westwood, 1859) ♀ with pretty colouration: NW-Sarawak, Gunung Gading N.P. [photo by Francis Seow-Choen]; 468. Haaniella grayii (Westwood, 1859) prettily mottled ♀ posessing remarkable camouflage on a lichenose trunk: NW-Sarawak, Gunung Gading N.P. [photo by Francis Seow-Choen]; 469. Haaniella grayii (Westwood, 1859) ♂: NWSarawak, Gunung Gading N.P. [photo by Francis Seow-Choen].


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PLATE 18. Live insects 470. Haaniella parva Günther, 1944 mating couple: NW-Sumatra, Prov. Sumatera Utara, Berastagi 1220 m, IV.2013 [photo by Francis Seow-Choen]; 471. Haaniella parva Günther, 1944 ♂: NW-Sumatra, Prov. Sumatera Utara, Berastagi 1220 m, IV.2013 [photo by Francis Seow-Choen].



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PLATE 19. Live insects 472. Haaniella echinata (Redtenbacher, 1906) mating couple: Sarawak, Lambir Hills National Park [photo by Francis SeowChoen]; 473. Haaniella echinata (Redtenbacher, 1906) ♂: Sabah, Sepilok Orang Utan Rehabilitation Centre [photo by Francis Seow-Choen]; 474. Haaniella echinata (Redtenbacher, 1906) ♀ with white diagonal stripe on tegmina rarely seen in ♀♀: Sarawak, Mulu N.P., 2009 [@ Paul Bertner].


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PLATE 20. Live insects 475. Haaniella echinata (Redtenbacher, 1906) prettily mottled ♀: Sabah, Mount Kinabalu, Poring Hot Springs ca. 500 m [photo by Francis Seow-Choen]; 476. Haaniella echinata (Redtenbacher, 1906) couple: Sarawak, Mulu National Park [photo by Francis Seow-Choen].



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PLATE 21. Live insects 477. Haaniella saussurei Kirby, 1904 mating couple: N-Sarawak, Bako National Park [photo by Francis Seow-Choen]; 478. Haaniella saussurei Kirby, 1904 ♀: N-Sarawak, Bako National Park [photo by Francis Seow-Choen]; 479. Haaniella saussurei Kirby, 1904 mating couple: N-Sarawak, Batang Ai N.P. [photo by Francis Seow-Choen].


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PLATE 22. Live insects 480. Haaniella saussurei Kirby, 1904 ♀: N-Sarawak, Bako National Park [photo by Francis Seow-Choen]; 481. Haaniella saussurei Kirby, 1904 ♀: N-Sarawak, Batang Ai N.P. [photo by Francisis Seow-Choen].



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PLATE 23. Live insects 482. Haaniella scabra (Redtenbacher, 1906) mating couple: N-Sabah, Mount Kinabalu National Park, ca. 1500 m [photo by Francis Seow-Choen]; 483. Haaniella scabra (Redtenbacher, 1906) ♀ with a dark longitudinal stripe along dorsal body surface: Sabah, Crocker Range National Park [photo by Francis Seow-Choen]; 484. Haaniella scabra (Redtenbacher, 1906) ♀: Captive reared from Sabah, Mount Kinabalu, VIII.2014 [photo by Frank Hennemann].


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PLATE 24. Live insects 485. Heteropteryx dilatata (Parkinson, 1798) ♀: Captive reared from Peninsular Malaysia, Perak, Tapah Hills, III.2010 [photo by Frank Hennemann]; 486. Heteropteryx dilatata (Parkinson, 1798) mating couple: West Malaysia, Perak, Kijang Falls [photo by Francis Seow-Choen]; 487. Heteropteryx dilatata (Parkinson, 1798) ♂: Captive reared from Peninsular Malaysia, Perak, Tapah Hills, VIII.2012 [photo by Frank Hennemann]; 488. Heteropteryx dilatata (Parkinson, 1798) penultimate instar ♂ nymph with unusual colouration: Captive reared from Peninsular Malaysia, Perak, Tapah Hills, VIII.2012 [photo by Frank Hennemann].



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PLATE 25. Maps showing the records of Haaniella-species.


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PLATE 26. Maps showing the records of Haaniella-species and Heteropteryx dilatata (Parkinson, 1798).



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