Tepakiphasma ngatikuri, a new genus and species of stick insect (Phasmatodea) from the Far North of New Zealand

Tepakiphasma ngatikuri, a new genus and species of stick insect (Phasmatodea) from the Far North of New Zealand THOMAS R. BUCKLEY 1, SVEN BRADLER 2 1 Landcare Research, Private Bag 92170, Auckland, New Zealand. E-mail: buckleyt@landcareresearch.co.nz 2 Johann-Friedrich-Blumenbach-Institut für Zoologie und Anthropologie, Georg-August-Universität Göttingen, Berliner Strasse 28, 37073 Göttingen, Germany. ABSTRACT We describe a new genus and species of stick insect from Northland, New Zealand, Tepakiphasma ngatikuri, gen. nov., sp. nov. We have classified this genus as a member of Phasmatidae, Phasmatinae, Acanthoxylini, and due to the presence of certain key synapomorphies it is phylogenetically placed within the Australasian clade Lanceocercata. A number of character states differentiate Tepakiphasma from other New Zealand Acanthoxylini genera including the number and arrangement of teeth on the claspers and a perforate egg capitulum or capitular cone. Like many New Zealand phasmatodeans the known host plants of T. ngatikuri include species of Myrtaceae. This genus appears to have an extremely limited geographic distribution and is known from only two specimens collected in the Te Paki / North Cape area at the northernmost tip of mainland New Zealand. This discovery further emphasises the importance of the Te Paki / North Cape area in New Zealand biodiversity. The phasmatodean fauna of New Zealand now contains 10 genera and 23 described species. KEY WORDS Acanthoxylini, Lanceocercata, Phasmatidae, Phasmatinae, Te Paki. INTRODUCTION The Te Paki/North Cape area at the very northern tip of New Zealand is of high biodiversity value because of the large numbers of endemic plant (e.g., Davidson et al. 1969; Druce et al. 1979; de Lange et al. 2003; von Konrat & Braggins 2005), vertebrate (e.g., Chapple et al. 2008) and invertebrate species (e.g., Gardner 1967; Goulstone et al. 1993; Larochelle & Larivière 2005; Winterbourn 2009). Furthermore, due to the extremely degraded habitat in this area (Gardner & Bartlett 1980; Lux et al. 2009) many of these endemic species are threatened with extinction (Lux et al. 2009). Many vertebrate species have become extinct due to human induced environmental changes and information on the pre-human invertebrate fauna is lacking, with the exception of land molluscs (Brook 1999). Because the Te Paki/North Cape area is a centre of endemism, two collecting trips were undertaken to survey the phasmatodean fauna. In December 2008 we collected a pair of late instar nymphs in Radar Bush, Te Paki Recreation Reserve. These specimens were taken back to the laboratory and reared through to adulthood. Analysis of morphological characters indicated the specimens belong to a new species and genus. This discovery represents the second new phasmatodean species from the Te Paki/North Cape area, the first being an undescribed species of Clitarchus Stål discussed by Buckley et al. (2010a). MATERIAL AND METHODS Specimens and analysis of characters The specimens studied here are lodged in the New Zealand Arthropod Collection (NZAC), Auckland, New Zealand. Higher level nomenclature follows Günther (1953) instead of more recent classifications for reasons outlined in Klug and Bradler (2006). Measurements were made using electronic digital callipers. Specimens were studied and illustrations were prepared using LEICA MZ75, ZEISS Stemi SV6 and ZEISS Stemi SV11 stereomicroscopes at different magnifications and with a drawing tube (camera lucida). Terminology of adult morphological characters follows Beier (1968) and Littig (1942) and that of egg characters Clark Sellick (1997, 1998). The holotype and paratype of this newly described species are stored in the NZAC in 70% and 100% ethanol respectively, as this method of preservation facilitates inspection of the terminalia better than in pinned specimens and ensures the preservation of DNA in 100% ethanol. DNA sequence data from the paratype will be published at a later date. Because the genus is known from only one species and two specimens the generic description is essentially identical to the species description. For this reason we give a diagnosis for the genus, a full species description and a discussion of characters justifying erection of a new genus. 118

New genus of New Zealand stick insect Taxonomy Phasmatodea: Phasmatidae: Phasmatinae: Acanthoxylini Tepakiphasma gen. nov. Type species: Tepakiphasma ngatikuri sp. nov., by present designation Diagnosis : Antennae extend well beyond fore femora. Small, lateral foliaceous lobes at posterior end of abdominal tergum VII. Each clasper with two teeth, one larger tooth at base of clasper and one at apex of clasper, opposing inner teeth touching in repose. : Antennae extend well beyond fore femora. Cerci only slightly extending beyond paraprocts. Flanges present on dorsal carina of 1 st tarsus. Egg: with perforated capitulum or capitular cone. Etymology The name Tepakiphasma is formed from Te Paki, the only region where the genus has so far been found, and the stem word for Phasmatodea. Tepakiphasma ngatikuri sp. nov. Type material: Holotype (NZAC03005884), NEW ZEALAND: ND: Kauri Bush, Te Paki. 34º 28 S, 172 º 45 E, beating Metrosideros perforata, 9 December 2008, T.R. Buckley, D. Seldon, R.A.B. Leschen (NZAC). Paratype (NZAC03005995), NEW ZEALAND: ND: Kauri Bush, Te Paki. 34º 28 S, 172 º 45 E, ex Metrosideros perforata at night, 9 December 2008, T.R. Buckley, D. Seldon, R.A.B. Leschen (NZAC). Other material examined: 102 eggs from paratype female laid in captivity. Description General: Medium sized stick insect, gracile and apterous. Body colour brown. Spines absent and tubercles sparse on body. Body colour mottled from light grey to dark brown. Head: Prognathous, dorsoventrally flattened and longer than wide, unarmed and darker on ventral surface. Gula present, shorter than wide. Ocelli absent. Eyes prominent, hemispherical, grey and black. Antenna filiform with 16 flagellomeres, lightly setose, reaching well beyond end of fore femora. Each flagellomere darker near apex. Scapus oval shaped and dorsoventrally flattened, especially towards proximal end. Pedicellus cylindrical and less than half the length of scapus. 1 st flagellomere more than twice as long as pedicellus and longer than 2 nd flagellomere. Slight basal swelling on ventral surface of 7 th flagellomere. Labial palps with three articles, slightly dorsoventrally flattened. Maxillary palps with five articles, terminal article longest. Galea with medially directed trichome area on apex, galealobulus not elongated. Stipes with three apical teeth. Thorax: Prothorax slightly shorter than head, longer than wide, unarmed with dark median line running length of pronotum. Openings of pair of defensive glands slit-like, indistinct, located laterally at the front margin of the pronotum. Prothoracic basisternite short, of equal length as furcasternite, both sternites without spines and tubercles. Mesothorax longer than wide and more than four times longer than prothorax. Mesonotum with several small tubercles and darker patches at posterior margin. Mesothoracic episternum elongated, significantly longer than epimeron, with small, rounded tubercles. Episternal suture present. Mesothoracic basisternite unarmed with several very small tubercles. Mesothoracic furcasternite with small tubercle. Metathorax shorter than mesothorax, tergum with darker patches at posterior margin. Episternum elongated, darkly coloured in anterior region with row of small tubercles, with episternal suture. Epimeron short with single indistinct tubercle. Metanotum and metasternum unarmed with scarce small tubercles. Abdomen: Ten segmented abdomen unarmed. First abdominal segment (median segment or segmentum medianum) fused to metathorax, boundary indistinct, tergum of first abdominal segment significantly shorter than metanotum and subsequent terga of abdomen. Abdominal terga II VIII longer than wide, tergum X less the half length of IX. Tergum VII with small foliaceous lobes at posterior end. Darker patches at lateral posterior margins of abdominal terga and lighter patches at central posterior margin. Pair of small tubercles at posterior margins of terga II to VII. Spines and tubercles absent on sterna. Tergum VIII broadens posteriorly and tergum IX narrows posteriorly, lateral margins approach each other on ventral side. Margins of basal part of tergum X also strongly bent downwards, touching on ventral side, not fused. Sternum IX transversally divided, posterior part (poculum, subgenital plate) free, not connected to dorsal part of corresponding segment, slightly smaller than anterior part. Tip of subgenital plate pointed with notch and reaching less than half way along 9 th tergum. Vomer absent, remnants visible as sclerotized area between paraprocts. Paraprocts and epiproct largely concealed beneath lateral margins of tergum X. Claspers formed by tergum X, short, with pair of opposing teeth at apex and massive enlarged pair at base touching in repose. Cerci dorsoventrally flattened, elongated, slightly attenuated at apex and projecting beyond 10 th abdominal segment. Legs: Femora and tibiae with mid carina on ventral surface, appearing pentagonal in cross section with setae running along carinae. Setae increasing in density from femora to tibiae to tarsi and especially numerous and long on pretarsus and ungues. Fore coxae with several tubercles. Trochanters small, fused to femora. Fore femora compressed basally and 119

curved to accommodate head during catalepsy, with four small spines along posterior-ventral carina. Pair of apical spines present. Fore tibiae approximately equal in length to fore femora and unarmed except for apical teeth. Tarsomeres all shorter than femora and tibiae, and in decreasing order of 1 st, 5 th, 2 nd, 3 rd, and 4 th. Mid coxae with several small tubercles. Trochanters as in fore legs. Mid femora unarmed except for two small spines on latero-ventral carinae near apex. Pair of apical spines present. Mid tibiae unarmed except for pair of apical spines. Tarsi as for fore leg. Hind coxae unarmed. Hind trochanters small, fused to femora. Hind femora unarmed except for apical spines. Hind tibiae unarmed except for pair of apical spines. Tarsi as for fore leg. General: Medium sized stick insect, moderately robust and apterous. Body colour brown. Spines absent and tubercles sparse on body. Body colour mottled from light grey to dark brown. Head: Prognathous, dorsoventrally flattened and longer than wide. Ocelli absent. Eyes prominent, hemispherical, darker than rest of head. Short gula present. Interocular and anterior area of ventral surface darker. Gena also darker. Antenna filiform with 20 flagellomeres covered in setae, reaching beyond end of fore femora. Scapus oval shaped and dorsoventrally flattened, especially towards proximal end. Pedicellus cylindrical, slightly dorsoventrally flattened and less than half the length of scapus. 1 st flagellomere longer than 2 nd. Light coloured basal swelling on dorsal surface of 9 th flagellomere. Terminal flagellomere longest. Labial palps with three articles slightly dorsoventrally flattened. Galea slender, with trichome area on apex, galealobulus present, not elongated. Stipes with three teeth: one apical tooth and two subapical teeth. Maxillary palps with five articles, terminal article longest. Labrum, glossa and paraglossa light red to pink. Thorax: Prothorax slightly shorter than head, longer than wide. Pronotum rugose with several small tubercles. Small darker patch at median of anterior and posterior margins. Openings of pair of defensive glands located laterally at the front margin of the pronotum, slit-like, indistinct. Episternum and epimeron with dark or body coloured tubercles. Prothoracic basisternite short, of equal length as furcasternite. Several small tubercles on basisternite and furcasternite. Episternum and epimeron with dark or body coloured tubercles. Prothoracic basisternite short, of equal length as furcasternite. Several small tubercles on basisternite and furcasternite. Mesothorax longer than wide and over four times longer than prothorax. Mesonotum with approximately six irregular rows of spines or tubercles running from anterior to posterior margin with six dark larger tubercles in anterior half. Episternum largely elongated, epimeron short. Tubercles present on episternum and epimeron. Episternal suture present. Mesothoracic basisternite with many small tubercles usually lightly coloured with 120 dark margins. Several larger tubercles, lightly coloured with larger dark margins. Metathorax + median segment (1 st abdominal segment) marginally shorter than mesothorax. Metanotum with many small tubercles usually lightly coloured with dark margins. Several larger tubercles, lightly coloured with larger dark margins. Posterior and anterior region of metanotum with more extensive lighter areas. Episternum and epimeron as in mesothorax. Tubercles present on metathoracic basisternite and furcasternite. Abdomen: Ten segmented abdomen with tergal spines absent. First abdominal segment (segmentum medianum) fused to metathorax, shorter than metanotum, boundary indistinct. Terga IV and V light grey colour, other terga mottled brown / grey with smaller light grey patches. Spines absent or reduced to tubercles on sterna. Row of tubercles running perpendicular to body near posterior margin of abdominal terga II and III. Pair of larger tubercles near posterior margin of abdominal terga II, III and IV. Terga V to VIII with three larger tubercles near posterior margin. Tergum II marginally shorter than III, increasing in size from III to VI. Terga VII and VIII of equal length, and IX shorter than VIII, and X being the shortest tergum. Tubercles on abdominal sterna, larger on sterna VI and VII. Sterna IV and V with more light grey colouration, other sterna darker brown / grey with smaller light grey patches. Operculum (abdominal sternum VIII) boat shaped with carina along ventral surface, keeled. Single lateral carina along each side of abdominal operculum. Tip of operculum pointed and reaching half way along tergum X. Terminal margin of tergum X truncated, with epiproct protruding slightly beyond end. Paraprocts triangular and only slightly longer than wide and very dark. Gonapophyses largely covered by operculum. Gonapophyses VIII (GpVIII) reaching to end of gonapophysis IX. Gp VIII slender, pointed at apex. Gp IX leaf like, stout. No olistheter (tongue-and-groove-mechanism) formed by valves. Gonoplac finger-like, with setae at tip, extending well beyond other valves to posterior margin of tergum IX. Gonangulum prominent, triangular. Cerci flattened and foliaceous, slightly longer than paraprocts and protruding just beyond tergum X. Setae on surface of cerci, more dense around margins. Dorsal and ventral sides of each cercus with darker, more setose patch in interior, anterior half. Outside margins of cerci curving outwards towards tip with inside margins straight. Legs: Femora and tibiae as in, appearing pentagonal in cross section with setae running along carina. Setae increasing in density from femora to tibiae to tarsi and especially numerous and long on pretarsi and ungues. Ungues smooth, not pectinate. Fore coxae with several large tubercles. Trochanters in all legs small, fused to femora. Fore femora compressed basally and curved to accommodate head in resting position, and bright pink coloured at base. Ventro-posterior carina with row of 7 teeth, with apical tooth the largest of all fore leg teeth,

New genus of New Zealand stick insect Figures 1-2. Photographs of male (holotype) and female (paratype) of Tepakiphasma ngatikuri gen. nov., sp. nov. in copula. Fig. 1, male (holotype) and female (paratype) in copula; Fig. 2, male (holotype) clasping female (paratype) at base of operculum (abdominal sternum 8). 121 80954 NZ Entomologist Book_artic121 121 8/3/10 12:28:47 PM

pair of dorsal carinae almost fused at anterior end, with gentle undulating profile. Pair of apical spines always present. Fore tibiae approximately equal in length to fore femora and unarmed except for dorsal carinae with gentle undulating profile and apical teeth. Tarsomeres all shorter than femora and tibiae, and in decreasing order of 1 st, 5 th, 2 nd, 3 rd, and 4 th. Mid coxae with tubercles. Mid femora with 2-3 small teeth along dorsal carinae, four teeth along latero-ventral carinae including larger tooth on each carina near apex. Pair of apical spines always present. Mid tibiae with large flange on dorsoposterior carina near proximal end. Pair of apical spines always present. Large dorsal flange near distal end of 1 st tarsomere. Other tarsi as for fore leg. Hind coxae with several tubercles. Hind femora with four teeth along the mid-ventral carinae, pair of small teeth near proximal end of latero-ventral carinae and larger pairs near distal end of carina, 2-3 teeth along each dorsal carina. Pair of apical spines always present. Hind tibiae with pair of apical teeth and large flange on posterior dorsal carina near proximal end, or with single dorsal tooth, apical spines always present. Large dorsal flange near distal end of 1 st tarsomere. Other tarsi as for fore leg. Egg: Slightly elongated along anterior-posterior axis, laterally flattened, opercular angle close to zero. Capsule smooth with scattered small pits, darkly coloured. Keel distinct, running from opercular collar to micropylar plate. Micropylar plate darkly coloured, elliptical and mostly in posterior half. Micropylar cup lightly coloured. Opercular collar smooth. Operculum with capitular cone, flat topped with apical projection or large and conical. Anterior surface of capitulum perforated. Notes on variation As mentioned in the description, the structure of the capitulum shows some variation. Some eggs have a conical capitulum and others have a flat capitulum with a central apical projection resembling a stalk (Figs. 9, 11). Etymology The specific name was selected by the Ngāti Kurī people who are kaitiaki (guardians) of the Te Paki/North Cape area. Type locality and distribution The species is known only from the type locality, near the bottom of the Kauri Bush Track in Radar Bush, Te Paki Recreation Reserve. This locality is known either as Radar Bush or Kauri Bush. The holotype and paratype were collected from two Metrosideros perforata (Myrtaceae) vines, both growing on trees, within a few metres of each other. 122 DISCUSSION Taxonomic status and classification The antennae of Tepakiphasma are longer than the fore femora and this character state is shared with the New Zealand genera Acanthoxyla Uvarov, Clitarchus, and Pseudoclitarchus Salmon, which are placed in Phasmatidae, Phasmatinae, Acanthoxylini. Phylogenetic studies show both the Phasmatidae and the Phasmatinae to be non-monophyletic (Whiting et al. 2003; Bradler 2009; Buckley et al. 2009, 2010b) and so this taxonomic placement is provisional only. Tepakiphasma is a member of the clade Lanceocercata (Bradler 2001) due to the presence of key apomorphies including leaf-like cerci, absence of a vomer, claspers orientated medially against each other and the male clasping the female at the base of the operculum (abdominal sternum 8) (Fig. 2). We have erected a new genus, because Tepakiphasma displays a number of character states not possessed by any other genus of New Zealand stick insect. The most striking character state difference between Tepakiphasma and the remaining New Zealand genera is the morphology of the egg. Clark-Sellick (1997) described the opercular extensions of [ ] most Acanthoxylini as simply raised opercula that cannot be termed capitula. However, Clark-Sellick (1997) also described the structure in question as a closed conical capitulum in Clitarchus and Pseudoclitarchus and as a conical capitular structure in Argosarchus. We follow the latter interpretation, because a perforated operculum would be equivalent to an open capsule and consequently a non-viable egg. The perforated capitulum or capitular cone (Fig. 14) is an apomorphy with regard to all other New Zealand phasmatodean taxa. However, perforated capitula are known from stick insects from other subfamilies in other geographic areas (e.g., Clark Sellick 1997; Baker & Chandrapatya 2001). In Acanthoxyla, Clitarchus and Pseudoclitarchus, the female cerci are all much longer, significantly extending posteriorly beyond the paraprocts (Salmon 1948, 1991; Jewell & Brock 2002), whereas in Tepakiphasma the cerci are only slightly longer than the paraprocts. The relative size of the cerci and paraprocts are similar between Argosarchus and Tepakiphasma; however the cerci of Tepakiphasma are much more rounded apically (Fig. 5, 9) than in Argosarchus. Acanthoxyla lacks males (Salmon 1991), and therefore we are unable to compare the male of Tepakiphasma with males of this genus. However, the number of teeth on the claspers differs between Clitarchus, Pseudoclitarchus, Argosarchus and Tepakiphasma. Clitarchus males have between three and five teeth on each clasper (Fig. 5; Salmon 1991; Buckley & Bradler unpublished data), Pseudoclitarchus has a single tooth per clasper (Salmon 1991) and Tepakiphasma has two teeth on

New genus of New Zealand stick insect each clasper (Figs. 3, 4). The claspers of Argosarchus are different from all remaining Acanthoxylini as they possess a cluster of long spine-like teeth at the apex (Salmon 1991). Therefore, the suite of putative autapomorphies displayed by Tepakiphasma and the lack of unambiguously synapomorphic characters with other New Zealand taxa are consistent with its generic status. Natural history and conservation The type locality consists of regenerating forest dominated by various broadleaf trees, Kunzea ericoides, Agathis australis, and the podocarps Dacrydium cupressinum, Halocarpus kirkii, Podocarpus totara and the tree Libocedrus plumosa (Gardner & Bartlett 1980). This area occupies approximately 20 hectares only and is of outstanding biodiversity value because it has the greatest plant diversity of all remaining forest fragments in the Te Paki/North Cape area (Gardner & Bartlett 1980). Furthermore, this forest fragment and the immediate surrounding area is the type locality for a number of beetle (Holloway 1961; Larochelle & Larivière 2005), mirid hemipteran (Eyles 2005), spider (Forster 1970), skink (Chapple et al. 2008), and plant species (von Konrat & Braggins 2005), and is the only known locality of several undescribed species of land snail (e.g., Goulstone et al. 1993). The new, and as yet undescribed, species of Clitarchus discussed by Buckley et al. (2010a) is widespread in the Te Paki/North Cape area including Radar Bush. Both specimens were found on Metrosideros perforata as final instar nymphs, and were observed to feed on this host shortly after collection. In the laboratory they were reared on Lophomyrtus bullata (Myrtaceae) through to adulthood and then kept alive on Metrosideros excelsa (Myrtaceae) for several months. Although T. ngatikuri can survive on at least three species of Myrtaceae, we have not yet collected it from Leptospermum scoparium or Kunzea ericoides (both Myrtaceae), both of which are abundant in the Te Paki area and are known host plants for many other New Zealand Phasmatodea (Salmon 1991; Buckley et al. 2010b). Despite searches at a number of other sites in the Te Paki/North Cape area, this species has not been located beyond Radar Bush. Potential threats to T. ngatikuri include predation by adventive rats, possums and vespulid wasps, all known predators of New Zealand Figures 3-9. Illustrations of the male and female terminalia of Tepakiphasma ngatikuri gen. nov., sp. nov. and male terminalia of Clitarchus hookeri. Fig. 3, male (holotype) of T. ngatikuri, terminal region, ventral view; Fig. 4, male of T. ngatikuri, abdominal tergum 10 (clasper), posterior view; Fig. 5, male of T. ngatikuri, terminal region, lateral view; Fig. 6, male of C. hookeri, terminal region, ventral view; Fig. 7, male of C. hookeri, abdominal tergum 10 (clasper), posterior view; Fig. 8, male of C. hookeri, terminal region, lateral view; Fig. 9, female (paratype) of T. ngatikuri, terminal region, lateral view; cer, cercus; ep, epiproct; gon, gonangulum; gpl, gonoplac; op, operculum; par, paraproct; pha, phallus; poc, poculum; s8, s9, abdominal sternum 8, 9; sti8, abdominal stigmum 8; t8 t10, abdominal tergum 8 10; tho, thorns resp. thorn pads. 123

Figures 10-14. Photographs of two eggs of Tepakiphasma ngatikuri gen. nov., sp. nov. laid by the paratype female. Fig. 10, lateral view of fi rst egg; Fig. 11, dorsal view of fi rst egg; Fig. 12, lateral view of second egg; Fig. 13, dorsal view of second egg; Fig. 14, anterior view of second egg showing the perforated capitulum resp. capitular cone. 124

Table 1. Body dimensions (mm) of holotype and paratype of Tepakiphasma ngatikuri gen. nov., sp. nov. New genus of New Zealand stick insect Male Female Body 76 104 Head 4 5 Antennae 30 25 Pronotum 3 4 Mesonotum 18 25 Metanotum incl. median segment 15 19 Abdomen without median segment 39.5 51 Fore femur 23.5 28 Mid femur 18.5 21 Hind femur 21 22 Fore tibia 27 28 Mid tibia 19 19 Hind tibia 23 20 stick insects (Cowan & Moeed 1987; Brockie 1992; Buckley, pers. obs.). A mammalian control programme is currently underway in the Radar Bush area, and the discovery of a vulnerable endemic stick insect species strongly suggests this programme should be continued long term. In recent years significant fires have occurred in the Te Paki area and destruction of the type locality also remains a threat (Lux et al. 2009). We recommend that surveys of the Te Paki/North Cape area be undertaken urgently to assess the distribution of T. ngatikuri and if necessary conservation management plans put into action. ACKNOWLEDGEMENTS We acknowledge the support of Ngāti Kurī and Te Aupōuri in granting access to the Te Paki/North Cape area, and in particular Ngāti Kurī for selecting the specific name. We thank Dave Seldon and Rich Leschen for helping collect the type specimens. Thanks to Robert Hoare, Rich Leschen and Tony Jewell for comments on the manuscript and many useful discussions on taxonomy. We also thank two anonymous reviewers for helpful comments on earlier versions of the manuscript. Birgit Rhode and Robert Hoare provided photographs. The Department of Conservation issued collecting permits and helped with field work and logistics, in particular Lester Bridson, Janeen Collings, and Patrick Whaley. This work was funded by the Foundation for Research, Science and Technology through the Defining New Zealand Land Biota OBI and an Ernst Mayr Travel Award from Harvard University (to TRB). REFERENCES Baker GT, Chandrapatya A. 2001. Morphology of the chorion of Diapheromera femorata (Say) (Phasmida: Heteronemiidae). Proceedings of the Entomological Society of Washington 103: 849-853. Beier M. 1968. Phasmida (Stab- oder Gespenstheuschrecken). In: Handbuch der Zoologie IV (2) 2/10 (eds, Helmcke, J.-G., Starck, D., Wermuth, H). Walter De Gruyter & Co, Berlin. Bradler S. 2001. The Australian stick insects, a monophyletic group within the Phasmatodea? Zoology 104, Suppl. III: 69. Bradler S. 2009. Phylogeny of the stick and leaf insects (Insecta: Phasmatodea). Species, Phylogeny & Evolution 2: 3-139. Brockie R. 1992. A Living New Zealand Forest. David Bateman, Glenfield, Auckland. Brook FJ. 1999. Stratigraphy, landsnail faunas, and palaeoenvironment history of coastal dunefields at Te Werahi, northernmost New Zealand. Journal of the Royal Society of New Zealand 29: 361-393. Buckley TR, Attanayake D, Bradler S. 2009. Extreme convergence in stick insect evolution: phylogenetic placement of the Lord Howe Island tree lobster. Proceedings of the Royal Society of London, B 276: 1055-1062. Buckley TR, Marske K, Attanayake D. 2010a. Phylogeography and ecological niche modelling of the New Zealand stick insect Clitarchus hookeri (White) support survival in multiple coastal refugia. Journal of Biogeography, in press. Buckley TR, Attanayake D, Nylander JAA, Bradler S. 2010b. The phylogenetic placement and biogeographical origins of the New Zealand stick 125

insects (Phasmatodea). Systematic Entomology, in press. Chapple DG, Patterson GB, Bell T, Daugherty CH. 2008. Taxonomic revision of the New Zealand copper skink (Cyclodina aenea: Squamata: Scincidae) species complex, with descriptions of two new species. Journal of Herpetology 42: 437-452. Clark Sellick JT 1997. Descriptive terminology of the phasmid egg capsule, with an extended key to the phasmid genera based on egg structure. Systematic Entomology 22: 97-122. Clark Sellick JT 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. Cowan PE, Moeed A. 1987. Invertebrates in the diet of brushtail possums, Trichosurus vulpecula, in lowland podocarp/broadleaf forest, Orongorongo Valley, Wellington, New Zealand. New Zealand Journal of Zoology 14: 163-177. Davidson J, Grant-Mackie JA, Morton JE, Rattenbury JA. 1969. North Cape: a scientific case for conservation. Tane 15: 5-11. de Lange PJ, Heenan PB, Dawson MI. 2003. A new species of Leucopogon (Ericaceae) from the Surville Cliffs, North Cape, New Zealand. New Zealand Journal of Botany 41: 13-21. Druce AP, Bartlett JK, Gardner RO. 1979. Indigenous vascular plants of the serpentine area of Surville Cliffs and adjacent cliff tops, northwest of North Cape, New Zealand. Tane 25, 187-206. Eyles AC. 2005. Revision of New Zealand Orthotylinae (Insecta: Hemiptera: Miridae). New Zealand Journal of Zoology 32: 181 215. Forster RR. 1970. The spiders of New Zealand. Part III. Desidae, Dictynidae, Hahniidae, Amaurobioididae, Nicodamidae. Otago Museum Bulletin 3. Gardner NW. 1967. Descriptions of six new species of land snails from the far north of New Zealand. Transactions of the Royal Society of New Zealand Zoology 8: 215-220. Gardner RO, Bartlett JK. 1980. Forest flora of the North Cape region. Tane 26: 223-234. Goulstone JF, Mayhill PC, Parish GR. 1993. An illustrated guide to the land Mollusca of the Te Paki Ecological Region, New Zealand. Tane 34: 1-32. Günther K. 1953. Über die taxonomische Gliederung und geographische Verbreitung der Insektenordnung der Phasmatodea. Beitr Entomol 3: 541-563. Holloway BA. 1961. A systematic revision of the New Zealand Lucanidae (Insecta: Coleoptera). Dominion Museum Bulletin 20. Jewell T, Brock PD. 2002. A review of the New Zealand stick insects: new genera and synonymy, keys, and a catalogue. Journal of Orthoptera Research 11: 189-197. 126 Klug R, Bradler S. 2006. The pregenital abdominal musculature in phasmids and its implications for the basal phylogeny of Phasmatodea (Insecta: Polyneoptera). Organisms, Diversity and Evolution 6: 171-184. Larochelle A, Larivière MC. 2005. Harpalini (Insecta: Coleoptera: Carabidae: Harpalinae). Fauna of New Zealand, vol. 53. Manaaki Whenua Press, Lincoln, New Zealand. Littig KS. 1942. External anatomy of the Florida walking stick Anisomorpha buprestoides Stoll. The Florida Entomologist 3: 33-41. Lux J, Holland W, Rate S, Beadel S. 2009. Natural areas of Te Paki ecological district: reconnaissance survey report for the protected natural areas programme. Unpublished report, Department of Conservation, Whangarei, New Zealand. Salmon JT. 1948. New genera, species and records of Orthoptera from the Three Kings Islands New Zealand. Records of the Auckland Institute and Museum 3: 291-311. Salmon JT. 1991. The Stick Insects of New Zealand. Reed, Auckland, New Zealand. von Konrat MJ, Braggins JE. 2005. Frullania wairua, a new and seemingly rare liverwort species from Northland, New Zealand. New Zealand Journal of Botany 43: 885-893. Winterbourn MJ. 2009. A new genus and species of Leptophlebiidae (Ephemeroptera) from northern New Zealand. New Zealand Journal of Zoology 36: 423-430. Whiting MF, Bradler S, Maxwell T. 2003. Loss and recovery of wings in stick insects. Nature 421: 264-267.