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1 Zootaxa 3900 (4): Copyright 2014 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) Revision of the stick insect genus Clitarchus Stål (Phasmatodea: Phasmatidae): new synonymies and two new species from northern New Zealand THOMAS R. BUCKLEY 1,2,3, SHELLEY S. MYERS 4,2,3 & SVEN BRADLER 5 1 Landcare Research, Private Bag 92170, Auckland, New Zealand. buckleyt@landcareresearch.co.nz 2 School of Biological Sciences, The University of Auckland, Private Bag 92019, Auckland, New Zealand 3 Allan Wilson Centre, Auckland, New Zealand 4 Landcare Research, Private Bag 92170, Auckland, New Zealand. shelley_m5@hotmail.com 5 Johann-Friedrich-Blumenbach-Institut für Zoologie und Anthropologie, Georg-August-Universität Göttingen, Berliner Strasse 28, Göttingen, Germany. sbradle@gwdg.de Abstract We describe two new species of Clitarchus Stål from Northland, New Zealand. Clitarchus rakauwhakanekeneke sp. nov. is described from the Poor Knights Islands and Clitarchus tepaki sp. nov. is described from the Te Paki / North Cape area and the Karikari Peninsula at the northernmost tip of New Zealand. Two new synonymies are proposed including Clitarchus multidentatus Brunner (syn. nov.) and Clitarchus tuberculatus Salmon (syn. nov.) as synonyms of Clitarchus hookeri (White). Clitarchus magnus Brunner, recorded from Thailand, is transferred to Ramulus Saussure and given the replacement name Ramulus changmaiense nom. nov. The holotype of C. multidentatus was recorded as being collected from New Caledonia; however we believe this is a labelling error and the specimen was collected from New Zealand. These taxonomic changes render Clitarchus endemic to New Zealand and consisting of three species; C. hookeri, C. rakauwhakanekeneke and C. tepaki. Keys to the adult males and females of Clitarchus species are given in addition to notes on host plants, ecology and geographic distributions. Key words: Lanceocercata, Phasmatinae, Acanthoxylini, Poor Knights Islands, Te Paki Introduction The New Zealand stick insect species Clitarchus hookeri (White) is found throughout most of the North Island and coastal regions of the north and east of the South Island (Buckley et al. 2010a). This species was among the first stick insects to be described from New Zealand based on specimens collected on the voyage of the Erebus and Terror, which visited the Bay of Islands, Northland, in 1841 (White 1846). Clitarchus hookeri was initially placed in the genus Phasma Lichtenstein, but was transferred to Clitarchus Stål (Stål 1875) of which the type species is C. laeviusculus Stål by subsequent designation of Kirby (1904). Further species of Clitarchus Stål were described from New Zealand by Colenso (1885), Hutton (1899), Brunner (1907) and Salmon (1991). Other stick insect species from New Caledonia and Thailand were placed in Clitarchus by Brunner (1907). Most recently, Salmon (1991) described the New Zealand species C. tuberculatus Salmon for female specimens of Clitarchus that were brown coloured with abundant tubercles on the body. Subsequent genetic studies (Trewick et al. 2005; Buckley et al. 2008, 2010a) showed that individual females matching the description of C. tuberculatus could not be differentiated genetically from C. hookeri suggesting that C. tuberculatus is a synonym of C. hookeri (see, also, Jewell & Brock 2002), and it was treated as such by Trewick et al. (2005), Buckley et al. (2008, 2010a), but not formalised. Clitarchus is placed in the Phasmatidae, Phasmatinae, Acanthoxylini by Günther (1953) and subsequent cataloguers (Otte and Brock 2005). Phylogenetic studies (Buckley et al. 2009; 2010b; Bradler 2009) show that along with all other New Zealand stick insects, Clitarchus is a member of the Australasian clade Lanceocercata (Bradler 2001). Clitarchus is sister to Pseudoclitarchus Salmon and this clade in turn is related to Acanthoxyla Uvarov and Tepakiphasma Buckley & Bradler (Buckley et al. 2008; 2010b; Dunning et al. 2013; Trewick et al. 2008). Accepted by B. Mantovani: 17 Nov. 2014; published: 24 Dec

2 Despite the revision of Salmon (1991), it has been known for some time that a potential new species of Clitarchus is present on the Poor Knights Islands off the eastern coast of Northland (Watt 1982). The genetic divergence of the Poor Knights Clitarchus was demonstrated in the phylogeographic study of Buckley et al. (2010a). This same study also indicated the presence of a further undescribed species in the Te Paki/North Cape area, at the northern tip of the North Island. The occurrence of two undescribed species and the presence of likely synonymies indicate that a revision of the genus is required (Jewell and Brock 2002; Buckley et al. 2010a). Many New Zealand stick insects demonstrate extreme variation in colouration and the size and distribution of spines and tubercles (Salmon 1991; Trewick et al. 2005; Buckley et al. 2010a). For these reasons, attention to the structures of the terminalia is likely to yield more robust information on species limits. Here we present morphological data from species of Clitarchus, describe two new species, redescribe C. hookeri, and formalize previously proposed synonymies. Material and methods Taxonomic and morphological studies Abbreviations AMNZ BMNH LUNZ MONZ NHMW NZAC Auckland War Memorial Museum, Auckland, New Zealand Natural History Museum, London, United Kingdom Lincoln University Entomology Research Museum, Canterbury, New Zealand Te Papa Tongarewa Museum of New Zealand, Wellington, New Zealand Naturhistorisches Museum Wien, Vienna, Austria New Zealand Arthropod Collection, Landcare Research, Auckland, New Zealand Specimens were collected from throughout the geographic distribution of Clitarchus in New Zealand and detailed examinations made of these along with specimens already housed in the NZAC. Other non-type material from MONZ, LUNZ, and AMNZ was also inspected to confirm the classification described here. All available primary type specimens were also directly inspected. Locality codes for New Zealand follow Crosby et al. (1998). We follow the higher level nomenclature of Günther (1953) instead of more recent classifications for reasons outlined in Klug & 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 and with a drawing tube (camera lucida). We use terminology of adult morphological characters following Beier (1968) and Littig (1942) and that of egg characters following Clark Sellick (1997, 1998). Results Taxonomy Order PHASMATODEA Jacobson & Bianchi Family PHASMATIDAE Gray Subfamily PHASMATINAE Gray Tribe ACANTHOXYLINI Bradley & Galil Genus Clitarchus Stål, 1875 Diagnosis. : Antennae reaching beyond tip of fore femora, claspers with 2 5 internal teeth, cerci greater than twice as long as paraprocts. 452 Zootaxa 3900 (4) 2014 Magnolia Press BUCKLEY ET AL.

3 : Antennae reaching beyond tip of fore femora. Operculum (abdominal sternum VIII) reaching beyond end of tergum IX, but not beyond base of paraprocts. Opercular organ absent or forming a sclerotized ridge rather than a spine. Tips of cerci acute and at least three times as long as paraprocts. The above diagnoses will allow Clitarchus to be differentiated from all other genera of New Zealand Phasmatodea. Type species. Clitarchus laeviusculus Stål by subsequent designation of Kirby 1904: 339. Species included. Clitarchus hookeri (White, 1846); Clitarchus rakauwhakanekeneke Buckley, Myers & Bradler, sp. nov.; Clitarchus tepaki Buckley, Myers & Bradler, sp. nov. Description. General: Medium sized stick insects, moderately robust and apterous. Body colour highly variable, most commonly light green, but also brown, grey with mottled combinations of different shades of the above colours. Body unarmed or with tubercles or short spines more common on dorsal side. Head: Prognathous, dorsoventrally flattened. Gula present. Lateral sutures present yet indistinct. Ocelli absent. Eyes prominent, hemispherical. Antenna filiform with flagellomeres, lightly setose, reaching slightly beyond end of fore femora. Scapus oval-shaped and dorsoventrally flattened, especially towards proximal end. Pedicellus cylindrical and half the length of scapus. 1 st flagellomere over twice as long as pedicellus and longer than 2 nd flagellomere. Terminal flagellomere longest and longer than scapus. 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. 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. Mesothorax longer than prothorax. Mesothoracic episternum elongated, significantly longer than epimeron. Episternal suture present. Metathorax + median segment (1 st abdominal segment) marginally shorter than or approximately equal in length to mesothorax. Metathoracic episternum elongated, significantly longer than epimeron, with row of small, rounded tubercles. Abdomen: Ten-segmented abdomen. 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 IX longer than wide. 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 transversely divided, posterior part (poculum, subgenital plate) free, not connected to dorsal part of corresponding segment, slightly smaller than anterior part. Vomer absent, remnants visible as sclerotized area between paraprocts. Paraprocts and epiproct largely concealed beneath posterior margin of tergum X, rounded at end and approximately equal in length. Claspers formed by tergum X, short, row of 3 5 dark tipped teeth. Cerci unsegmented, dorsoventrally flattened, elongated with rounded end, covered in setae, projecting well beyond 10 th abdominal segment. Phallic organ lobiform with several membranous, indistinct lobes with weak or no sclerotization. Legs: Femora and tibiae appearing pentagonal in cross section with setae running along carinae. Setae increasing in density from femur to tibia to tarsus and especially and numerous and long on pretarsus and uncus. Femora and tibiae with variable spines along carinae. Trochanters small, fused to femora. Fore femora compressed basally and curved to accommodate head. Pair of apical spines always present. Fore tibiae approximately equal in length to fore femora. Tarsus with five tarsomeres, all shorter than femora and tibiae. Mid femora with pair of apical spines. Hind femora with pair of apical spines. General: Medium sized stick insects, moderately robust and apterous. Body colouration as for male. Body unarmed or with tubercles, stout, short spines or slender sharp spines more common on dorsal side. Head: Prognathous, dorsoventrally flattened and longer than wide. Gula present, shorter than wide. Lateral sutures indistinct. Ocelli absent. Eyes prominent, hemispherical. Antenna filiform with flagellomeres, covered in setae, reaching just 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 twice as long as 2 nd. 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 dark brown, glossa and paraglossa very dark. Thorax: Prothorax slightly shorter than head, longer than wide. Openings of pair of defensive glands located laterally at the front margin of the pronotum, slit-like, indistinct. Prothoracic basisternite short, of equal length as furcasternite. Mesothorax longer than wide and greater REVISION OF CLITARCHUS Zootaxa 3900 (4) 2014 Magnolia Press 453

4 than three times longer than prothorax. Episternum elongated and epimeron short. Wing remnants present. Metathorax + median segment (1 st abdominal segment) marginally shorter than mesothorax. Episternum elongated and epimeron short. Abdomen: Ten-segmented abdomen. First abdominal segment (segmentum medianum) fused to metathorax, shorter than metanotum, boundary indistinct. Tergum II marginally shorter than III, terga III VI approximately equal in size, terga VIII to X with gradual and slight decrease in length with X being the shortest. Operculum (abdominal sternum VIII) boat shaped with carina along ventral surface, forming a sclerotized ridgelike opercular organ at base. Tip of operculum acute or rounded. Terminal margin of tergum X truncated, with epiproct protruding slightly beyond end. Paraprocts posteriorly rounded and only slightly longer than wide. Terminal ventral area posterior to ovipositor valves (gonapophyses and gonoplacs) twice as long as wide with median groove and carina either side. Gonapophyses VIII (GpVIII) reaching nearly to end of Gp IX. Gonoplac finger like, with many setae at tip, extending well beyond other valves to anterior margin of tergum X. Gonangulum well posterior to Gp VIII and Gp IX with broad, flat posterior margin. Cerci unsegmented, flattened and foliaceous and protruding 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. Legs: Femora and tibiae pentagonal in cross section with setae running along carina. Setae increasing in density from femora to tibiae. Femora and tibiae with variable spines along carinae Fore femora compressed basally and curved to accommodate head. Pair of apical spines present. Tarsus with five tarsomeres, all shorter than femora and tibiae. Terminal tarsomere bearing two non-pectinate claws. Mid femora with pair of apical spines. Mid tibiae with pair of apical spines. Hind femora with pair of apical spines. Hind tibiae with pair of apical spines. Egg: Slightly elongated along anterior-posterior axis, laterally flattened, opercular angle close to zero. Capsule rough with scattered small pits and low ridges. Colour mottled light grey to dark brown. Keel distinct, running from opercular collar to micropylar plate. Micropylar plate elliptical and variable in size. Opercular collar with blunt spines, prominent, low or absent. Operculum with pyramidal, capitular cone. Clitarchus hookeri (White, 1846) (Figs 1a, 2a,b, 3a, 4a, 5a, b, Tables 1, 2) hookeri: White, 1846, p. 24 (Phasma). laeviusculus: Stål, 1875, p. 82; Ragge, 1965, p. 39, synonym of C. hookeri. coloreus: Colenso, 1885, p. 151; Brunner, 1907, p. 151, synonym of C. hookeri. minimus: Colenso, 1885, p. 151; Brunner, 1907, p. 237, synonym of C. hookeri. reductus: Hutton, 1899, p. 55; Salmon, 1991, p. 82, synonym of C. hookeri. interruptelineatus: Brunner, 1907, p. 236; Brock, 1997, p. 236, synonym of C. hookeri. multidentatus, Brunner, 1907, p. 237, syn. n. tuberculatus: Salmon, 1991, p. 85, syn. n. Type material examined. Bacillus hookeri Holotype (BMNH) [HOLO-TYPE // N Zeal. B of Is. / // Bacillus hookeri White // Bacillus hookeri White New Zealand // BMNH(E) #844984]; Clitarchus laeviusculus Stål Lectotype (NHMW) [Coll. Br. v. W. Neu-Seeland Boucard // det. Br. v. W. Clitarchus laeviusculus // // 8947 // Phas. Inv. Nr. 418]; Clitarchus interruptelineatus Brunner Lectotype (NHMW) [Neu Seeland. Auckland. Coll. Redt. // Collectio Br. v. W. // det. Br. v. W. Clitarchus interruptelineatus // Phas. Inv. Nr. 419]; Clitarchus multidentatus Brunner Holotype. (NHMW) [Coll. Br. v. W. Neu-Caledon Deyrolle // det. Br. v. W. Clitarchus multidentatus // 5463 // // Phas. Inv. Nr. 420]; Clitarchus tuberculatus Salmon Holotype. (MONZ) [South Karori, Wellington, 2 Jan 1943, J.T. Salmon]. Additional material examined. See Appendix 1. Diagnosis. : Row of 3 5 teeth on claspers, all approximately equal in size with occasional much smaller teeth adjacent, tergum IX slightly less or greater than twice the length of tergum X. : Slender spines absent from dorsal surface, opercular organ with very weakly sclerotized carina. Egg: Egg width mm. Description. General: Medium-sized stick insect, moderately robust and apterous. Body green to mottled brown and grey. Most individuals without tubercles, or less commonly large tubercles or rarely blunt, stout spines present on dorsal surface. Head: Prognathous, dorsoventrally flattened and longer than wide with broad dark strip 454 Zootaxa 3900 (4) 2014 Magnolia Press BUCKLEY ET AL.

5 running along ventral lateral sides. Gula present, shorter than wide. Vertex of head unarmed or less commonly with a few small tubercles. Lateral sutures present yet indistinct. Ocelli absent. Eyes prominent, hemispherical. Antenna filiform with flagellomeres, lightly setose, reaching slightly beyond end of fore femora. Scapus oval shaped and dorsoventrally flattened, especially towards proximal end. Pedicellus cylindrical and half the length of scapus. 1 st flagellomere over twice as long as pedicellus and longer than 2 nd flagellomere. Slight basal swelling at base of 8 th or 9 th flagellomere. Terminal flagellomere longest and longer than scapus. 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. Mesothorax longer than wide and approximately 4 times longer than prothorax. Mesonotum unarmed or with several body coloured tubercles, with dark median line and longitudinal stripes at anterior and posterior margins. Mesothoracic episternum elongated, significantly longer than epimeron, sometimes with small, rounded tubercles. Episternal suture present. Mesothoracic basisternite and furcasternite unarmed with several very small tubercles. Metathorax + median segment (1st abdominal segment) approximately equal in length to mesothorax with dark longitudinal stripe along median of ventral surface, usually continuous. Metanotum and metasternum unarmed or occasionally with scarce small tubercles. Metathoracic episternum elongated, significantly longer than epimeron, sometimes with small, rounded 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 IX longer than wide, tergum IX slightly less or greater than twice the length of tergum X. 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. Short longitudinal dark stripes at posterior margins of tergites and occasionally with small tubercles on terga II V. Sterna unarmed. 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 blunt with shallow notch and reaching less than half way along 9 th tergum, with broad lateral carina on ventral surface. Vomer absent, remnants visible as sclerotized area between paraprocts. Paraprocts and epiproct largely concealed beneath posterior margin of tergum X, rounded at end and approximately equal in length. Claspers formed by tergum X, short, row of 3 5 dark tipped teeth or variable size. Occasionally smaller teeth present offset from row of larger teeth. Cerci dorsoventrally flattened, elongated with rounded end, covered in setae, approximately equal in length to sternum X and projecting well beyond 10 th abdominal segment. Phallic organ lobiform with several membranous, indistinct lobes with dorsal sclerite only weakly sclerotized. Legs: Femora and tibiae appearing pentagonal in cross section with setae running along carinae. Setae increasing in density from femur to tibia to tarsus and especially numerous and long on pretarsus and uncus. Fore coxa usually with single tubercle. Trochanters small, fused to femora. Fore femora compressed basally and curved to accommodate head, with 3 6 short spines on latero-ventral carinae. Pair of apical spines always 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, 2 nd, 5 th, 3 rd, 4 th. Mid coxae unarmed. Mid femur with 0 4 small spines along ventral carina, 0 4 teeth along latero-ventral carina, dorsal carina unarmed. Pair of apical spines always present. Mid tibia unarmed except for pair of apical spines. Tarsus as for fore leg. Hind coxa unarmed. Hind femur with 0 3 small spines along median of ventral surface and 1 3 spines on latero-ventral carina. Pair of apical spines always present. Hind tibia unarmed except for pair of apical spines. Tarsus as for fore leg. General: Medium-sized stick insect, moderately robust and apterous. Body green to mottled brown and grey. Body mostly unarmed, but sometimes tubercles and rarely, blunt spines present on dorsal side. Head: Prognathous, dorsoventrally flattened and longer than wide with broad dark strip running along ventral lateral sides. Gula present, shorter than wide. Vertex of head usually unarmed, or with a few tubercles and rarely with stout, blunt spines. Lateral sutures indistinct. Ocelli absent. Eyes prominent, hemispherical. Antenna filiform with flagellomeres covered in setae, reaching just 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 twice as long as 2 nd. Slight basal swelling at base of 9 th or 11 th flagellomere. Terminal flagellomere longest. Labial palps with three articles slightly dorsoventrally flattened. REVISION OF CLITARCHUS Zootaxa 3900 (4) 2014 Magnolia Press 455

6 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 dark brown, glossa and paraglossa very dark. Thorax: Prothorax slightly shorter than head, longer than wide. Pronotum unarmed or with few small tubercles and dark median line. Openings of pair of defensive glands located laterally at the front margin of the pronotum, slit-like, indistinct. Furcasternite, basisternite, prothoracic episternum and epimeron unarmed. Prothoracic basisternite short, of equal length to furcasternite. Mesothorax longer than wide and greater than three times longer than prothorax. Mesonotum unarmed or with very few tubercles. Mesothoracic episternum elongated, episternal suture with row of small tubercles or unarmed. Epimeron usually unarmed or with very few tubercles. Mesosternum unarmed or less commonly with few tubercles. Wing remnants present. Metathorax + median segment (1st abdominal segment) marginally shorter than mesothorax. Metanotum unarmed and with dark median stripe thicker at anterior and posterior margins. Metathoracic episternum elongated, episternal suture usually with row of small tubercles. Metathoracic epimeron usually unarmed with few very small tubercles. Metasternum unarmed. Abdomen: Ten-segmented abdomen unarmed. First abdominal segment (segmentum medianum) fused to metathorax, shorter than metanotum, boundary indistinct. Spines very small or absent from sternites. Posterior margins of terga with dark strip medially, sometimes flanked by lighter patches and darker patches more laterally. Terga II and III same length, terga III VI approximately equal in size, terga IX and X shorter with X being the shortest. Operculum (abdominal sternum VIII) boat shaped with carina along ventral surface, forming a weakly sclerotized ridge-like opercular organ at base and not sclerotized laterally. Tip of operculum usually acute and not reaching end of tergum X. Terminal margin of tergum X truncated, with epiproct protruding slightly beyond end. Paraprocts posteriorly rounded and only slightly longer than wide. Terminal ventral area posterior to ovipositor valves twice as long as wide with median groove in posterior half and carina either side. Gonapophyses VIII (GpVIII) reaching nearly to end of Gp IX. Gonoplac finger like, with many setae at tip, extending beyond other valves but not as far as anterior margin of tergum X. Gonangulum well posterior to Gp VIII and Gp IX with broad, flat posterior margin. Cerci acute, flattened, foliaceous, over 3 times longer than paraprocts and protruding well 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 edge of cerci usually straight with inside edge curving outwards towards tip. Legs: Femora and tibiae pentagonal in cross section with setae running along carinae. Setae increasing in density from femora to tibiae. Fore coxa with 1 or more tubercles. Fore femora compressed basally and curved to accommodate head, external latero-ventral carina with row of 0 6 teeth, dorsal and ventral carina unarmed. Pair of apical spines present. Fore tibiae slightly shorter, slightly longer or same length as fore femora and unarmed except for apical teeth. Tarsomeres all shorter than femora and tibiae, and in decreasing order of 1 st, 2 nd, 5 th, 3 rd, 4 th. Terminal (5 th ) tarsomere bearing two non-pectinate claws. Mid coxa usually unarmed. Mid femur with 0 3 teeth along ventral carinae, 1 2 teeth along latero-ventral carinae and dorsal carina unarmed. Pair of apical spines present. Mid tibia occasionally with single tooth near proximal end of ventral carinae and dorsal carina unarmed. Pair of apical spines present. Tarsus as for fore leg. Hind coxa usually unarmed. Hind femur with 0 3 teeth on ventral carina, 1 2 teeth along latero-ventral carina, dorsal carina unarmed. Pair of apical spines present. Hind tibia with pair of apical teeth and otherwise unarmed or with single dorsal tooth, apical spines present. Tarsus as for fore leg. Egg: (Fig. 5a,b,c,d, Table 2) Slightly elongated along anterior-posterior axis, laterally flattened, opercular angle close to zero. Egg width mm. Capsule rough with scattered small pits and low ridges. Colour mottled light grey to dark brown. Keel distinct, running from opercular collar to micropylar plate. Micropylar plate elliptical and variable in size. Opercular collar with low blunt spines, sometimes absent. Operculum with pyramidal, capitular cone. Notes on synonyms and type specimens. Unfortunately the type material of C. coloreus (Colenso), C. minimus (Colenso) and C. reductus Hutton cannot be located (Jewell & Brock 2002) making definitive conclusion about the status of these names problematic. The description of C. coloreus differentiates this species from C. hookeri solely on the basis of colour and spination of the femora (Colenso 1885), both characters which vary extensively within and among Clitarchus species. The type locality of C. coloreus is Waipawa, Hawkes Bay. Our collections in Hawkes Bay have shown only the presence of C. hookeri. Therefore we have maintained the synonymy of Brunner (1907) where C. coloreus is considered a junior synonym of C. hookeri. 456 Zootaxa 3900 (4) 2014 Magnolia Press BUCKLEY ET AL.

7 FIGURE 1. Habitus photos of (a) mating pair of Clitarchus hookeri from Great Barrier Island; (b) female and male of Clitarchus rakauwhakanekeneke sp. nov., Tawhiti Rahi, Poor Knights Islands; (c) male of Clitarchus tepaki sp. nov., Kohuronaki, Te Paki. REVISION OF CLITARCHUS Zootaxa 3900 (4) 2014 Magnolia Press 457

8 FIGURE 2. Illustrations of terminalia of Acanthoxylini males; left: lateral view; right: posterior view of abdominal tergum 10 (clasper). (a,b) Clitarchus hookeri; (c,d) C. rakauwhakanekeneke sp. nov.; (e,f) C. tepaki sp. nov.; (g,h) Pseudoclitarchus sentus; (i,j) Tepakiphasma ngatikuri. cer, cercus; svii six, abdominal sternum VII IX; stiviii, abdominal stigmum VIII; tvii tx, abdominal tergum VII X; poc, poculum; tho, thorns. Scale: 2 mm. 458 Zootaxa 3900 (4) 2014 Magnolia Press BUCKLEY ET AL.

9 FIGURE 3. Illustrations of terminalia of Acanthoxylini males, ventral view. (a) Clitarchus hookeri; (b) C. rakauwhakanekeneke; (c) C. tepaki; (d) Pseudoclitarchus sentus; (e) Tepakiphasma ngatikuri. cer, cercus; epi, epiproct; par, paraproct; tix, tx, abdominal tergum XI, X; tho, thorns. Scale: 1 mm. Clitarchus minimus was described from a single specimen about 8.5 lines (18 mm) long, smooth and green. Although Colenso (1885) believed the specimen to be an adult, we agree with Hutton (1898) that it is most likely a nymph based on the reported body length. No adult New Zealand stick insect is nearly that short (Salmon 1991). From this description and other characters listed in Colenso (1885) we are unable to determine if the now lost type is a nymph of C. hookeri or even another genus of New Zealand stick insect. Hutton (1898) in fact placed the name as incertae sedis. However, we have retained the synonymy of Salmon (1991) as there is insufficient evidence to challenge Salmon s placement of the name. The type locality of C. minimus is Waipawa, Hawkes Bay of the eastern North Island, well within the geographic distribution of C. hookeri (Buckley et al. 2010a). REVISION OF CLITARCHUS Zootaxa 3900 (4) 2014 Magnolia Press 459

10 FIGURE 4. Illustrations of terminalia of Acanthoxylini females, lateral view. (a) Clitarchus hookeri; (b) C. rakauwhakanekeneke; (c) C. tepaki; (d) Pseudoclitarchus sentus; (e) Tepakiphasma ngatikuri; (f) Acanthoxyla inermis. cer, cercus; gpl, gonoplac; op, operculum; opo, opercular organ; svii, sviii, abdominal sternum VII, VIII; stiviii, abdominal stigmum VIII; tvii tx, abdominal tergum VII X. Scale: 2 mm. 460 Zootaxa 3900 (4) 2014 Magnolia Press BUCKLEY ET AL.

11 FIGURE 5. Lateral (a) and dorsal (b) views of eggs from Clitarchus hookeri, (c f) C. rakauwhakanekeneke from Warawara Forest, Northland (c,d) and from Otari-Wiltons Bush, Wellington (e,f), and (g,h) C. tepaki. Scale: 1 mm. The type locality of C. reductus is within Canterbury and all specimens of Clitarchus we have observed from this region are C. hookeri. The character states described by Hutton (1899) fall within the range of variation observed within C. hookeri. The male of C. reductus was not described because males of C. hookeri are not known from the South Island (Buckley et al. 2010a). These findings therefore support Salmon s (1991) synonymy of C. reductus with C. hookeri. Ragge (1965) synonymized Clitarchus laeviusculus Stål with C. hookeri. The lectotype of Clitarchus laeviusculus is a female and listed as being from Canterbury by Hutton (1899). We have studied the lectotype and agree with this synonymy. Brock (1997) synonymized C. interruptelineatus Brunner with C. hookeri. The lectotype of C. interruptelineatus is a male and our inspection of the claspers of the lectotype show that they match those of C. hookeri, supporting the synonymy of Brock (1997). One of the paralectotypes was collected from Great Barrier Island, and C. hookeri is recorded from this locality. We have examined the female holotype of C. multidentatus, and the relevant characters match those of C. hookeri even though the label data reports the specimen was collected from New Caledonia. Because there are no further records of Clitarchus from New Caledonia, we believe the specimen was mislabeled and was in fact collected from New Zealand. We have therefore placed C. multidentatus as a junior synonym of C. hookeri. Brunner (1907) described the species Clitarchus magnus from South East Asia. Hennemann et al. (2008) noted that Clitarchus magnus Brunner was not a species of Clitarchus at all, but most likely belonged in the Clitumninae, tentatively suggesting Ramulus Saussure as an appropriate genus. Inspection of an image of the female holotype of Clitarchus magnus does indeed demonstrate strong similarity to Ramulus and other Clitumninae genera. However, the name Ramulus magnus (Brunner, 1907: 190) is preoccupied by the species initially described as Clitumnus magnus Brunner and treated as a member of Ramulus by Otte & Brock (2005). Therefore we have selected a replacement name for Clitarchus magnus Brunner, 1907: 236 and transferred it to Ramulus. The replacement name REVISION OF CLITARCHUS Zootaxa 3900 (4) 2014 Magnolia Press 461

12 is Ramulus changmaiense nom. nov. The above synonymies and replacement name render Clitarchus endemic to New Zealand. TABLE 1. Range of body dimensions (mm) of Clitarchus species. The following sample sizes were used: C. hookeri male (n=10), C. hookeri female (n=10), C. rakauwhakanekeneke sp. nov. male (n=2), C. rakauwhakanekeneke sp. nov. female (n=4), C. tepaki sp. nov. male (n=8), C. tepaki sp. nov. female (n=10). C. hookeri C. rakauwhakanekeneke sp. nov. C. tepaki sp. nov. Female Body Head Antennae Pronotum Metanotum Mesonotum Abdomen Fore femur Mid femur Hind femur Fore tibia Mid tibia Hind tibia Male Body Head Antennae Pronotum Metanotum Mesonotum Abdomen Fore femur Mid femur Hind femur Fore tibia Mid tibia Hind tibia TABLE 2. Range of egg dimensions (mm) of Clitarchus species. The following sample sizes were used: C. hookeri (n=31), C. rakauwhakanekeneke sp. nov. (n=26), C. tepaki sp. nov. (n=30). C. hookeri C. rakauwhakanekeneke sp. nov. C. tepaki sp. nov. Height 4.5 ( ) 5.0 ( ) 4.8 ( ) Width 1.4 ( ) 3.0 ( ) 2.4 ( ) Height/width ratio 1.6 ( ) 1.6 ( ) 2 ( ) Operculum height 0.6 ( ) 0.7 ( ) 0.7 ( ) Operculum width 1.0 ( ) 1.3 ( ) 1.3 ( ) 462 Zootaxa 3900 (4) 2014 Magnolia Press BUCKLEY ET AL.

13 The most recently described species of Clitarchus is C. tuberculatus Salmon (1991). Salmon (1991) believed C. tuberculatus to be a new species based on the colouration and presence of tubercles on the body. Salmon (1991) also believed the species to be parthenogenetic and he apparently found no males associated with the C. tuberculatus females or males that matched the morphology of the female. However, our own research and that of Trewick et al. (2005) has shown populations of what are clearly C. hookeri containing females matching the type of C. tuberculatus. We have also observed many males that are brown coloured and with many tubercles, like the females of C. tuberculatus, and we therefore conclude that these different morphs represent a single species. Distribution and ecology. Clitarchus hookeri is found from the southern end of the Aupouri Peninsula south, throughout most of the rest of the North Island (Buckley et al. 2010a). It appears to be absent from some areas of the central North Island. On the South Island its range is more restricted and it is generally less abundant being found mainly in coastal areas from Northwest Nelson through Marlborough, Kaikoura, and Canterbury areas, south to Dunedin (Buckley et al. 2010a). It has also been introduced to Great Britain (Brock 1987). The species is a geographic parthenogen and males have not been recorded from the South Island (Buckley et al. 2010a; Morgan- Richards et al. 2010). The species is polyphagous and has been observed feeding on Metrosideros perforata, Metrosideros excelsa, Leptospermum scoparium, Kunzea spp. (all Myrtaceae), Muehlenbeckia australis (Polygonaceae), Rubus sp. (Rosaceae) and Coprosma spp. (Rubiaceae) (Salmon 1991; Buckley et al. 2010a). It is most commonly collected from Leptospermum scoparium. The adventive plant species it has been observed feeding on include Erica cruenta (Ericaceae) and Rubus armeniacus (Rosaceae). The species can be successfully reared on Metrosideros excelsa, despite it being uncommonly collected from this tree. Information on development and lifecycle is given in Stringer (1969, 1970) and Salmon (1991). Studies on parthenogenesis and evolutionary history are given in Buckley et al. (2008, 2009, 2010a,b), Buckley & Leschen (2013), Dunning et al. (2013), Morgan-Richards & Trewick (2005) and Morgan-Richards et al. (2010), Myers et al. (2013) and Trewick et al. (2005, 2008). Clitarchus rakauwhakanekeneke Buckley, Myers & Bradler, sp. nov. (Fig. 1b, 2c,d, 3b, 4b, 5c f, Tables 1,2) Official Māori name: Rākau Whakanekeneke Material examined. Holotype. (NZAC) [NEW ZEALAND, ND, Poor Knights Is, Tawhiti Rahi, 8 Dec 1980, JC Watt, beating from Leptospermum ericoides] [NZAC ] [HOLOTYPE Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler] Paratypes. (NZAC) [NEW ZEALAND, ND, Poor Knights Is, Tawhiti Rahi, 4 Dec 1980, R Kleinpaste] [NZAC ] [PARATYPE Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler]; (NZAC) [NEW ZEALAND, ND, Aorangi, Poor Knights Islands, Urupa Point, 12 Nov 1981, J C Watt] [NZAC ] [PARATYPE Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler]; (NZAC) [NEW ZEALAND, ND, Aorangi, Poor Knights Islands, , , 9 Jun 2006, T R Buckley D Seldon] [NZAC ] [PARATYPE Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler]; (NZAC) [NEW ZEALAND, ND, Poor Knights Is, Aorangi, , , June 9 th 2006, T R Buckley, D Seldon] [NZAC ] [PARATYPE Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler]; (NZAC) [NEW ZEALAND, ND, Tawhiti Rahi, Poor Knights Is, Main Trk, top of plateau, , , 17 Dec 2009, T R Buckley R Leschen D Seldon] [NZAC ] [PARATYPE Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler]; (NZAC) [NEW ZEALAND ND Tawhiti Rahi Poor Knights Is Main Trk top of plateau 17 Dec 2009 T R Buckley R Leschen D Seldon beating S E] [NZAC ] [PARATYPE Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler]; Additional material examined. See Appendix 1. Diagnosis. : Row of 3 prominent teeth on claspers; inner tooth massively enlarged with middle tooth slightly closer to outer tooth. Tergum IX less than twice the length of tergum X. : Slender spines present on dorsal surface of head, thorax or abdomen, opercular organ ridge-like and flanked by sclerotized patch. Egg: Egg width mm. Description. General: Medium-sized stick insect, moderately robust and apterous. Body green to mottled REVISION OF CLITARCHUS Zootaxa 3900 (4) 2014 Magnolia Press 463

14 brown and grey. Tubercles present, more common on dorsal side. Head: Prognathous, dorsoventrally flattened and longer than wide with broad dark strip running along ventral lateral sides. Gula present, shorter than wide. Vertex of head with a few small tubercles. Lateral sutures present yet indistinct. Ocelli absent. Eyes prominent, hemispherical. Antenna filiform with flagellomeres, lightly setose, reaching slightly beyond end of fore femora. Scapus oval shaped and dorsoventrally flattened, especially towards proximal end. Pedicellus cylindrical and half the length of scapus. 1 st flagellomere over twice as long as pedicellus and longer than 2 nd flagellomere. Slight basal swelling at base of 9 th flagellomere. Terminal flagellomere longest and longer than scapus. 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 teeth: one apical tooth and two subapical 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 to furcasternite, both sternites without spines. Mesothorax longer than wide and more than 3 times longer than prothorax. Mesonotum unarmed with dark longitudinal stripes at anterior and posterior margins, sometimes continuous. Mesothoracic episternum elongated, significantly longer than epimeron, with row of small, rounded tubercles. Episternal suture present. Mesothoracic basisternite and furcasternite unarmed with several very small tubercles. Metathorax + median segment (1st abdominal segment) slightly shorter than mesothorax with dark longitudinal stripes at anterior and posterior margins. Metanotum and metasternum unarmed or with scarce small tubercles. Metathoracic episternum elongated, significantly longer than epimeron, with row of small, rounded 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 IX longer than wide, tergum IX slightly less than twice the length of tergum X. 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. Short longitudinal dark stripes at posterior margins of terga with sparse, small tubercles. Sparse very small tubercles on sterna. Sternum IX transversely divided, posterior part (poculum, subgenital plate) free, not connected to dorsal part of corresponding segment, slightly smaller than anterior part. Tip of subgenital plate blunt with shallow notch and reaching less than half way along 9 th tergum, with broad lateral carina on ventral surface. Vomer absent, remnants visible as sclerotized area between paraprocts. Paraprocts and epiproct largely concealed beneath posterior margin of tergum X, rounded at end and approximately equal in length. Claspers formed by tergum X, short, row of 3 dark tipped teeth, inner tooth enlarged. Middle tooth slightly closer to outer tooth. Sometimes a 4 th asymmetric small tooth developed. Cerci dorsoventrally flattened, elongated with rounded end, covered in setae, slightly shorter than tergum X and projecting well beyond 10 th abdominal segment. Phallic organ lobiform with several membranous, indistinct lobes; dorsal sclerite only weakly sclerotized. Legs: Femora and tibiae appearing pentagonal in cross section with setae running along carinae. Setae increasing in density from femur to tibia to tarsus and especially and numerous and long on pretarsus and uncus. Fore coxae with several short blunt spines. Trochanters small, fused to femora. Fore femora compressed basally and curved to accommodate head, with 0 3 short spines on latero-ventral carinae. Pair of apical spines always 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, 2 nd, 5 th, 3 rd, 4 th. Mid coxa unarmed. Mid femur with 0 3 small spines along ventral carina, 0 4 teeth along latero-ventral carina, dorsal carina unarmed. Pair of apical spines always present. Mid tibia unarmed except for pair of apical spines. Tarsomeres all shorter than femora and tibiae, and in decreasing order of 1 st, 5 th,2 nd, 3 rd, 4 th. Hind coxa unarmed. Hind femur with 2 small spines along median of ventral surface and pair of spines on latero-ventral carinae near apex. Pair of apical spines always present. Hind tibia unarmed except for pair of apical spines. Tarsus as for fore leg. General: Medium-sized stick insect, moderately robust and apterous. Body green to mottled brown and grey. Spines and tubercles present, more common on dorsal side. Head: Prognathous, dorsoventrally flattened and longer than wide with broad dark strip running along ventral lateral sides. Gula present, shorter than wide. Vertex of head armed with a few short spines and / or tubercles. Lateral sutures indistinct. Ocelli absent. Eyes prominent, hemispherical. Spines and tubercles absent from ventral surface of head. Antenna filiform with flagellomeres covered in setae, reaching just beyond end of fore femora. Scapus oval-shaped and dorsoventrally flattened, especially towards proximal end. Pedicellus cylindrical, slightly dorsventrally flattened and less than half 464 Zootaxa 3900 (4) 2014 Magnolia Press BUCKLEY ET AL.

15 the length of scapus. 1 st flagellomere twice as long as 2 nd. Slight basal swelling at base of 10 th or 11 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 dark brown. Glossa and paraglossa very dark. Thorax: Prothorax slightly shorter than head, longer than wide. Stout spines and tubercles on pronotum with dark median line. Openings of pair of defensive glands located laterally at the front margin of the pronotum, slit-like, indistinct. A few small tubercles on furcasternite, basisternite, prothoracic episternum and epimeron. Prothoracic basisternite short, of equal length as furcasternite. Mesothorax longer than wide and greater than three times longer than prothorax. Mesonotum armed with spines and with row of larger spines near lateral margins. Mesothoracic episternum elongated with episternal suture with row of spines, in posterior third row of spines runs slightly ventrally. Mesothoracic epimeron short with several spines or tubercles. Mesothoracic furcasternite with tubercles. Wing remnants present. Metathorax + median segment (1 st abdominal segment) marginally shorter than mesothorax. Metanotum with irregular rows of small spines with row of larger spines along lateral area. Dark median stripe thicker at anterior and posterior margins. Metathoracic episternum elongated; episternal suture with row of spines in anterior two thirds, in posterior third row of spines runs slightly ventrally. Metathoracic epimeron short with several spines or tubercles. Tubercles present on metasternum. Abdomen: Tensegmented abdomen with very small tergal spines present. First abdominal segment (segmentum medianum) fused to metathorax, shorter than metanotum, boundary indistinct. Spines very small or absent from sterna. Spines usually absent from anterior region of terga, pair of small spines near posterior margin of terga. Spines reducing in size posteriorly. Posterior margins of terga with dark strip medially, sometimes flanked by lighter patches and darker patches more laterally. Tergum II marginally shorter than III, terga III VI approximately equal in size, terga VIII to X with gradual and slight decrease in length with X being the shortest. Operculum (abdominal sternum VIII) boat-shaped with carina along ventral surface, forming a sclerotized ridge-like opercular organ at base. Tip of operculum acute or rounded and reaching slightly beyond half way of tergum X. Terminal margin of tergum X truncated, with epiproct protruding slightly beyond end. Paraprocts posteriorly rounded and only slightly longer than wide. Terminal ventral area posterior to ovipositor valves twice as long as wide with median groove and carina either side. Gonapophyses VIII (GpVIII) reaching nearly to end of Gp IX. Gonoplac finger like, with many setae at tip, extending well beyond other valves to anterior margin of tergum X. Gonangulum well posterior to Gp VIII and Gp IX with broad, flat posterior margin. Cerci flattened and foliaceous, over 3 times longer than paraprocts and protruding well 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 edge of cerci straight with inside edge curving outwards towards tip. Legs: Femora and tibiae pentagonal in cross section with setae running along carinae. Setae increasing in density from femora to tibiae. Fore coxa with several spines and tubercles. Fore femur compressed basally and curved to accommodate head, external latero-ventral carina with row of 0 2 teeth, dorsal and ventral carina unarmed. Pair of apical spines present. Fore tibiae slightly shorter than 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. Terminal (5 th ) tarsomere bearing two non-pectinate claws. Mid coxa with few spines or tubercles. Mid femur with 0 4 teeth along ventral carinae, 1 4 teeth along latero-ventral carinae and 0 2 teeth along dorsal carina. Pair of apical spines present. Mid tibia with single tooth near proximal end of ventral carinae and 0 1 tooth on dorsal carina. Pair of apical spines present. Tarsus as for fore leg. Hind coxa with few tubercles or small spines. Hind femur with 1 4 teeth on ventral carina, 1 4 teeth along latero-ventral carina, 0 2 teeth along dorsal carina. Pair of apical spines present. Hind tibia with pair of apical teeth and otherwise unarmed or with single dorsal tooth, apical spines present. Tarsus as for fore leg. Egg (Fig. 5e, f, Table 2): Slightly elongated along anterior-posterior axis, laterally flattened, opercular angle close to zero. Egg width mm. Capsule rough with scattered small pits and prominent ridges. Colour mottled light to dark brown. Keel distinct, running from opercular collar to micropylar plate. Micropylar plate elliptical and variable in size. Opercular collar with blunt spines. Operculum with pyramidal, capitular cone. Distribution and ecology. The occurrence of a possible endemic species of stick insect on the Poor Knights was first noted by Watt (1982), where it was referred to as Glitarchus [sic] aff. hookeri (p. 294). Watt referred to this population as possibly an endemic species or a definable geographic race. The population was referred to as Clitarchus sp. nov. 2 by Buckley et al. (2010a) and given a Threat Listing of Range Restricted and Island Endemic by Buckley et al. (2012). Clitarchus rakauwhakanekeneke sp. nov. is restricted to the Poor Knights REVISION OF CLITARCHUS Zootaxa 3900 (4) 2014 Magnolia Press 465