ArticlePDF Available

Revision of the stick insect genus Clitarchus Stål (Phasmatodea: Phasmatidae): New synonymies and two new species from northern New Zealand

Authors:

Abstract and Figures

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 Cli-tarchus 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.
Content may be subject to copyright.
Accepted by B. Mantovani: 17 Nov. 2014; published: 24 Dec. 2014
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3900 (4): 451
482
www.mapress.com
/
zootaxa
/
Article
451
http://dx.doi.org/10.11646/zootaxa.3900.4.1
http://zoobank.org/urn:lsid:zoobank.org:pub:032197C5-5266-43D9-AFE2-CB746A93E1F0
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. E-mail: 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. E-mail: shelley_m5@hotmail.com
5
Johann-Friedrich-Blumenbach-Institut für Zoologie und Anthropologie, Georg-August-Universität Göttingen, Berliner Strasse 28,
37073 Göttingen, Germany. E-mail: 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 Cli-
tarchus 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
Terro r, 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).
BUCKLEY ET AL.
452
·
Zootaxa 3900 (4) © 2014 Magnolia Press
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 Auckland War Memorial Museum, Auckland, New Zealand
BMNH Natural History Museum, London, United Kingdom
LUNZ Lincoln University Entomology Research Museum, Canterbury, New Zealand
MONZ Te Papa Tongarewa—Museum of New Zealand, Wellington, New Zealand
NHMW Naturhistorisches Museum Wien, Vienna, Austria
NZAC 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.
Zootaxa 3900 (4) © 2014 Magnolia Press
·
453
REVISION OF CLITARCHUS
♀: 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 18–21 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 20–22 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
BUCKLEY ET AL.
454
·
Zootaxa 3900 (4) © 2014 Magnolia Press
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 ridge-
like 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. / 43 70. //
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 // 9.8947
// 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 // 59. 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 1.1–1.7 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
Zootaxa 3900 (4) © 2014 Magnolia Press
·
455
REVISION OF CLITARCHUS
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 19
20 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
20–22 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.
BUCKLEY ET AL.
456
·
Zootaxa 3900 (4) © 2014 Magnolia Press
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 1.1–1.7 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.
Zootaxa 3900 (4) © 2014 Magnolia Press
·
457
REVISION OF CLITARCHUS
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.
BUCKLEY ET AL.
458
·
Zootaxa 3900 (4) © 2014 Magnolia Press
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.
Zootaxa 3900 (4) © 2014 Magnolia Press
·
459
REVISION OF CLITARCHUS
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).
BUCKLEY ET AL.
460
·
Zootaxa 3900 (4) © 2014 Magnolia Press
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.
Zootaxa 3900 (4) © 2014 Magnolia Press
·
461
REVISION OF CLITARCHUS
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
BUCKLEY ET AL.
462
·
Zootaxa 3900 (4) © 2014 Magnolia Press
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).
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.
Female
Body 77.9–100.3 60.7–75.5 62.4–80.8
Head 4.6–6.6 4.8–6 4.5–5.5
Antennae 20–27.5 13.8–17.6 19.0–20.0
Pronotum 3.4–4.6 3.2–4.3 2.8–4.3
Metanotum 15.3–19.6 10.6–14.3 11.4–14.8
Mesonotum 13.6–19 10.4–12.9 10.9–14.2
Abdomen 40.3–50.5 31.6–39.4 32.8–42
Fore femur 19.8–24.6 15.3–19 16.3–19.3
Mid femur 13.3–17.3 10.8–12.9 10.1–13.2
Hind femur 16.1–21 13–15.2 13–15.8
Fore tibia 19.7–24.7 14.4–18 16.1–19.7
Mid tibia 12.5–15.3 9.5–10.3 10.4–12.4
Hind tibia 15.5–19.5 12.5–13.1 13–15.3
Male
Body 59–71.3 51.2–56.3 48.5–56.8
Head 3.4–4.3 3.3–3.9 3.0–4.25
Antennae 20.2–28 16.1–18.5 16.7–21
Pronotum 2.2–3 2.9–3 1.95–2.7
Metanotum 11.1–14.3 9.6–10.9 9.1–10.2
Mesonotum 11–14.3 9.6–10.6 8.7–10.5
Abdomen 30–35.4 25.7–28 25.4–30.5
Fore femur 17–20.7 14.1–16 13.3–15.9
Mid femur 12.5–15 10.3–11.4 10.2–12
Hind femur 15–19 12.8–14.3 12.6–14.6
Fore tibia 16.7–20.7 13.6–15.6 13.6–16
Mid tibia 11.3–13.3 9.5–9.7 8.3–11.3
Hind tibia 14.2–18 11.3–12.6 11.6–14.7
C. hookeri C. rakauwhakanekeneke sp. nov. C. tepaki sp. nov.
Height 4.5 (4.0
4.9) 5.0 (4.7
5.3) 4.8 (4.6
4.9)
Width 1.4 (1.1
1.7) 3.0 (2.8
3.3) 2.4 (2.2
2.7)
Height/width ratio 1.6 (1.4
1.8) 1.6 (1.5
1.8) 2 (1.8
2.2)
Operculum height 0.6 (0.2
0.9) 0.7 (0.5
1.0) 0.7 (0.5
1.0)
Operculum width 1.0 (1.0
1.7) 1.3 (1.1
1.4) 1.3 (1.2
1.4)
Zootaxa 3900 (4) © 2014 Magnolia Press
·
463
REVISION OF CLITARCHUS
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] [NZAC03005301] [HOLOTYPE Clitarchus
rakauwhakanekeneke Buckley, Myers and Bradler]
Paratypes. ♀ (NZAC) [NEW ZEALAND, ND, Poor Knights Is, Tawhiti Rahi, 4 Dec 1980, R Kleinpaste]
[NZAC03000077] [PARATYPE Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler]; ♂ (NZAC) [NEW
ZEALAND, ND, Aorangi, Poor Knights Islands, Urupa Point, 12 Nov 1981, J C Watt] [NZAC03005385]
[PARATYPE Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler]; ♀ (NZAC) [NEW ZEALAND, ND,
Aorangi, Poor Knights Islands, 35 28.9, 174 44, 9 Jun 2006, T R Buckley D Seldon] [NZAC03000466]
[PARATYPE Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler]; ♀ (NZAC) [NEW ZEALAND, ND,
Poor Knights Is, Aorangi, 35 28.9, 174 44, June 9
th
2006, T R Buckley, D Seldon] [NZAC04028338] [PARATYPE
Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler]; ♀ (NZAC) [NEW ZEALAND, ND, Tawhiti Rahi,
Poor Knights Is, Main Trk, top of plateau, 35 27.457, 174 44.257, 17 Dec 2009, T R Buckley R Leschen D Seldon]
[NZAC03009511] [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 35 27.457 S 174 44.257 E] [NZAC04097333] [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 2.8–3.3 mm.
Description. ♂ General: Medium-sized stick insect, moderately robust and apterous. Body green to mottled
BUCKLEY ET AL.
464
·
Zootaxa 3900 (4) © 2014 Magnolia Press
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 20–21 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 20–22
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
Zootaxa 3900 (4) © 2014 Magnolia Press
·
465
REVISION OF CLITARCHUS
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: Ten-
segmented 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 2.8–3.3 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
BUCKLEY ET AL.
466
·
Zootaxa 3900 (4) © 2014 Magnolia Press
Islands 22 km from the eastern coast of Northland. It has been collected on both main islands of Tawhiti Rahi and
Aorangi, including trips by the senior author and collaborators in 2006 (Aorangi), 2010 (Tawhiti Rahi) and 2013
(Tawhiti Rahi). It is unknown if it survives on any of the smaller islands and vegetated rock stacks, many of which
would have been connected to Tawhiti Rahi and Aorangi during Pleistocene sea level lowering. Our own field
research and label data from NZAC specimens shows it has been collected from Metrosideros perforata,
Metrosideros excelsa, Leptospermum scoparium, Kunzea spp. (all Myrtaceae), Leucopogon fasciculatus
(Ericaceae), Coprosma sp. (Rubiaceae), “tussock” and other monocots. It has been successfully reared on
Metrosideros excelsa. Given the abundance of host plants on the islands and absence of introduced mammalian and
vespid wasp predators the survival of the species seems assured, contingent on no major environmental disturbance
to the islands. Information on phylogenetic relationships with other species of Clitarchus is given in Buckley et al.
(2010a), Buckley & Leschen (2013), and Dunning et al. (2013).
Etymology. The specific name rakauwhakanekeneke was selected by the Ngatiwai people of Northland who
are kaitiaki (guardians) of the Poor Knights Islands. The name is formed from the words “rakau” and
“whakanekeneke” which translate from Māori as “stick” and “mobile” respectively.
Clitarchus tepaki Buckley, Myers & Bradler, sp. nov.
(Figs 1c, 2e, f, 3c, 4c, 5g, h, Tables 1, 2)
Official Māori name: Whē O Ngāti Kurī
Material examined. Holotype. ♂ (NZAC) [NEW ZEALAND ND Kohuronaki Reserve 11 Dec 2008 T R
Buckley RL1412 beating kanuka 35 9.853 S 173 16.332E] [NZAC04097536] [HOLOTYPE Clitarchus tepaki
Buckley, Myers & Bradler]
Paratypes. ♀, eggs (NZAC) [NEW ZEALAND, ND, North Cape Scientific Reserve, 34 24.467, 173 0.431, 18
Dec 2007, TR Buckley D Seldon R Hoare] [NZAC03005528] [PARATYPE Clitarchus tepaki Buckley, Myers &
Bradler]; ♀, eggs (NZAC) [NEW ZEALAND, ND, Kauri Bush Trk, Te Paki, 34 28.391, 172 45.842, 19 Dec 2007,
TR Buckley D Seldon R Hoare] [NZAC03005527] [PARATYPE Clitarchus tepaki Buckley, Myers & Bradler]; ♂
(NZAC) [NEW ZEALAND, ND, North Cape Scientific Reserve, 34 24.467, 173 0.431, 18 Dec 2007, TR Buckley
D Seldon R Hoare] [NZAC03004927] [PARATYPE Clitarchus tepaki Buckley, Myers & Bradler]; ♀ (NZAC)
[NEW ZEALAND, ND, North Cape Scientific Reserve, 34 24.467, 173 0.431, 18 Dec 2007, TR Buckley D Seldon
R Hoare] [NZAC03004931] [PARATYPE Clitarchus tepaki Buckley, Myers & Bradler]
Additional material examined. See Appendix 1.
Diagnosis. ♂: Row of 2–3 teeth on claspers, inner tooth massively enlarged with middle tooth and outer tooth
paired, tergum IX greater than twice the length of tergum X.
♀: Slender spines absent from dorsal surface, opercular organ ridge-like and flanked by sclerotized patch.
Egg: Egg width 2.2–2.7 mm.
Description. ♂ General: Medium-sized stick insect, moderately robust and apterous. Body green to mottled
brown, or dark grey. Most individuals without tubercles, or less commonly large tubercles or blunt, stout spines
present on dorsal surface. 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 unarmed or less commonly with
a few small tubercles. Lateral sutures present yet indistinct. Ocelli absent. Eyes prominent, hemispherical. Antenna
filiform with 18–20 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 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 over 3 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
Zootaxa 3900 (4) © 2014 Magnolia Press
·
467
REVISION OF CLITARCHUS
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 same length as 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 tergum II. Abdominal terga II–IX longer than wide, tergum IX 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 terga 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 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 posterior margin of tergum
X, rounded at end and approximately equal in length. Claspers formed by tergum X, with usually 3 but sometimes
2 or 4 dark tipped teeth. Inner tooth much larger than paired middle and outer 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, dorsal sclerite
only weakly sclerotized. Legs: Femora and tibia appearing pentagonal in cross section with setae running along
carinae. Setae increasing in density from femora to tibiae to tarsi and especially and numerous and long on
pretarsus and uncus. Fore coxa usually with single tubercle. Trochanters small, fused to femora. Fore femur
compressed basally and curved to accommodate head, with 4 short spines on latero-ventral carinae. Pair of apical
spines always present. Fore tibia approximately equal in length to fore femur and unarmed except for apical teeth.
Tarsomeres all shorter than femora and tibiae, and in decreasing order of 1
st
, 2
nd
, 3
rd
, 5
th
, 4
th
. Mid coxa 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
carinae. 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 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 spines. Lateral sutures
indistinct. Ocelli absent. Eyes prominent, hemispherical. Antenna filiform with 20–22 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 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. 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. Prosternum, prothoracic episternum and epimerum 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. Basisternite unarmed.
Mesosternum unarmed or less commonly with few tubercles. Wing remnants present. Metathorax + median
segment (1st abdominal segment) equal in size or marginally shorter than mesothorax. Metanotum unarmed and
with dark median stripe thicker at anterior and posterior margins. Methathoracic episternum elongated with
episternal suture usually with row of small tubercles. Metathoracic epimeron usually unarmed or with few very
small tubercles. Metasternum unarmed. Abdomen: Ten-segmented abdomen, unarmed. First abdominal segment
BUCKLEY ET AL.
468
·
Zootaxa 3900 (4) © 2014 Magnolia Press
(segmentum medianum) fused to metathorax, shorter than metanotum, boundary indistinct. Sternites unarmed.
Posterior margins of terga with dark strip medially, sometimes flanked by lighter patches and darker patches more
laterally. Terga II slightly shorter than III, 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
prominent, sclerotized ridge-like opercular organ at base, flanked by small lateral grooves or sclerotized patch. Tip
of operculum usually acute and not reaching of 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. Gonoplacs finger-like, with many
setae at tip, extending beyond other valves but not as far as anterior margin of terga 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 femur 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 tibia approximately same length as fore
femur 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 usually unarmed. Mid
femur with 0–3 teeth along ventral carina, 1–2 teeth along latero-ventral carina and dorsal carina unarmed. Pair of
apical spines present. Mid tibia occasionally with single tooth near proximal end of ventral carina 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, 1–3 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. 5g, h, Table 2). Slightly elongated along anterior-posterior axis, laterally flattened, opercular angle
close to zero. Egg width 2.2–2.7 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.
Ecology and distribution. This species has a disjunct distribution where it is found in the Te Paki / North
Cape area and also at the Paraawanui area of the Karikari Peninsula. Within the Te Paki / North Cape it has been
found in many locations, including Karatia, Kohuronaki, Spirits Bay, Tom Bowling Bay, Waikuku Flat, Unuwhao,
and North Cape Scientific Reserve. South of Te Paki along the Aupouri Peninsula it is replaced by C. hookeri. On
Karikari Peninsula it has so far only been found on the volcanic uplands of Paraawanui and elsewhere is replaced
by C. hookeri. Clitarchus tepaki sp. nov. has been observed feeding on Leptospermum scoparium, Kunzea spp.,
Muelhenbeckia australis, Metrosideros bartletii and M. perforata and the host plant range may be larger as it can
be successfully reared on Metrosideros excelsa. Leptospermum scoparium and Kunzea spp. are extremely common
throughout the Te Paki / North Cape and Karikari areas therefore large areas of suitable habitat exist. However, C.
tepaki sp. nov. is less common in the drier scrub and gumland areas, even when large amounts of Leptospermum
scoparium and Kunzea spp. are present, and is more common within and at the margins of forest remnants such as
Kauri Bush (Radar Bush) and Kohuronaki. The species appears to be abundant despite the presence of mammalian
predators and vespid wasps. It was given a Threat listing of “Range Restricted” by Buckley et al. (2012).
Information on phylogeographic relationships are given in Buckley et al. (2010a), Buckley & Leschen (2013), and
Dunning et al. (2013).
Etymology. The official Māori name, Whē O Ngāti Kurī, was selected by the Ngāti Kurī people of Northland
whose rohe (area of interest) includes the Te Paki and North Cape area. The specific name refers to the Te Paki area
that includes the type locality and is a well known area of endemism for many invertebrate groups (see below).
Zootaxa 3900 (4) © 2014 Magnolia Press
·
469
REVISION OF CLITARCHUS
Identification keys
Key to the males of Clitarchus
(1) Claspers with 2–4 teeth, inner tooth massively enlarged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(2)
- Claspers with 3–5 teeth of approximately equal size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hookeri
(2) Tergum IX less than twice the length of tergum X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rakauwhakanekeneke
- Tergum IX greater than twice the length of tergum X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. tepaki
Key to the females of Clitarchus
(1) Sharp, slender spines absent from thoracic tergites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2)
- Sharp, slender spines present on thoracic tergites. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rakauwhakanekeneke
(2) Opercular organ consisting of sclerotized ridge and flanked by small lateral sclerotized patches with grooves . . . . . . .C. tepaki
- Opercular organ only very weakly sclerotized and not flanked by lateral sclerotized patches . . . . . . . . . . . . . . . . . . C. hookeri
Discussion
Classification and morphological characters. Among the New Zealand stick insects, the Acanthoxylini is
diagnosed by having antennae longer than the fore femora (Bradley & Galil 1977). The remaining genera,
traditionally placed in the polyphyletic Pachymorphinae (Buckley et al. 2009), have shorter antennae and this
character was previously used to justify assignment of the New Zealand genera to either Phasmatinae or
Pachymorphinae (e.g., Günther 1953). Although antennal length varies across the Phasmatodea and is clearly a
poor character for defining higher level taxa (Bradler 2009), among the New Zealand species, it can be used to
delimit the Acanthoxylini from the remaining genera.
In general, male postgenital claspers of stick insects are characterized by the presence of a movable sclerotised
vomer on the venter of abdominal segment X and a pair of thorn pads at the hind margin of tergum X (Bradler
2009; Bradler et al. 2014; Gottardo & Vallotto 2014). In Lanceocercata, which also include the Acanthoxylini, the
vomer is reduced and the specialised tergum X serves as clasper alone, often with thorn pads transformed into few
distinctive spines or teeth (Bradler 2009; Buckley et al. 2010b). Within the Acanthoxylini Pseudoclitarchus
Salmon is diagnosed by having a single tooth at the apex of each clasper (Figs 2h, 3d). Clitarchus has at least three
teeth on each clasper (Figs 2b, 3a). Clitarchus rakauwhakanekeneke sp. nov. has three prominent teeth with the
basal tooth enlarged, and the median and apical teeth usually paired (Figs 2d, 3b). Clitarchus tepaki sp. nov. also
has three teeth, with the basal tooth massively enlarged and usually well separated from the two paired apical teeth,
more so than in C. rakauwhakanekeneke sp. nov. (Figs 2f, 3c). The large basal tooth is a character state also shared
with Tepakiphasma ngatikuri Buckley and Bradler (Figs 2j, 3d; Buckley & Bradler 2010); however this species
only ever has only two teeth with the smaller tooth at the clasper apex. Because males of Acanthoxyla Uvarov are
unknown (Salmon 1955, 1991; Buckley et al. 2008; Morgan-Richards & Trewick 2005) this character cannot be
examined in this genus. This pattern suggests that the enlarged basal tooth may be apomorphic for a clade
comprising Clitarchus, Acanthoxyla, Pseudoclitarchus and Tepakiphasma (Fig. 6). The clasper structure of
Argosarchus is highly divergent from the other New Zealand Acanthoxylini with a cluster of spine-like teeth at the
apex (cf. Fig. 6; Salmon 1991), consistent with its sister-group relationship to Acanthoxyla, Tepakiphasma,
Pseudoclitarchus and Clitarchus (Buckley et al. 2010b; Dunning et al. 2013). We infer that the enlarged basal tooth
is plesiomorphic among Clitarchus spp. and was secondarily lost or reduced in males of C. hookeri (Figs 2, 3, 6).
All of the New Zealand genera, with the exception of Spinotectarchus Salmon, have a carina, tubercle or spine
at the base of the operculum (e.g., Fig. 4), where the male clasps the female during copulation (Buckley et al.
2010b). We have termed this structure the opercular organ (Buckley et al. 2010b) and it is most prominent in
Acanthoxyla, where in some species it forms a long, posteriorly curved spine (Salmon 1955, 1991). This character
appears to be unique within the Phasmatodea, where it is likely an apomophy for the main New Zealand clade.
Species from the Platycraninae also have a sclerotized structure at the base of the operculum, but it runs laterally
along the anterior margin of the operculum rather than anteriorly-posteriorly. This difference in structure along
with the lack of any phylogenetic relationship between the Platycraninae and the New Zealand species (Buckley et
BUCKLEY ET AL.
470
·
Zootaxa 3900 (4) © 2014 Magnolia Press
al. 2009; 2010b) suggests an evolutionary convergence. In Clitarchus, the opercular organ appears as a tubercle or
sclerotized ridge, with smaller, lateral lines of sclerotization in C. tepaki sp. nov. (Fig. 4c) and C.
rakauwhakanekeneke sp. nov. (Fig. 4b). The opercular organ in many C. tepaki individuals is similar to that of
Acanthoxyla inermis Salmon, but in the latter it clearly forms a spine-like structure (Fig. 4f; see, also, Salmon
1991).
Species of Clitarchus have traditionally been differentiated on the basis of colouration and degree of body
tuberculation (e.g., Colenso 1885; Brunner 1907; Salmon 1991). All three species of Clitarchus display
considerable levels morphological variation. This variation is expressed in terms of body colour, presence, size and
abundance of tubercles on the body, the presence of carinae and spination on the legs and the presence of
abdominal flanges. Therefore, these characters are poor indicators of species boundaries. Superior resolution of
species differentiation can be obtained by the shape and size of the claspers and the arrangement of teeth on the
claspers. Females can be differentiated using the relative size of the cerci and shape of the opercular organ.
FIGURE 6. Phylogeny of Acanthoxylini following Buckley et al. (2010b) and Dunning et al. (2013), illustrating the evolution
of male clasper morphology. The massively enlarged basal tooth evolved in the last common ancestor of Tepakiphasma,
Acanthoxyla, Pseudoclitarchus and Clitarchus (synapomorphy) and was reduced in C. hookeri. The male of Acanthoxyla is not
known.
Biogeography and conservation status. The North Cape region was isolated from the area of Northland south
of the Aupouri Peninsula for much of the Pliocene (Ballance and William 1982). This isolation has facilitated
allopatric speciation of many endemic species of plant (e.g., Davidson et al. 1969; Druce et al. 1979; Gardner &
Bartlett 1980; de Lange et al. 2003; von Konrat & Braggins 2005), vertebrate (e.g., Chapple et al. 2008) and
invertebrate species (e.g., Holloway 1961; Gardner 1967; Goulstone et al. 1993; Larochelle & Larivière 2005;
Marshall & Barker 2007; Winterbourn 2009; Hoare 2010; Seldon & Leschen 2011; Vink et al. 2011; Buckley &
Zootaxa 3900 (4) © 2014 Magnolia Press
·
471
REVISION OF CLITARCHUS
Leschen 2013), including genetically divergent clades (e.g., Buckley & Leschen 2013; Morgan-Richards et al.
2001). The invertebrate fauna of this area also includes the recently described phasmid Tepakiphasma ngatikuri
(Buckley & Bradler 2010), and so the present revision brings the number of newly described stick insects from this
small geographic area to two. The Karikari Peninsula was also isolated during the Pliocene (Ballance and William
1982), but has fewer endemic species, possibly due to more extensive habitat degradation and the near absence of
forest. Although, C. tepaki does not appear to be threatened due to the presence of multiple populations across the
Te Paki / North Cape area and Karikari Peninsula, the presence of yet another endemic species in northern New
Zealand reinforces the biological uniqueness of this area and supports the continuation of ongoing conservation
efforts.
The Poor Knights Islands are remnants of a volcanic system that dates back to the Late Miocene (Hayward
1991). Unlike many New Zealand offshore islands they were isolated from the mainland during the Last Glacial
Maximum and possibly for as long as 2 million years (Hayward 1991). This isolation has allowed many endemic
species of invertebrates to evolve (Watt 1982), although there is only one endemic plant taxon (de Lange and
Cameron 1999). As with the Te Paki / North Cape area, the presence of another endemic invertebrate species
supports conservation and biosecurity efforts to preserve the biota of these islands.
Acknowledgements
We acknowledge Ngatiwai for supporting our research on the Poor Knights Islands, in particular Hori Parata, Clive
Stone and Tui Shortland. Access to Te Paki and North Cape sites was facilitated by Ngāti Kurī. Other iwi assisted
with access to sites including; Te Rarawa, Te Aupōuri, Ngāi Takoto, Ngāti Rehia, Ngāti Kuta, Patukeha, Ngāti
Kawau, Ngāti Kura, Ngai Toro, Ngāti Rahiri, Ngāti Whātua, Te Roroa, Ngāti Kahu, Te Uri-O-Hau, and Ngāpuhi.
Special thanks to Dave Seldon for collecting the specimen that first indicated C. tepaki was a new species. We
thank Dave Seldon, Robert Hoare, Rich Leschen, Christina Painting, Megan Iminitoff, Jamie Stavert, Elizabeth
Cunningham and Christina Rowe for help in the field. Leonie Clunie and Grace Hall provided technical assistance
and Birgit Rhode took the automontage images. Tony Jewell provided much useful discussion on New Zealand
stick insect biology and taxonomy. Thanks to Robert Hoare, Rich Leschen and three anonymous reviewers for
comments on the manuscript. The Department of Conservation issued collecting permits and helped with field
work, in particular Lester Bridson, Patrick Whaley, Andrea Booth, Bryce Lummis, Gail Townsend, Donna
Stuthridge, Janeen Collings, Carolyn Smith and Ian Stringer. Access to collections and loan of specimens was
provided by Ricardo Palma (MONZ) Judith Marshall (BMNH), Susanne Randolf and Ulrike Aspöck (NHMW).
Specimens were collected under permits issued by the Department of Conservation (NO-21083-FAU, NO-23262-
FAU and NO-15555-FAU). This work was funded by grants from the Royal Society of New Zealand Marsden
Fund (LCR302, LCR0502 and LCR0901), the Allan Wilson Centre, the University of Auckland and Core funding
for Crown Research Institutes from the Ministry of Business, Innovation and Employment’s Science and
Innovation Group.
References
Ballance, P.F. & Williams, P.W. (1982) The geomorphology of Auckland and Northland. In: Soons, J.M. & Selby, M.J. (Eds.),
Landforms of New Zealand. Longman Paul; Hong Kong, pp. 210–232.
Beier, M. (1968) Phasmida (Stab- oder Gespenstheuschrecken). In: Helmcke, J.-G., Starck, D., & Wermuth), Handbuch der
Zoologie IV. Walter De Gruyter & Co, Berlin, Germany, pp. 1–56.
Bradler, S. (2001) The Australian stick insects, a monophyletic group within the Phasmatodea? Zoology, 104 (Supplement III),
pp. 69.
Bradler, S. (2009) Phylogeny of the stick and leaf insects (Phasmatodea). Species, Phylogeny and Evolution, 2, 3–139.
Bradler, S., Robertson, J.A. & Whiting, M.F. (2014) A molecular phylogeny of Phasmatodea with emphasis on Necrosciinae,
the most species-rich subfamily of stick insects. Systematic Entomology, 39, 205–222.
http://dx.doi.org/10.1111/syen.12055
Bradley, J.C. & Galil, B.S. (1977) The taxonomic arrangement of the Phasmatodea with keys to the subfamilies and tribes.
Proceedings of the Entomological Society of Washington, 79, 176–208.
Brock, P.D. (1987) A third New Zealand stick insect (Phasmatodea) established in the British Isles, with notes on the other
BUCKLEY ET AL.
472
·
Zootaxa 3900 (4) © 2014 Magnolia Press
species, including a correction. In Mazzini, M., Scali, V. (Eds). 1
st
International Symposium on Stick Insects. University of
Siena, pp. 125–132.
Brock, P.D. (1997) Taxonomic changes relating to New Zealand stick insects. Phasmid Studies, 6, 21–23.
Brunner von Wattenwyl, K. (1907) Die Insektenfamilie der Phasmiden. Phasmidae Anareolatae (Clitumnini, Lonchodini,
Bacunculini). Wilhelm Engelmann, Leipzig, Germany, pp. 181–340.
Buckley, T.R., Attanayake, D., Park, D.-C., Ravindran, S., Jewell, T.R. & Normark, B.B. (2008) Investigating hybridization in
the parthenogenetic New Zealand stick insect Acanthoxyla (Phasmatodea) using single-copy nuclear loci. Molecular
Phylogenetics and Evolution, 47, 335–349.
http://dx.doi.org/10.1016/j.ympev.2008.02.016
Buckley, T.R., 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, B, 276, 1055–1062.
http://dx.doi.org/10.1098/rspb.2008.1552
Buckley, T.R., 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, 37, 682–695.
http://dx.doi.org/10.1111/j.1365-2699.2009.02239.x
Buckley, T.R., Attanayake, D., Nylander, J.A.A. & Bradler, S. (2010b) The phylogenetic placement and biogeographical
origins of the New Zealand stick insects (Phasmatodea). Systematic Entomology, 35, 207–225.
http://dx.doi.org/10.1111/j.1365-3113.2009.00505.x
Buckley, T.R. & Bradler, S. (2010) Tepakiphasma ngatikuri, a new genus and species of stick insect (Phasmatodea) from the
Far North of New Zealand. New Zealand Entomologist, 33, 118–126.
http://dx.doi.org/10.1080/00779962.2010.9722200
Buckley, T.R., Palma, R.L., Johns, P.M., Gleeson, D.M., Heath, C.G., Hitchmough, R.A. & Stringer, I.A.N. (2012) The
conservation status of small or less well known groups of New Zealand terrestrial invertebrates. New Zealand
Entomologist, 35, 137–143.
http://dx.doi.org/10.1080/00779962.2012.686319
Buckley, T.R. & Leschen, R.A.B. (2013) Comparative phylogenetic analysis reveals long term isolation of lineages on the
Three Kings Islands, New Zealand. Biological Journal of Linnean Society, 108, 361–377.
http://dx.doi.org/10.1111/j.1095-8312.2012.02009.x
Chapple, D.G., Patterson, G.B., Bell, T. & Daugherty, C.H. (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.
http://dx.doi.org/10.1670/07-110.1
Clark Sellick, J.T. (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.
http://dx.doi.org/10.1046/j.1365-3113.1997.d01-30.x
Clark Sellick, J.T. (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
Colenso, W. (1885) A description of some newly discovered New Zealand insects believed to be new to science. Transactions
of the New Zealand Institute, 17, 151–155.
Crosby, T.K., Dugdale, J.S. & Watt, J.C. (1998) Area codes for recording specimen localities in the New Zealand subregion.
New Zealand of Zoology, 25, 175–183.
http://dx.doi.org/10.1080/03014223.1998.9518148
Davidson, J., Grant-Mackie, J.A., Morton, J.E. & Rattenbury, J.A. (1969) North Cape: a scientific case for conservation. Tane ,
15, 5–11.
de Lange, P. & Cameron, E.K. (1999) The vascular flora of Aorangi Island, Poor Knights Islands, northern New Zealand. New
Zealand Journal of Botany, 37, 433–468.
http://dx.doi.org/10.1080/0028825X.1999.9512646
de Lange, P.J., Heenan, P.B. & Dawson, M.I. (2003) A new species of Leucopogon (Ericaceae) from the Surville Cliffs, North
Cape, New Zealand. New Zealand Journal of Botany, 41, 13–21.
http://dx.doi.org/10.1080/0028825X.2003.9512829
Druce, A.P., Bartlett, J.K. & Gardner, R.O. (1979) Indigenous vascular plants of the serpentine area of Surville Cliffs and
adjacent cliff tops, northwest of North Cape, New Zealand. Tan e , 25, 187–206.
Dunning, L.T., Thomson, G., Dennis, A.B., Sinclair, B.J., Newcomb, R.D. & Buckley, T.R. (2013) Positive selection in
glycolysis among Australasian stick insects. BMC Evolutionary Biology, 13, 1–215.
http://dx.doi.org/10.1186/1471-2148-13-215
Gardner, N.W. (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, R.O. & Bartlett, J.K. (1980) Forest flora of the North Cape region. Tan e , 26, 223–234.
Gottardo, M. & Vallotto, D. (2014) External macro- and micromorphology of the male of the stick insect Hermarchus leytensis
(Insecta: Phasmatodea) with phylogenetic considerations. Comptes Rendus Biologies, 337, 258–268.
Zootaxa 3900 (4) © 2014 Magnolia Press
·
473
REVISION OF CLITARCHUS
http://dx.doi.org/10.1016/j.crvi.2014.02.005
Goulstone, J.F., Mayhill, P.C. & Parish, G.R. (1993) An illustrated guide to the land Mollusca of the Te Paki Ecological Region,
New Zealand. Ta n e , 34, 1–32.
Günther, K. (1953) Über die taxonomische Gliederung und geographische Verbreitung der Insektenordnung der Phasmatodea.
Beiträge zur Entomologie, 3, 541–563.
Hayward, B.W. (1991) Geology and geomorphology of the Poor Knights Islands, Northern New Zealand. Tane , 33, 23–37.
Hennemann, F.H., Conle, O.V. & Zhang, W. (2008) Catalogue of the stick and leaf-insects (Phasmatodea) of China, with a
faunistic analysis, review of recent ecological and biological studies and bibliography (Insects: Orthoptera: Phasmatodea).
Zootaxa, 1735, 1–77.
Hoare, R.J.B. (2010) Izatha (Insecta: Lepidoptera: Gelechioidea: Oecophoridae). Fauna of New Zealand. Vol. 65. Manaaki
Whenua Press, Lincoln, New Zealand, 201 pp.
Holloway, B.A. (1961) A systematic revision of the New Zealand Lucanidae (Insecta: Coleoptera). Dominion Museum Bulletin,
20, 1–139.
Hutton, F.W. (1898) The Phasmidae of New Zealand. Transactions of the New Zealand Institute, 30, 160–166.
Hutton, F.W. (1899) Revision of New Zealand Phasmidae. Transactions of the New Zealand Institute, 31, 55–59.
Jewell, T. & Brock, P.D. (2002) A review of the New Zealand stick insects: new genera and synonymy, keys, and a catalogue.
Journal of Orthoptera Research, 11, 189–197.
http://dx.doi.org/10.1665/1082-6467(2002)011[0189:AROTNZ]2.0.CO;2
Kirby, W.F. (1904) A synonymic catalogue of Orthoptera. Vol. 1. Longmans & Co., London, 501 pp.
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.
http://dx.doi.org/10.1016/j.ode.2005.08.004
Larochelle, A. & Larivière, M.C. (2005) Harpalini (Insecta: Coleoptera: Carabidae: Harpalinae). Fauna of New Zealand. Vol.
53. Manaaki Whenua Press, Lincoln, New Zealand, 160 pp.
Littig, K.S. (1942) External anatomy of the Florida walking stick Anisomorpha buprestoides Stoll. The Florida Entomologist,
3, 33–41.
http://dx.doi.org/10.2307/3492435
Marshall, B.A. & Barker, G.M. (2007) A revision of New Zealand landsnails of the genus Cytora Kobelt & Mollendorff, 1897
(Mollusca: Gastropoda: Pupinidae). Tuhinga, 18, 49–113.
Morgan-Richards, M., Trewick, S.A. & Wallis, G.P. (2001) Chromosome races with Pliocene origins: evidence from mtDNA.
Heredity, 86, 303–312.
http://dx.doi.org/10.1046/j.1365-2540.2001.00828.x
Morgan-Richards, M. & Trewick, S.A. (2005) Hybrid origin of a parthenogenetic genus? Molecular Ecology, 14, 2133–2142.
http://dx.doi.org/10.1111/j.1365-294X.2005.02575.x
Morgan-Richards, M., Trewick, S.A. & Stringer, I.A. (2010) Geographic parthenogenesis and the common tea-tree stick insect
of New Zealand. Molecular Ecology, 19, 1227–1238.
http://dx.doi.org/10.1111/j.1365-294X.2010.04542.x
Myers, S.S., Trewick, S.A. & Morgan-Richards, M. (2013) Multiple lines of evidence suggest mosaic polyploidy in the hybrid
parthenogenetic stick insect lineage Acanthoxyla. Insect Conservation and Diversity, 6, 537–548.
http://dx.doi.org/10.1111/icad.12008
Otte, D. & Brock, P.D. (2005) Phasmida Species File: A Catalog to the Stick Insects of the World (The Orthopterists Society),
504 pp.
Ragge, D.R. (1965) Grashoppers, crickets and cockroaches of the British Isles. Wayside and Woodland series, Warne, London,
299 pp.
Salmon, J.T. (1955) The genus Acanthoxyla (Phasmidae). Transactions of the Royal Society of New Zealand, 82, 1149–1156.
Salmon, J.T. (1991) The Stick Insects of New Zealand. Reed, Auckland, New Zealand. 124 pp.
Seldon, D.S. & Leschen, R.A.B. (2011) Revision of the Mecodema curvidens species complex (Coleoptera, Carabidae,
Broscini). Zootaxa, 2829, 1–45.
Stål, C. (1875) Recensio Orthopterorum. Revue critique des Orthoptères décrits par Linné, de Geer et Thunberg. Vol. 3. P.A.
Norstedt & Söner, Stockholm, 105 pp.
Stringer, I.A.N. (1969) Blastokinesis and embryology of the phasmid Clitarchus hookeri. Tane, 15, 41–52.
Stringer, I.A.N. (1970) The nymphal and imaginal stages of the bisexual stick insect Clitarchus hookeri. New Zealand
Entomologist, 4, 85–95.
http://dx.doi.org/10.1080/00779962.1970.9722927
Trewick, S.A., Goldberg, J. & Morgan-Richards, M. (2005) Fewer species of Argosarchus and Clitarchus stick insects
(Phasmida: Phasmatinae): evidence from mitochondrial and nuclear DNA sequences. Zoologica Scripta, 34, 483–491.
http://dx.doi.org/10.1111/j.1463-6409.2005.00204.x
Trewick, S.A., Morgan-Richards, M. & Collins, L.J. (2008) Are you my mother? Phylogenetic analysis reveals orphan hybrid
stick insect genus is part of a monophyletic New Zealand clade. Molecular Phylogenetics and Evolution, 48, 799–808.
http://dx.doi.org/10.1016/j.ympev.2008.05.025
Vink, C.J., Fitzgerald, B.M., Sirvid, P.J. & Dupérré, N. (2011) Reuniting males and females: redescriptions of Nuisiana arboris
BUCKLEY ET AL.
474
·
Zootaxa 3900 (4) © 2014 Magnolia Press
(Marples 1959) and Cambridgea reinga Forster & Wilton 1973 (Araneae: Desidae, Stiphidiidae). Zootaxa, 2739, 41–50.
von Konrat, M.J. & Braggins, J.E. (2005) Frullania wairua, a new and seemingly rare liverwort species from Northland, New
Zealand. New Zealand Journal of Botany, 43, 885–893.
http://dx.doi.org/10.1080/0028825X.2005.9512998
Watt, J.C. (1982) Terrestrial arthropods from the Poor Knights Islands, New Zealand. Journal of the Royal Society of New
Zealand, 12, 283–320.
http://dx.doi.org/10.1080/03036758.1982.10415350
White, A. (1846) The Zoology of the voyage of H.M.S. Erebus and Terror, 1. Insects of New Zealand. E.W. Janson, London,
United Kingdom, 27 pp.
Winterbourn, M.J. (2009) A new genus and species of Leptophlebiidae (Ephemeroptera) from northern New Zealand. New
Zealand Journal of Zoology, 36, 423–430.
http://dx.doi.org/10.1080/03014223.2009.9651475
APPENDIX 1. Material examined.
Specimen data follows the following format: Crosby et al. (1998) area code, number of individuals, gender / eggs, locality,
NZAC accession code, latitude, longitude, date of collection, collectors.
Clitarchus hookeri
ND, 1, Herekino Nth Head, Rangikohu Rd, far end, NZAC03009381, 23 Dec 2010, D S Seldon; ND, 1, East Herekino,
Kaitaia Walkway, NZAC03009469, 35 9.803, 173 16.197, 16 Jan 2010, TR Buckley D Seldon R Hoare; ND, eggs, East
Herekino, Kaitaia Walkway, NZAC03014415, 35 9.803, 173 16.197, 16 Jan 2010, TR Buckley D Seldon R Hoare; ND, 1,
Paihia, Opua SF, NZAC03009447, 35 17.312, 174 05.220, 17 Mar 2011, R Leschen N Lord; ND, 1, ♀, Whakaangi,
NZAC03006378, 34 56.863, 173 32.684, 18 Jan 2010, TR Buckley D Seldon R Hoare; ND, 1, ♀, Gum Hole Reserve,
NZAC03006389, 34 58.024, 173 22.828, 17 Jan 2010, TR Buckley D Seldon R Hoare; ND, 1, ♀, Paranui Scenic Reserve,
NZAC03006384, 35 4.122, 173 26.441, 17 Jan 2010, TR Buckley D Seldon ; ND, 1, ♀, Junction of SH10 and Takou Bay Rd, 2
km N of Te Whau, Kerikeri, NZAC03000090, 35 8.211, 173 52.775, 23 Mar 2005, K Hill D Marshall; ND, 1, Junction of SH10
and Takou Bay Rd, 2 km N of Te Whau, Kerikeri, NZAC03000057, 35 8.211, 173 52.775, 23 Mar 2005, K Hill D Marshall;
ND, 1, ♀, Mangamuka Gorge Scenic Reserve, roadside, NZAC03000323, 35 9.486, 173 25.521, 6 Feb 2006, TR Buckley R
Hoare; ND, 1, ♂, Mangamuka Gorge Scenic Reserve, roadside, NZAC03000500, 35 9.486, 173 25.521, 6 Feb 2006, TR
Buckley R Hoare; ND, 1, ♂, Mangamuka Gorge Scenic Reserve, roadside, NZAC03000125, 35 9.486, 173 25.521, 6 Feb 2006,
TR Buckley R Hoare; ND, 1, ♂, Lake Ngatu, NZAC03005489, 35 1.622, 173 11.935, 12 Dec 2008, TR Buckley R Leschen D
Seldon; ND, 1, ♀, Lake Ngatu, NZAC03005458, 35 1.622, 173 11.935, 12 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1,
♂, Lake Ngatu, NZAC03005460, 35 1.622, 173 11.935, 12 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1, ♂, Lake Ngatu,
NZAC03005463, 35 1.622, 173 11.935, 12 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1, ♂, Lake Ngatu,
NZAC03005472, 35 1.622, 173 11.935, 12 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1, ♂, Lake Ngatu,
NZAC03005468, 35 1.622, 173 11.935, 12 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1, ♀, Lake Ngatu,
NZAC03005475, 35 1.622, 173 11.935, 12 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1, ♀, Lake Ngatu,
NZAC03005490, 35 1.622, 173 11.935, 12 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1, ♂, Lake Ngatu, Kaitaia,
NZAC04030718, 5 Jan 1963, E S Gourlay; ND, 1, ♀, eggs, East Herekino, Kaitaia Walkway, NZAC03005572, 35 9.803, 173
16.197, 14 Jan 2010, TR Buckley D Seldon R Hoare; ND, 1, Ahipara, road to Gumlands, NZAC03000129, 35 11.069, 173
7.314, 6 Feb 2006, TR Buckley R Hoare; ND, 1, ♂, Ahipara, road to Gumlands, NZAC03000351, 35 11.069, 173 7.314, 6 Feb
2006, TR Buckley R Hoare; ND, 1, ♂, Ahipara, road to Gumlands, NZAC03000396, 35 11.069, 173 7.314, 6 Feb 2006, TR
Buckley R Hoare; ND, 1, Ahipara, Reef Point, NZAC03005280, 10 Feb 1988, G Messenger; ND, 2, Ahipara, Herekino Bush,
NZAC03005319, Jan 1987, G Messenger; ND, 1, ♀, Herekino Gorge, Kaitaia Awaroa Rd, NZAC03000386, 35 12.3000, 173
11.015, 7 Feb 2006, TR Buckley R Hoare; ND, 1, Herekino Gorge, Kaitaia Awaroa Rd, NZAC03000359, 35 12.3000, 173
11.015, 7 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, Herekino Forest Track, Herekino Forest, NZAC03000127, 35 12.587, 173
11.650, 7 Feb 2006, TR Buckley R Hoare; ND, 1, ♂, Herekino Forest Track, Herekino Forest, NZAC03000113, 35 12.587, 173
11.650, 7 Feb 2006, TR Buckley R Hoare; ND, 1, ♂, Opua Forest, Oromahoe Rd, NZAC04028289, 35 19.251, 174 03.936, 14
Dec 2006, K Hill, D Marshall; ND, 1, ♂, Opua Forest, Oromahoe Rd, NZAC04028293, 35 19.251, 174 03.936, 14 Dec 2006, K
Hill, D Marshall; ND, 1, ♂, Opua Forest, Oromahoe Rd, NZAC04028229, 35 19.251, 174 03.936, 14 Dec 2006, K Hill, D
Marshall; ND, 1, ♀, Opua Forest, Oromahoe Rd, NZAC04028313, 35 19.251, 174 03.936, 14 Dec 2006, K Hill, D Marshall;
ND, 1, ♂, Pawarenga, Warawara Forest, NZAC03004868, 35 21.497, 173 15.044, 15 Dec 2007, TR Buckley D Seldon R
Hoare; ND, 1, ♂, Pawarenga, Warawara Forest, NZAC03004783, 35 21.497, 173 15.044, 15 Dec 2007, TR Buckley D Seldon
R Hoare; ND, 1, ♂, Pawarenga, Warawara Forest, NZAC03004907, 35 21.497, 173 15.044, 15 Dec 2007, TR Buckley D
Seldon R Hoare; ND, 1, ♂, Pawarenga, Warawara Forest, NZAC03005009, 35 21.497, 173 15.044, 15 Dec 2007, TR Buckley
D Seldon R Hoare; ND, 1, Pawarenga, Warawara Forest, NZAC03005021, 35 21.497, 173 15.044, 15 Dec 2007, TR Buckley D
Seldon R Hoare; ND, 1, Eggs, Warawara Forest, NZAC03004871, 35 22.159, 173 17.002, 15 Dec 2007, TR Buckley D Seldon
R Hoare; ND, 1, ♀, Warawara Forest, NZAC03004982, 35 22.159, 173 17.002, 15 Dec 2007, TR Buckley D Seldon R Hoare;
ND, 1, ♀, eggs, Whakaruangangana gumland, ca 3km SW of Kaikohe, NZAC03005530, 35 26.67, 173 45.31, 3 Feb 2007, R
Zootaxa 3900 (4) © 2014 Magnolia Press
·
475
REVISION OF CLITARCHUS
Hoare; ND, 1, Corner Signal Station Rd & SH12, Pakia rest area, Omapere, NZAC03000117, 35 32.265, 173 22.951, 5 Feb
2006, TR Buckley R Hoare; ND, 1, ♀, Corner Signal Station Rd & SH12, Pakia rest area, Omapere, NZAC03000116, 35
32.265, 173 22.951, 5 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, Corner Signal Station Rd & SH12, Pakia rest area, Omapere,
NZAC03000348, 35 32.265, 173 22.951, 5 Feb 2006, TR Buckley R Hoare; ND, 1, ♂, Mangamuka Gorge Scenic Reserve,
roadside, NZAC03000370, 35 9.486, 173 25.521, 6 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, Mangamuka Gorge Scenic
Reserve, roadside, NZAC03000363, 35 9.486, 173 25.521, 6 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, eggs, Herekino East,
NZAC03005547, 35 9.853, 173 16.332, 12 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1, ♂, Herekino East,
NZAC03005264, 35 9.853, 173 16.332, 12 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1, ♂, Herekino East,
NZAC03005317, 35 9.853, 173 16.332, 12 Dec 2008, TR Buckley R Leschen D Seldon; ND, 5, Waipoua Forest,
NZAC03005367, Mar 1973; ND, 1, ♀, Waipoua, NZAC04030748, Mar 1923, Milligan; ND, 1, ♂, Waipoua, NZAC04030772,
Mar 1923, Milligan; ND, 1, Fanal I, Mokohinau Is, NZAC03005289, 18 Mar 2008, I A Stringer; ND, 1, Omahuta State Forest,
NZAC03005330, Oct 1974, J S Dugdale; ND, 1, ♀, Roadside SH12, 2 km N of Babylon Coast Rd, Dargaville,
NZAC03000369, 35 53.758, 173 48.367, 6 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, Roadside SH12, 2 km N of Babylon
Coast Rd, Dargaville, NZAC03000366, 35 53.758, 173 48.367, 6 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, Trounson Kauri
Park, NZAC03000344, 35 43.228, 173 38.960, 4 Feb 2006, TR Buckley R Hoare; ND, 1, ♂, Trounson Kauri Park,
NZAC03000346, 35 43.228, 173 38.960, 4 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, Trounson Kauri Park, NZAC03000345,
35 43.228, 173 38.960, 4 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, Trounson Kauri Park, NZAC03000336, 35 43.228, 173
38.960, 4 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, Trounson Kauri Park, NZAC03000330, 35 43.228, 173 38.960, 4 Feb
2006, TR Buckley R Hoare; ND, 1, ♂, Trounson Kauri Park, NZAC03000342, 35 43.228, 173 38.960, 4 Feb 2006, TR Buckley
R Hoare; ND, 1, ♀, Trounson Kauri Park, NZAC03000383, 35 43.228, 173 38.960, 4 Feb 2006, TR Buckley R Hoare; ND, 1,
Trounson Kauri Park, NZAC03000416, 35 43.228, 173 38.960, 4 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, Trounson Kauri
Park, NZAC03000021, 35 43.228, 173 38.960, 4 Feb 2006, TR Buckley R Hoare; ND, 1, ♀, Roadside, Cove Rd, Robert Hastie
Reserve, NZAC03000457, 36 3.616, 174 34.358, 27 Apr 2006, TR Buckley D Seldon; ND, 1, ♀, Maitahi Wetland Scientific
Reserve, Maitahi Rd, NZAC03004952, 35 52.01, 173 44.15, 7 Feb 2007, R Hoare; ND, 1, ♀, Memorial Park, Dargaville,
NZAC03000352, 35 55.360, 173 51.263, 5 Feb 2006; ND, 1, ♀, Brynderwyn, NZAC03000066, 36 4.327, 174 43.544, 7 Feb
2006, TR Buckley R Hoare; ND, 1, ♂, Brynderwyn, NZAC03000040, 36 4.327, 174 34.544, 7 Feb 2006, TR Buckley R Hoare;
ND, 1, ♀, Brynderwyn, NZAC03000016, 36 4.327, 174 34.544, 7 Feb 2006, TR Buckley R Hoare; ND, 1, ♂, Brynderwyn,
NZAC03000094, 36 4.327, 174 34.544, 7 Feb 2006, TR Buckley R Hoare; ND, 1, ♂, Brynderwyn, NZAC03000074, 36 4.327,
174 34.544, 7 Feb 2006, TR Buckley R Hoare; ND, 1, Tangihua Forest lodge, NZAC03009433, 35 53 06.7 , 174 07 34.5, 6 Dec
2012, TR Buckley D Seldon; ND, 1, Tangihua Forest lodge, NZAC03009370, 35 53 06.7 , 174 07 34.5, 6 Dec 2012, TR
Buckley D Seldon; ND, 1, Tangihua Forest lodge, NZAC03009475, 35 53 06.7 , 174 07 34.5, 6 Dec 2012, TR Buckley D
Seldon; ND, 1, Tangihua Forest lodge, NZAC03009451, 35 53 06.7 , 174 07 34.5, 6 Dec 2012, TR Buckley D Seldon; ND, 1,
Tangihua Forest lodge, NZAC03009415, 35 53 06.7 , 174 07 34.5, 6 Dec 2012, TR Buckley D Seldon; ND, 1, Tangihua Forest
lodge, NZAC03009367, 35 53 06.7 , 174 07 34.5, 6 Dec 2012, TR Buckley D Seldon; ND, 2, Whale Bay, NZAC03009517, 35
33.750, 174 29.967, 16 Dec 2009, TR Buckley R Leschen D Seldon; ND, 1, Omapere, NZAC03009513, 35.538937, 173
385054, 6 Apr 2010, C Carlton R Leschen; ND, 2, Brattys Bush Scenic Res, Tutukaka, NZAC03009425, 35 36.739, 174
29.106, 16 Dec 2009, TR Buckley R Leschen D Seldon; ND, 1, Bream Head, Peach Cove Trk, Whangarei Heads,
NZAC03009490, 7 Dec 2010, D Seldon; ND, 1, Bream Head, Peach Cove Trk, Whangarei Heads, NZAC03009403, 7 Dec
2010, D Seldon; ND, 1, Bream Head, Peach Cove Trk, Whangarei Heads, NZAC03009359, 7 Dec 2010, D Seldon; ND, 1,
Bream Head, Peach Cove Trk, Whangarei Heads, NZAC03009358, 7 Dec 2010, D Seldon; ND, 1, Bream Head, Peach Cove
Trk, Whangarei Heads, NZAC03009401, 7 Dec 2010, D Seldon; ND, 1, Bream Head, Peach Cove Trk, Whangarei Heads,
NZAC03009491, 7 Dec 2010, D Seldon; ND, 1, Brynderwyn Trk, East side from Waipu Cove, NZAC03009402, 6 Dec 2010, D
S Seldon; ND, 1, ♀, Coppermine I, Hen and Chicken Is, track campsite to Lighthouse, NZAC03000027, 35 53.196, 174
46.698, 3 Mar 2005, Ian Stringer; ND, 1, ♀, Pouto Peninsula, Oneroa Rd, nr Te Ta Point, NZAC03006388, 29 Jan 2010, P De
Lange; ND, 1, ♀, Pouto Peninsula, Oneroa Rd, nr Te Ta Point, NZAC03006410, 29 Jan 2010, P De Lange; ND, 1, ♀, Pouto
Peninsula, Oneroa Rd, nr Te Ta Point, NZAC03006401, 29 Jan 2010, P De Lange; ND, 1, ♀, Pouto Peninsula, Oneroa Rd, nr Te
Ta Point, NZAC03006375, 29 Jan 2010, P De Lange; ND, 1, ♂, Marotiri Isd, Wh. (Marotere), NZAC04030798, 15 Mar 1931,
E S Gourlay; AK, 1, ♂, NW of Mangawhai, NZAC03004991, 36.11354, 174.55425, 5 Jan 2008, R Leschen; AK, 1, Mt
Tamahunga, NZAC03005315, 5 Jan 1982, J C Watt; AK, 1, ♂, Home Bay, Motuora I, NZAC03000358, 36 30.304, 174
47.6114, 14 Mar 2004, R Gardner-Gee; AK, 1, ♀, Still Bay, Motuora I, NZAC03000331, 36 30.283, 174 47.396, 16 Mar 2005,
R Gardner-Gee; AK, 1, ♂, Still Bay, Motuora I, NZAC03000380, 36 30.283, 174 47.396, 18 Mar 2005, R Gardner-Gee; AK, 1,
♀, Tapora, West of Wellsford, NZAC03000406, 36 21.100, 174 18.120, 18 Dec 2005, K Hill D Marshall; AK, 1, ♀, Tapora,
West of Wellsford, NZAC03000350, 36 21.100, 174 18.120, 18 Dec 2005, K Hill D Marshall; AK, 1, ♀, Mount Auckland,
NZAC03006385, 36 27.505, 174 28.022, 8 Jan 2010, TR Buckley R Leschen D Seldon; AK, 1, ♀, Wenderholm Regional Park,
Waiwera, NZAC03000408, 36 32.251, 174 42.317, 18 Dec 2005, D F Ward; AK, 1, ♂, Puhoi Motorway extension, Puhoi,
NZAC03000353, 36 30.708, 174 39.746, 27 Jun 1905, R Symcock; AK, 1, ♀, Puhoi Motorway extension, Puhoi,
NZAC03000487, 36 30.7, 174 39.8, May 2006, R Symcock; AK, 1, Mt Auckland, NZAC03005318, 17 Nov 1973, J S
Dugdale; AK, 1, ♂, Motuora I., NZAC04028301, 36 30, 174 47, Feb 2007, R Gardner-Gee; AK, 1, ♂, Motuora I.,
NZAC04028201, 36 30, 174 47, Feb 2007, R Gardner-Gee; AK, 1, ♂, Motuora I., NZAC04028290, 36 30, 174 47, Feb 2007, R
Gardner-Gee; AK, 1, ♀, Motuora I., NZAC04028208, 36 30, 174 47, Feb 2007, R Gardner-Gee; AK, 1, ♀, Motuora I.,
NZAC04028256, 36 30, 174 47, Feb 2007, R Gardner-Gee; AK, 1, ♀, Motuora I., NZAC04028261, 36 30, 174 47, Feb 2007, R
BUCKLEY ET AL.
476
·
Zootaxa 3900 (4) © 2014 Magnolia Press
Gardner-Gee; AK, 1, ♀, eggs, Wright Rd, Redvale, Albany, NZAC03005529, 36 41.7, 174 41.8, 29 Feb 2008, R Hoare; AK, 1,
♂, North shore, NZAC03000354, 27 Jun 1905, S Hibberd; AK, 1, Bethells, Matuku Res, NZAC03005352, Nov 1983, G Hall;
AK, 1, Bethells, Matuku Res, NZAC03005292, 15 Oct 1983, J C Watt; AK, 1, ♀, Oratia, Auckland, NZAC04030706, 13 Apr
1959, A M Richards; AK, 1, ♀, Oratia, Auckland, NZAC03005464, 36 54, 174 36, Nov 2004, J Drummond M Stanley; AK, 1,
Oratia, Auckland, NZAC03005476, 36 54, 174 36, Nov 2004, J Drummond M Stanley; AK, 1, Auckland, NZAC03005354,
Feb 1956, A Richards; AK, 1, ♂, Auckland, NZAC04030586, 18 Feb 1942, C G Taylor; AK, 1, Massey, Auckland, off Don
Bucks Rd, NZAC03005361, 22 Mar 1980; AK, 1, ♀, Titirangi, Auckland, NZAC04030744, 5 Oct 1927, E S Gourlay; AK, 2,
Titirangi, NZAC03005365, 6 Feb 1977, G W Ramsay; AK, 1, Titirangi, NZAC03005357, 18 Jan 1990, G W Ramsay; AK, 1,
Titirangi, NZAC03005383, Feb 1993, G W Ramsay; AK, 1, Titirangi, NZAC03005351, 4 Apr 1994, G W Ramsay; AK, 1,
Titirangi, NZAC03005377, Feb 1999, G W Ramsay; AK, 1, Titirangi, NZAC03005376, Apr 1992, G W Ramsay; AK, 4,
Titirangi, NZAC03005295, 18 Feb 1982, B G Bennett; AK, 1, ♀, Titirangi, NZAC04030532, Mar 1942, M W Carter; AK, 1,
♀, Titirangi, NZAC04030671, Mar 1942, M W Carter; AK, 1, Exhibition Drive, Titirangi, NZAC03005345, 26 Mar 1993, G W
Ramsay; AK, 1, Exhibition Drive, Titirangi, NZAC03005328, Nov 1991, G W Ramsay; AK, 1, ♂, Waiheke Is,
NZAC04030663, 4 Jan 1942, C Chamberlain; AK, 1, ♂, Waiheke Is, NZAC04030563, 4 Jan 1942, C Chamberlain; AK, 1, ♂,
Noises Is, Otata I, NZAC03000119, 12 Dec 1977, J C Watt; AK, 1, Noises Is, Otata I, Audreys Track, NZAC03005368, 25 Oct
1979, M F Tocker B A Bradshaw; AK, 1, Noises Is, Motuhoropapa I, Snail Flat nr pit trap 8, NZAC03005371, 24 Apr 1979, I
A E Atkinson; AK, 1, Noises Is, Motuhoropapa I, NZAC03005291, 22 Aug 1978, G W Ramsay; AK, 1, ♂, Shona restoration
plot, Border Rd, Opanuku stream, Henderson, NZAC03000335, 36 53.294, 174 37.014, 13 Dec 2005, D F Ward; AK, 1, ♀,
Shona restoration plot, Border Rd, Opanuku stream, Henderson, NZAC03000375, 36 53.294, 174 37.014, 13 Dec 2005, D F
Ward; AK, 1, ♀, Gills Reserve, Albany, North Shore, NZAC03000332, 36 43.596, 174 41.502, 3 Apr 2004, K Green; AK, 1, ♀,
Gills Reserve, Albany, North Shore, NZAC03000403, 36 43.596, 174 41.502, 3 Apr 2004, K Green; AK, 1, Lucas Creek,
Albany, NZAC03000479, 36 43.407, 174 41.518, 7 Mar 2004, TR Buckley; AK, 1, ♀, Muriwai Beach, Maori Bay Trk,
NZAC03000436, 36 49.909, 174 25.579, 17 Nov 2003, P Langhoff; AK, 1, Hillsborough, Auckland, NZAC03005362, 6 Nov
1975, W Wouts; AK, 1, Auckland, Mt Albert Res Centre, NZAC03005366, Mar 1992; AK, 1, Mt Albert, MARC car park,
NZAC03005303, Jun 1986, P A Maddison; AK, 1, ♂, Owairaka, Auckland, NZAC04030546, Jul 1940, D Spiller; AK, 1, ♂,
Remuera, NZAC04030691, 12 Apr 1939, E Carr Smith; AK, 1, Auckland, Meadowbank, NZAC03005384, 7 Apr 1991, J
Broadbent; AK, 1, ♀, eggs, Maraetai, Omana Reg.Pk., NZAC03005573, 35 52.793, 174 2.526, 21 Feb 2008, N A Martin; AK,
1, ♂, Maraetai, Omana Reg.Pk., NZAC03006402, 36 52.793, 175 2.526, 22 Mar 2009, N A Martin; AK, 1, ♂, Anawhata,
Auckland, NZAC03005466, 36 55, 174 27, Dec 2004, J Drummond M Stanley; AK, 1, ♂, Anawhata, Auckland,
NZAC03005494, 36 55, 174 27, Dec 2004, J Drummond M Stanley; AK, 1, ♂, Parau Trk, Lower Nihoputu Reservoir,
Auckland, NZAC03000376, 36 57.966, 174 36.703, 20 Feb 2005, TR Buckley; AK, 1, ♀, Parau Trk, Lower Nihoputu
Reservoir, Auckland, NZAC03000398, 36 57.966, 174 36.703, 20 Feb 2005, TR Buckley; AK, 1, ♀, Tapapakanga Reg.Park,
Orere, NZAC03004877, 36 58.602, 175 15.379, 31 Mar 2008, TR Buckley D Seldon S Forgie; AK, 1, ♀, Tapapakanga
Reg.Park, Orere, NZAC03004913, 36 58.602, 175 15.379, 31 Mar 2008, TR Buckley D Seldon S Forgie; AK, 1, ♂,
Tapapakanga Reg.Park, Orere, NZAC03004923, 36 58.602, 175 15.379, 31 Mar 2008, TR Buckley D Seldon S Forgie; AK, 1,
♂, Tapapakanga Reg.Park, Orere, NZAC03004809, 36 58.602, 175 15.379, 31 Mar 2008, TR Buckley D Seldon S Forgie; AK,
1, ♀, Tapapakanga Reg.Park, Orere, NZAC03004973, 36 58.602, 175 15.379, 31 Mar 2008, TR Buckley D Seldon S Forgie;
AK, 1, ♀, eggs, Kakamatua Inlet, Waitakere Ranges, NZAC03005562, 36 59.98, 174 35.54, 6 Jan 2009, R Hoare; AK, 1, ♀,
Waharau Reg.Park, Hunua Ranges, NZAC03004889, 37 2.493, 175 17.422, 31 Mar 2008, TR Buckley S Forgie D Seldon R
Hoare; AK, 1, ♀, Waharau Reg.Park, Hunua Ranges, NZAC03004796, 37 2.493, 175 17.422, 31 Mar 2008, TR Buckley S
Forgie D Seldon R Hoare; AK, 1, ♀, Waharau Reg.Park, Hunua Ranges, NZAC03004800, 37 2.493, 175 17.422, 31 Mar 2008,
TR Buckley S Forgie D Seldon R Hoare; AK, 1, ♂, Waharau Reg.Park, Hunua Ranges, NZAC03004978, 37 2.493, 175 17.422,
31 Mar 2008, TR Buckley S Forgie D Seldon R Hoare; CL, 1, ♂, Whangapoua, 4.6 km N of SH25 turnoff and Te Rerenga,
Coromandel, NZAC03000089, 36 43.597, 175 36.875, 12 Mar 2005, TR Buckley; CL, 1, ♂, Whangapoua, 4.6 km N of SH25
turnoff and Te Rerenga, Coromandel, NZAC03000028, 36 43.597, 175 36.875, 12 Mar 2005, TR Buckley; CL, 1, ♂,
Whangapoua, 4.6 km N of SH25 turnoff and Te Rerenga, Coromandel, NZAC03000071, 36 43.597, 175 36.875, 12 Mar 2005,
TR Buckley; CL, 1, ♂, Whangapoua, 4.6 km N of SH25 turnoff and Te Rerenga, Coromandel, NZAC03000013, 36 43.597, 175
36.875, 12 Mar 2005, TR Buckley; CL, 1, ♂, Whangapoua, 4.8 km N of SH25 turnoff and Te Rerenga, Coromandel,
NZAC03000049, 36 43.597, 175 36.875, 12 Mar 2005, TR Buckley; CL, 1, ♀, Whangapoua, 4.6 km N of SH25 turnoff and Te
Rerenga, Coromandel, NZAC03000092, 36 43.597, 175 36.875, 12 Mar 2005, TR Buckley; CL, 1, ♀, Little Windy Hill, Great
Barrier Is, NZAC03000409, 18 Dec 2005, J Early; CL, 1, ♂, Little Windy Hill, Great Barrier I, NZAC03000095, 18 Dec 2005,
J Early; CL, 1, ♂, Little Barrier Island, NZAC04028349, 36 12, 175 5, 29 Nov 2006, TR Buckley, K Marske, D Seldon, R A B
Leschen; CL, 2, Little Barrier Island, NZAC03005346, Jan 1956, G Ramsay; CL, 2, Little Barrier Island, NZAC03005323, Jan
1956, G Ramsay; CL, 1, ♂, Little Barrier Island, NZAC04030666, Mar 1922, Milligan; CL, 1, ♂, Little Barrier Island,
NZAC04030627, Mar 1922, Milligan; CL, 1, ♀, Little Barrier , NZAC04030579, Mar 1922, Milligan; CL, 1, ♀, Little Barrier,
NZAC04030625, Mar 1922, Milligan; CL, 1, ♂, Little Barrier , NZAC04030651, Mar 1922, Milligan; CL, 1, ♀, Little Barrier
, NZAC04030751, Mar 1922, Milligan; CL, 1, ♀, Little Barrier , NZAC04030745, Mar 1922, Milligan; CL, 1, ♂, Little Barrier
, NZAC04030784, Mar 1922, Milligan; CL, 1, ♂, Little Barrier , NZAC04030862, Mar 1922, Milligan; CL, 1, ♂, Little Barrier
, NZAC04030828, Mar 1922, Milligan; CL, 1, ♂, Little Barrier , NZAC04030766, Mar 1922, Milligan; CL, 1, Little Barrier
Island, NZAC03005322, , CR?; CL, 1, ♀, Little Barrier Island, NZAC04030657, Jan 1932, A E Brookes; CL, 1, ♂, Little
Barrier Island, NZAC04030672, 26 Nov 1954, G W Ramsay; CL, 9, Eggs, Little Barrier Island, NZAC04030634, 26 Nov
Zootaxa 3900 (4) © 2014 Magnolia Press
·
477
REVISION OF CLITARCHUS
1954, G Ramsay; CL, 1, ♀, Little Barrier Island, NZAC04030637, Feb 1952, J T Salmon; CL, 1, ♀, Little Barrier Island,
NZAC04030613, Feb 1953, J T Salmon; CL, 1, ♂, Little Barrier Island, NZAC04030834, Feb 1953, J T Salmon; CL, 1, ♂,
Little Barrier Island, NZAC04030769, Feb 1953, J T Salmon; CL, 1, ♂, Little Barrier Island, NZAC04030697, Feb 1953, J T
Salmon; CL, 1, ♂, Little Barrier Island, NZAC04030700, 20 Feb 1953, J T Salmon; CL, 1, ♀, Little Barrier Island,
NZAC04030648, 22 Feb 1953, J T Salmon; CL, 5, Eggs, nymphs, Little Barrier Island, NZAC04030559, 22 Feb 1953, J T
Salmon; CL, 1, ♂, Little Barrier Island, NZAC04030763, 20 Feb 1953, J T Salmon; CL, 1, ♀, Little Barrier I, Awaroa Stream,
NZAC04028346, 36 13, 175 5, 29 Nov 2006, TR Buckley, K Marske, D Seldon; CL, 1, ♂, Little Barrier I, Awaroa Stream,
NZAC04028278, 36 13, 175 5, 29 Nov 2006, TR Buckley, K Marske, D Seldon; CL, 1, ♀, Little Barrier I, Awaroa Stream,
NZAC04028356, 36 13, 175 5, 29 Nov 2006, TR Buckley, K Marske, D Seldon; CL, 1, ♂, Little Barrier I, DoC Rangers
graden, NZAC04028326, 36 13.176, 175 3.409, 27 Nov 2006, TR Buckley, D Seldon; CL, 1, ♀, Bunk house, Little Barrier I,
NZAC03000384, 36 13.192, 175 3.436, 20 Jun 2005, D Seldon; CL, 1, ♂, Bunk house, Little Barrier I, NZAC03000404, 36
13.192, 175 3.436, 20 Jun 2005, D Seldon; CL, 1, ♀, Valley Track, Little Barrier Island, NZAC03000333, 36 12.940, 175
3.476, 7 Dec 2005, D Seldon; CL, 1, Valley Track, Little Barrier Island, NZAC03000452, 36 13, 175 3, 30 Nov 2006, TR
Buckley K Marske R Leschen D Seldon; CL, 1, ♂, Little Barrier I, Valley Track, NZAC04028277, 36 12, 175 3, 28 Nov 2006,
TR Buckley, D Seldon, K Marske, R A B Leschen; CL, 1, ♂, Flats, Little Barrier Island, NZAC03000360, 36 12.940, 175
3.476, 9 Dec 2005, D Seldon; CL, 1, ♂, Flats, Little Barrier Island, NZAC03000038, 36 12.940, 175 3.476, 8 Dec 2005, D
Seldon; CL, 1, ♂, Flats, Little Barrier Island, NZAC03000026, 36 12.940, 175 3.476, 8 Dec 2005, D Seldon; CL, 1, ♀, Great
Barrier I, between Gooseberry Flat & Mulberry Grove, NZAC04030643, 29 Dec 1979, G Lowe; CL, 1, ♂, Great Barrier I, road
between Gooseberry Flat & Mulberry Grove, NZAC04030810, 29 Dec 1979, G Lowe; CL, 1, ♂, Great Barrier I, road between
Gooseberry Flat & Mulberry Grove, NZAC04030724, 29 Dec 1979, G Lowe; CL, 1, ♀, Great Barrier I, road between
Gooseberry Flat & Mulberry Grove, NZAC04030694, 29 Dec 1979, G Lowe; CL, 2, Great Barrier I, Whangaparapara, Witheys
Track, NZAC03005374, 28 Mar 1978, R Archibald J C Watt; CL, 1, Great Barrier I, Kaiarara Valley, NZAC03005372, 25 Mar
1978, J C Watt; CL, 1, Great Barrier I, Kaiarara Valley, NZAC03005359, 26 Mar 1978, J C Watt; CL, 1, ♂, Great Barrier I,
Kaiarara Str, NZAC04030762, 23 Mar 1978, ESNZ Field trip; CL, 1, Great Barrier I, Mt Hobson, NZAC03005349, 26 Mar
1978, J C Watt; CL, 1, ♂, Wilson Bay, SH25 roadside, 2.5 km N of Tapu, Coromandel, NZAC03000412, 36 52.824, 175
25.687, 12 Feb 2005, TR Buckley; CL, 1, ♀, Wilson Bay, SH25 roadside, 2.5 km N of Tapu, Coromandel, NZAC03000399, 36
52.824, 175 25.687, 12 Feb 2005, TR Buckley; CL, 1, ♂, Wilson Bay, SH25 roadside, 2.5 km N of Tapu, Coromandel,
NZAC03000073, 36 52.824, 175 25.687, 12 Feb 2005, TR Buckley; CL, 1, Ruamahuaiti Is, Alderman Is, NZAC03005363, 12
Nov 1972, G W Ramsay; CL, 1, Ruamahuaiti Is, Alderman Is, NZAC03005360, 8 Nov 1972, G W Ramsay; CL, 1, ♀,
Ruamahuaiti Is, Alderman Is, NZAC03000096, 36 58.486, 176 4.877, 8 Nov 1972, G W Ramsay; CL, 3, Ruamahuanui I,
Alderman Is, NZAC03005337, 13 Nov 1972, G W Ramsay; CL, 1, ♂, Mercury Is, Korapuki, NZAC03000411, 36 39.503, 175
50.925, 19 Nov 2005, C J Green; CL, 1, ♂, Mercury Is, Korapuki, NZAC03000111, 36 39.503, 175 50.925, 19 Nov 2005, C J
Green; CL, 1, ♀, Mercury Is, Korapuki, NZAC03000131, 36 39.503, 175 50.925, 19 Nov 2005, C J Green; CL, 1, ♀, Mercury
Is, Korapuki, NZAC03000112, 36 39.503, 175 50.925, 19 Nov 2005, C J Green; CL, 1, ♀, Mercury Is, Korapuki,
NZAC03000124, 36 39.503, 175 50.925, 19 Nov 2005, C J Green; CL, 1, ♀, Mercury Is, Korapuki, NZAC03000128, 36
39.503, 175 50.925, 19 Nov 2005, C J Green; CL, 1, ♂, Mercury Is, Korapuki, NZAC03000121, 36 39.503, 175 50.925, 19
Nov 2005, C J Green; CL, 1, ♀, Mercury Is, Korapuki, NZAC03000361, 36 39.503, 175 50.925, 19 Nov 2005, C J Green; CL,
2, Mercury Is, Korapuki, NZAC03005334, 18 Nov 1972, G W Ramsay; CL, 1, above Lunch Bay, Red Mercury Is,
NZAC03005285, Aug 1971, J Johannesson; CL, 3, Ruamahuaiti I, Alderman Is, NZAC03005312, 12 Nov 1972, G W Ramsay;
CL, 1, ♀, Waiomu, NZAC03000485, 37 01.6, 175 31.5, 18 Dec 2006, R Hoare; CL, 1, ♀, 3.7 km N of Stony Bay, on road to
Stony Bay, Port Charles area, NZAC03000338, 36 30.717, 175 25.151, 20 Mar 2005, K Hill D Marshall; CL, 1, ♀, 3.7 km N of
Stony Bay, on road to Stony Bay, Port Charles area, NZAC03000118, 36 30.717, 175 25.151, 20 Mar 2005, K Hill D Marshall;
CL, 1, ♂, 3.7 km N of Stony Bay, on road to Stony Bay, Port Charles area, NZAC03000364, 36 30.717, 175 25.151, 20 Mar
2005, K Hill D Marshall; CL, 1, ♂, 3.7 km N of Stony Bay, on road to Stony Bay, Port Charles area, NZAC03000339, 36
30.717, 175 25.151, 20 Mar 2005, K Hill D Marshall; CL, 1, ♂, 3.7 km N of Stony Bay, on Rd to Stony Bay, Port Charles area,
NZAC03000058, 36 30.717, 175 25.151, 20 Mar 2005, K Hill D Marshall; CL, 1, ♀, 3.7 km N of Stony Bay, on Rd to Stony
Bay, Port Charles area, NZAC03000020, 36 30.717, 175 25.151, 20 Mar 2005, K Hill D Marshall; CL, 1, ♀, 3.7 km N of Stony
Bay, on Rd to Stony Bay, Port Charles area, NZAC03000101, 36 30.717N, 175 25.151E, 20 Mar 2005, K Hill D Marshall; CL,
1, ♂, Start of Piranui trk, Kopu-Hikuai Rd, NZAC03000365, Jan 2006, C Winks; CL, 1, ♀, Start of Piranui trk, Kopu-Hikuai
Rd, NZAC03000377, Jan 2006, C Winks; CL, 1, ♀, Start of Piranui trk, Kopu-Hikuai Rd, NZAC03000390, Jan 2006, C Winks;
CL, 1, Coromandel summit, Kopu-Hikuai road, NZAC03005286, Mar 1993, G W Ramsay; CL, 2, ♀, ♂, Kuaotona (Kuaotunu),
Wharekano Rd, rest area, near Whitianga, NZAC03000428, 36 45.322, 175 43.604, 20 Feb 2002, TR Buckley; CL, 4,
Kuaotona (Kuaotunu), Wharekano Rd, rest area, near Whitianga, NZAC03000488, 36 45.322, 175 43.604, 20 Feb 2002, TR
Buckley; CL, 1, ♂, Alderman Is, Ruamahuaiti I, NZAC03000374, 36 58.486, 176 4.877, 8 Nov 1972, G W Ramsay; CL, 1, ♂,
19 km N of Coromandel town, S of Colville, NZAC03000397, 20 Mar 2005, K Hill D Marshall; CL, 1, ♀, 19 km N of
Coromandel town, S of Colville, NZAC03000055, 20 Mar 2005, K Hill D Marshall; CL, 1, 19 km N of Coromandel town, S of
Colville, NZAC03000078, 20 Mar 2005, K Hill D Marshall; CL, 1, ♀, 19 km N of Coromandel town, S of Colville,
NZAC03000093, 20 Mar 2005, K Hill D Marshall; CL, 1, ♂, Kauaeranga Valley, Thames, NZAC04030675, 18 Jan 1960, J I
Townsend R Zondag; CL, 1, ♀, Kauaeranga River, Thames, NZAC04030780, 29 Nov 1953, J S Armstrong; CL, 1, ♀, Waihi,
NZAC03000385, 14 Jan 2005, C Winks; CL, 1, ♀, Waihi, NZAC03000418, 14 Jan 2005, C Winks; CL, 1, ♂, Mayor Island,
NZAC04030583, 28 Nov 1948, S A Rumsey; CL, 1, ♂, Mayor Island, NZAC04030538, 28 Nov 1948, S A Rumsey; CL, 1, ♀,
BUCKLEY ET AL.
478
·
Zootaxa 3900 (4) © 2014 Magnolia Press
Mayor Island, NZAC04030537, 28 Nov 1948, S A Rumsey; CL, 1, ♀, Mayor Island, NZAC04030552, 28 Nov 1948, S A
Rumsey; CL, 1, ♀, Mayor Island, NZAC04030622, 28 Nov 1948, S A Rumsey; CL, 1, ♂, Mayor Island, NZAC04030768, 28
Nov 1948, S A Rumsey; CL, 1, ♀, Mayor Island, NZAC04030685, 28 Nov 1948, S A Rumsey; CL, 1, Mayor Island, Panui
pen. On track to Western Bay, NZAC03009372, 30 Jan 2012, O R & C J Green; WO, 1, ♂, Waitakaruru Reserve, roadside,
NZAC03000122, 37 16.766, 175 19.442, 3 Mar 2005, TR Buckley D Seldon; WO, 1, ♀, Waitakaruru Reserve, roadside,
NZAC03000379, 37 16.766, 175 19.442, 3 Mar 2005, TR Buckley D Seldon; WO, 1, ♀, Waitakaruru Reserve, roadside,
NZAC03000329, 37 16.766, 175 19.442, 3 Mar 2005, TR Buckley D Seldon; WO, 1, ♀, Waitakaruru Reserve, roadside,
NZAC03000417, 37 16.766, 175 19.442, 3 Mar 2005, TR Buckley D Seldon; WO, 1, ♂, Mangakora Nature Walk, Mt Pirongia,
NZAC03000035, 37 58.052, 175 9.257, 28 Feb 2005, TR Buckley; WO, 1, ♂, Mangakora Nature Walk, Mt Pirongia,
NZAC03000065, 37 58.052, 175 9.257, 28 Feb 2005, TR Buckley; WO, 1, ♀, Mangakora Nature Walk, Mt Pirongia,
NZAC03000029, 37 58.052, 175 9.257E, 28 Feb 2005, TR Buckley; WO, 1, ♀, Mangakora Nature Walk, Mt Pirongia,
NZAC03000081, 37 58.052, 175 9.257, 28 Feb 2005, TR Buckley; WO, 1, ♂, Mangakora Nature Walk, Mt Pirongia,
NZAC03000087, 37 58.052, 175 9.257, 28 Feb 2005, TR Buckley ; WO, 1, ♂, Mt Karioi, Pirongia State Forest Pk, Start of
Waireke Track, NZAC04028310, 37 52.80, 174 47.54, 10 Dec 2006, TR Buckley, R Hoare, G Hall, D Seldon; WO, 1, ♀, Mt
Karioi, Pirongia State Forest Pk, Start of Waireke Track, NZAC04028250, 37 52.80, 174 47.54, 10 Dec 2006, TR Buckley, R
Hoare, G Hall, D Seldon; WO, 1, ♀, Mt Karioi, Pirongia State Forest Pk, Start of Waireke Track, NZAC04028219, 37 52.80,
174 47.54, 10 Dec 2006, TR Buckley, R Hoare, G Hall, D Seldon; WO, 1, ♀, Bridal Veil Falls, NZAC04028351, 37 54.44, 174
53.92, 10 Dec 2006, TR Buckley, R Hoare, G Hall, D Seldon; WO, 3, ♂, Cambridge, Maungakawa Scenic Reserve,
NZAC04028332, 37 51.592, 175 31.845, 25 Feb 2007, TR Buckley, R Hoare, G Hall; WO, 1, Cambridge, Maungakawa Scenic
Reserve, NZAC03005287, 37 51.592, 175 31.845, 25 Feb 2007, TR Buckley, R Hoare, G Hall; WO, 1, Cambridge,
Maungakawa Scenic Reserve, NZAC03005344, 37 51.592, 175 31.845, 25 Feb 2007, TR Buckley, R Hoare, G Hall; WO, 1, ♀,
Mt Te Aroha, bottom of Summit Rd, NZAC03000123, 37 31.216, 175 43.639, 3 Mar 2005, TR Buckley D Seldon ; WO, 1, ♀,
Mt Te Aroha, bottom of Summit Rd, NZAC03000400, 37 31.216, 175 43.639, 3 Mar 2005, TR Buckley D Seldon ; WO, 1, ♀,
Mt Te Aroha, bottom of Summit Rd, NZAC03000373, 37 31.216, 175 43.639, 3 Mar 2005, TR Buckley D Seldon ; WO, 1, ♂,
Mt Te Aroha, bottom of summit road, NZAC03000022, 37 31.216, 175 43.639, 3 Mar 2005, TR Buckley D Seldon; WO, 1, ♀,
Mt Te Aroha, bottom of summit road, NZAC03000036, 37 31.216, 175 43.639, 3 Mar 2005, TR Buckley D Seldon; WO, 1, ♂,
Mt Te Aroha, bottom of summit Rd, NZAC03000063, 37 31.216, 175 43.639, 3 Mar 2005, TR Buckley D Seldon; WO, 1, ♀,
Mt Te Aroha, bottom of summit Rd, NZAC03000085, 37 31.216, 175 43.639, 3 Mar 2005, TR Buckley D Seldon; WO, 1, ♂,
Mt Te Aroha, bottom of summit Rd, NZAC03000069, 37 31.216, 175 43.639, 3 Mar 2005, TR Buckley D Seldon; WO, 1, ♂,
Hakarimata Ra, Kauri Loop Trk, NZAC03005181, 37 36.705, 175 9.325, 22 Jan 2009, TR Buckley S Forgie G Hall R Hoare;
WO, 1, ♂, Hakarimata Ra, Kauri Loop Trk, NZAC03005395, 37 36.705, 175 9.325, 22 Jan 2009, TR Buckley S Forgie G Hall
R Hoare; WO, 1, ♂, Hakarimata Ra, Kauri Loop Trk, NZAC03005239, 37 36.705, 175 9.325, 22 Jan 2009, TR Buckley S
Forgie G Hall R Hoare; WO, 1, ♀, Hakarimata Ra, Kauri Loop Trk, NZAC03005417, 37 36.705, 175 9.325, 22 Jan 2009, TR
Buckley S Forgie G Hall R Hoare; WO, 1, ♀, Hakarimata Ra, Kauri Loop Trk, NZAC03005229, 37 36.705, 175 9.325, 22 Jan
2009, TR Buckley S Forgie G Hall R Hoare; WO, 1, ♀, Waipapa Scenic Reserve, Lake Waipapa, NZAC03000591, 38 17, 175
40, 23 Feb 2007, TR Buckley R Hoare G Hall; WO, 1, ♀, Waipapa Scenic Reserve, Lake Waipapa, NZAC04028274, 38 17,
175 40, 23 Feb 2007, TR Buckley, R Hoare, G Hall; WO, 1, ♂, Waipapa Scenic Reserve, Lake Waipapa, NZAC04028315, 38
17, 175 40, 23 Feb 2007, TR Buckley, R Hoare, G Hall; WO, 1, ♀, Waipapa Scenic Reserve, Lake Waipapa, NZAC04028304,
38 17, 175 40, 23 Feb 2007, TR Buckley, R Hoare, G Hall; WO, 1, ♀, Waipapa Scenic Reserve, Lake Waipapa,
NZAC04028352, 38 17, 175 40, 23 Feb 2007, TR Buckley, R Hoare, G Hall; WO, 1, Hillcrest, Hamilton, NZAC03005348, 21
Feb 1995, J Leathwick; WO, 1, ♂, Glen Massey, Ngaruawahia, NZAC03006395, 7 Dec 2009, C Winks; WO, 1, ♂, SH31, 3 km
E of Kawhia, NZAC03000059, 38 3.152, 174 51.836, 18 Mar 2005, K Hill D Marshall; BP, 1, ♂, Karangahake Gorge,
Waitawheta River, NZAC03005413, 37 25.721, 175 43.722, 9 Jan 2009, N A Martin; BP, 1, ♀, Karangahake Gorge,
Waitawheta River, NZAC03005267, 37 25.721, 175 43.722, 9 Jan 2009, N A Martin; BP, 1, ♀, Karangahake Gorge,
Waitawheta River, NZAC03005154, 37 25.721, 175 43.722, 9 Jan 2009, N A Martin; BP, 1, ♂, Karangahake Gorge,
Waitawheta River, NZAC03005451, 37 25.721, 175 43.722, 9 Jan 2009, N A Martin; BP, 1, ♀, Karangahake Gorge,
Waitawheta River, NZAC03005408, 37 25.721, 175 43.722, 9 Jan 2009, N A Martin; BP, 1, ♀, Karangahake Gorge,
Waitawheta River, NZAC03005488, 37 25.721, 175 43.722, 9 Jan 2009, N A Martin; BP, 1, ♀, Karangahake Gorge,
Waitawheta River, NZAC03005392, 37 25.721, 175 43.722, 9 Jan 2009, N A Martin; BP, 1, Eggs, Karangahake Gorge,
Waitawheta River, NZAC03005396, 37 25.721, 175 43.722, 9 Jan 2009, N A Martin; BP, 1, Mt Te Aroha, at gate,
NZAC03005347, 37 32, 175 44, 5 Nov 2005, R Leschen S Nomura; BP, 2, Mt Te Aroha, M.Hansen's residence,
NZAC03005343, 27 Feb 1992, R Henderson M Belsten J Dugdale; BP, 3, Mt Te Aroha, M.Hansen's residence,
NZAC03005329, R Henderson M Belsten J Dugdale; BP, 1, ♀, Katikati, Lindemann Track, Lindemann Road,
NZAC04028322, 37 32, 175 52, 8 Nov 2005, R Leschen; BP, 1, ♀, Katikati, Track at top of Lindeman Rd, NZAC03000542, 37
32.662, 175 52.397, 19 May 2006, B Chudleigh; BP, 1, ♀, Waitengaue Stm, Kaimai Ra, NZAC03000378, 37 30.288, 175
52.172, 21 Mar 2004, K Green; BP, 1, ♀, Waenga Bush, Lottin Pt Rd, NZAC03006394, 37 33.771, 178 9.135, 3 Dec 2009, TR
Buckley D Seldon R Hoare G Hall; BP, 1, ♀, Hicks Bay Motel, beach walk, NZAC03006386, 37 35.780, 178 18.257, 2 Dec
2009, TR Buckley D Seldon R Hoare G Hall; BP, 1, ♀, Te Koau, Hicks Bay, NZAC03006377, 37 35.996, 178 17.973, 30 Nov
2009, TR Buckley D Seldon R Hoare G Hall; BP, 1, ♀, Te Koau, Hicks Bay, NZAC03006392, 37 35.996, 178 17.973, 30 Nov
2009, TR Buckley D Seldon R Hoare G Hall; GB, 1, Te Koau, Hicks Bay, NZAC03005588, 37 35.996, 178 17.973, 4 Dec
2009, TR Buckley D Seldon R Hoare G Hall; GB, 1, Eggs, Te Koau, Hicks Bay, NZAC03005588, 37 35.996, 178 17.973, 4
Zootaxa 3900 (4) © 2014 Magnolia Press
·
479
REVISION OF CLITARCHUS
Dec 2009, TR Buckley D Seldon R Hoare G Hall; BP, 1, Lk Okataina, S end of Lk Okataina Rd, near lodge, NZAC03000114,
38 6.002, 176 25.756, 27 Mar 2006, TR Buckley D Seldon R Hoare; BP, 1, ♂, Lk Okataina, S end of Lk Okataina Rd, near
lodge, NZAC03000126, 38 6.002, 176 25.756, 27 Mar 2006, TR Buckley D Seldon R Hoare; BP, 1, ♂, Lk Okataina, S end of
Lk Okataina Rd, near lodge, NZAC03000395, 38 6.002, 176 25.756, 27 Mar 2006, TR Buckley D Seldon R Hoare; BP, 1, ♂,
Lk.Okataina, S end of Lk Okataina Rd, near lodge, NZAC03000465, 38 6.002, 176 25.756, 27 Mar 2006, TR Buckley D
Seldon R Hoare; BP, 1, ♂, Lk.Okataina, S end of Lk Okataina Rd, near lodge, NZAC03000475, 38 6.002, 176 25.756, 27 Mar
2006, TR Buckley D Seldon R Hoare; BP, 1, Te Pu, Mamaku, NZAC03005364, 14 Feb 1959; BP, 1, ♀, Moutohora Island,
NZAC03000328, 37 51.316, 176 58.447, Mar 2006; BP, 1, ♀, Moutohora Island, NZAC03000407, 37 51.316, 176 58.447, Mar
2006; BP, 1, ♀, Moutohora Island, NZAC03000349, 37 51.316, 176 58.447, Mar 2006; BP, 1, ♀, Moutohora Island,
NZAC03000025, 37 51.316, 176 58.447, Mar 2006; BP, 1, ♀, Moutohora Island, NZAC03000039, 37 51.316, 176 58.447, Mar
2006; BP, 1, Omaruparoa, NZAC03005308, 14 Apr 1992, J S Dugdale; BP, 1, Rereauira, NZAC03005321, 26 Jan 1993, J S
Dugdale; BP, 1, ♀, Te Koau, NZAC04030767, 15 Mar 1993, J S Dugdale; BP, 1, ♀, Te Koau, Waihiere Falls, NZAC04030865,
15 Mar 1993, T K Crosby; BP, 1, ♀, Te Koau, Waihiere Falls, NZAC04030752, 15 Mar 1993, T K Crosby; BP, 1, ♀, Te Koau,
Waihiere Falls, NZAC04030736, 15 Mar 1993, T K Crosby; BP, 1, ♀, Te Koau, Waihiere Falls, NZAC04030750, 15 Mar 1993,
T K Crosby; BP, 1, ♀, Te Koau, Waihiere Falls, NZAC04030755, 15 Mar 1993, T K Crosby; BP, 1, ♀, Te Koau, Waihiere Falls,
NZAC04030686, 15 Mar 1993, T K Crosby; BP, 1, ♀, Te Koau, Waihiere Falls, NZAC04030835, 15 Mar 1993, T K Crosby;
BP, 1, ♀, Karakatuwhero V., NZAC04030797, 17 Mar 1993, J S Dugdale; BP, 1, Hicks Bay? East Cape, NZAC03005339, Oct
1993, G Hall;TO, 1, ♀, Taupo, NZAC04030759, Jan 1945, J S Armstrong; GB, 1, ♀, Maori Track, Awaawaroa Strm, Lake
Waikaremoana, Te Urewera Nat.Pak., NZAC03000392, 38 48.978, 177 8.142, 29 Mar 2006, TR Buckley D Seldon; GB, 1, ♀,
Te Araroa Campground, East Cape, NZAC03000334, 37 36.384, 178 18.836, 3 Jan 2006, TR Buckley; GB, 1, ♀, East Cape
Lighthouse Trk, Otiki, NZAC03000327, 37 41.388, 178 32.836, 3 Jan 2006, TR Buckley; GB, 1, ♀, East Cape Lighthouse Trk,
Otiki, NZAC03000413, 37 41.388, 178 32.836, 3 Jan 2006, TR Buckley; GB, 1, ♀, Lake Whakamarino, Lake Waikaremoana,
NZAC04028325, 38 48.48, 177 9.23, 23 Feb 2007, TR Buckley, G Hall; GB, 2, ♀, Lake Whakamarino, Lake Waikaremoana,
NZAC04028280, 38 48.48, 177 9.23, 23 Feb 2007, TR Buckley, G Hall;HB, 1, ♀, Mohaka River, SH2 rest area,
NZAC04028292, 39 4.457, 177 7.706, 21 Feb 2007, TR Buckley, R Hoare, G Hall;HB, 1, ♀, Mohaka River, SH2 rest area,
NZAC04028334, 39 4.457, 177 7.706, 21 Feb 2007, TR Buckley, R Hoare, G Hall;HB, 1, ♀, Mohaka River, SH2 rest area,
NZAC04028340, 39 4.457, 177 7.706, 21 Feb 2007, TR Buckley, R Hoare, G Hall;HB, 2, ♀, Mohaka River, SH2 rest area,
NZAC04028283, 39 4.457, 177 7.706, 21 Feb 2007, TR Buckley, R Hoare, G Hall;HB, 1, ♀, Mohaka River, SH2 rest area,
NZAC04028210, 39 4.457, 177 7.706, 21 Feb 2007, TR Buckley, R Hoare, G Hall;TK, 1, ♀, Summit of Mt Messenger, SH 3,
NZAC03000130, 38 53.754, 174 35.950, 28 Feb 2005, TR Buckley;TK, 1, ♀, Summit of Mt Messenger, SH 3,
NZAC03000405, 38 53.754, 174 35.950, 28 Feb 2005, TR Buckley;TK, 1, ♀, Summit of Mt Messenger, SH 3,
NZAC03000367, 38 53.754, 174 35.950, 28 Feb 2005, TR Buckley;TK, 1, ♀, Summit of Mt Messenger, SH3,
NZAC03000041, 38 53.754, 174 35.950, 28 Feb 2005, TR Buckley;TK, 1, Summit of Mt Messenger, SH3, NZAC03000030,
38 53.754, 174 35.950, 28 Feb 2005, TR Buckley;TK, 1, ♀, Summit of Mt Messenger, SH 3, NZAC03000097, 38 53.754, 174
35.950, 28 Feb 2005, TR Buckley;TK, 1, ♂, Whangamomona Saddle, Awahou Ridge Track, top of saddle, NZAC03004988, 39
09.311, 174 41.986, 17 Jan 2008, TR Buckley R Leschen K Marske; RI, 1, ♀, Gentle Annie Rd, West of Kuripapango,
NZAC04028353, 39 23.77, 176 18.86, 20 Feb 2007, TR Buckley, R Hoare, G Hall; RI, 1, ♀, Gentle Annie Rd, West of
Kuripapango, NZAC04028228, 39 23.77, 176 18.86, 20 Feb 2007, TR Buckley, R Hoare, G Hall; RI, 1, ♀, Gentle Annie Rd,
West of Kuripapango, NZAC04028247, 39 23.77, 176 18.86, 20 Feb 2007, TR Buckley, R Hoare, G Hall; RI, 1, ♀, Gentle
Annie Rd, West of Kuripapango, NZAC04028305, 39 23.77, 176 18.86, 20 Feb 2007, TR Buckley, R Hoare, G Hall; RI, 1, ♀,
Gentle Annie Rd, West of Kuripapango, NZAC04028355, 39 23.77, 176 18.86, 20 Feb 2007, TR Buckley, R Hoare, G Hall;
WI, 1, ♀, Aramoho, NZAC04030775, Apr 1915; WI, 1, ♂, Atene Skyline Tk, north entrance, NZAC03004919, 39 43.616,
175.08.318, 18 Jan 2008, TR Buckley R Leschen K Marske; WI, 1, ♂, Atene Skyline Tk, north entrance, NZAC03004928, 39
43.616, 175.08.318, 18 Jan 2008, TR Buckley R Leschen K Marske; WI, 1, ♀, Atene Skyline Tk, north entrance,
NZAC03004964, 39 43.616, 175.08.318, 18 Jan 2008, TR Buckley R Leschen K Marske; WI, 1, ♂, Atene Skyline Tk, north
entrance, NZAC03004959, 39 43.616, 175.08.318, 18 Jan 2008, TR Buckley R Leschen K Marske; WI, 1, ♂, 5.7 km N of
Upokongaro, NZAC03000048, 39 50.970, 175 7.397, Dec 2005, K Hill D Marshall; WI, 1, ♀, Kahikatea walkway, Pohangina
Valley, NZAC03004960, 40 04.968, 175 54.400, 19 Jan 2008, TR Buckley R Leschen K Marske; WI, 1, ♀, Kahikatea
walkway, Pohangina Valley, NZAC03004972, 40 04.968, 175 54.400, 19 Jan 2008, TR Buckley R Leschen K Marske; WI, 1,
♀, Pohangina , NZAC03004971, 40 09.014, 175 46.227, 18 Jan 2008, TR Buckley RAB Leschen K Marske; WA, 1, ♀, Te Uri/
Dannevirke, NZAC03000391, 40 13.461, 176 16.322, 21 Dec 2005, K Hill D Marshall; WA, 1, ♀, Te Uri/Dannevirke,
NZAC03000340, 40 13.461, 176 16.322, 21 Dec 2005, K Hill D Marshall; WA, 1, ♀, Te Uri/Dannevirke, NZAC03000355, 40
13.461, 176 16.322, 21 Dec 2005, K Hill D Marshall; WA, 1, ♀, Te Uri, Dannevirke, NZAC03000075, 40 13.461, 176 16.322,
21 Dec 2005, K Hill D Marshall; WA, 1, ♀, 4.7 km N of Waihi Valley Rd, on road from Waimiro to Waipatiki, Wairarapa,
NZAC03000076, 40 23.779, 176 14.258, 13 Mar 2005, K Hill D Marshall; WA, 1, ♀, 4.7 km N of Waihi Valley Rd, on road
from Waimiro to Waipatiki, Wairarapa, NZAC03000072, 40 23.779, 176 14.258, 13 Mar 2005, K Hill D Marshall; WA, 1, ♀,
Puketoi Ra, Pahiatua-Pongaroa Rd, NE of Makuri, 6.7km N of road JCT in Makuri, NZAC03000490, 40 33.575, 176 3.275, 22
Dec 2005, K Hill D M Marshall; WA, 1, ♀, 8.5 km N of Akitio Esplanade on Coast Rd, NZAC03000484, 40 33.597, 176
24.302, 22 Dec 2005, K Hill D M Marshall; WA, 1, ♀, 8.5 km N of Akitio Esplanade on Coast Rd, NZAC03000472, 40 33.597,
176 24.302, 22 Dec 2005, K Hill D M Marshall; WA, 1, ♀, 8.5 km N of Akitio Esplanade on Coast Rd, NZAC03000481, 40
33.597, 176 24.302, 22 Dec 2005, K Hill D M Marshall; WA, 1, ♀, Bideford, Waterfalls Rd, NZAC03004953, 40 51.645, 175
BUCKLEY ET AL.
480
·
Zootaxa 3900 (4) © 2014 Magnolia Press
52.361, 27 Jan 2008, TR Buckley R Leschen K Marske; WA, 1, ♀, Tinui, 3 km E of junction of Castle Point Rd and Tinui Vly
Rd, NZAC03005443, 40 51.949, 176 8.720, 26 Jan 2008, TR Buckley R Leschen K Marske; WA, 1, ♀, Donnelly's clearing, Mt
Holdsworth, NZAC03004963, 40 54.029, 175 28.090, 29 Jan 2008, TR Buckley R Leschen K Marske; WA, 1, ♀, Fensham
Scenic Reserve, Carterton, NZAC03005369, 40 59.594, 175 30.141, 27 Jan 2008, TR Buckley R Leschen K Marske; WA, 1, ♀,
Ngaumu State Forest, NZAC03004955, 41 01.693, 175 57.411, 27 Jan 2008, TR Buckley R Leschen K Marske; WA, 1, ♀,
Putangirua Pinnacles, Haurangi Mountains, NZAC03004925, 41 26.982, 175 14.453, 24 Jan 2008, TR Buckley R Leschen K
Marske; WA, 1, ♀, Putangirua Pinnacles, Haurangi Mountains, NZAC03004880, 41 26.982, 175 14.453, 24 Jan 2008, TR
Buckley R Leschen K Marske; WA, 1, ♀, Putangirua Pinnacles, Haurangi Mountains, NZAC03004970, 41 26.982, 175 14.453,
24 Jan 2008, TR Buckley R Leschen K Marske; WA, 1, Putangirua Pinnacles, Haurangi Mountains, NZAC03004994, 41
26.982, 175 14.453, 24 Jan 2008, TR Buckley R Leschen K Marske; WA, 1, ♀, Putangirua Pinnacles, Haurangi Mountains,
NZAC03004985, 41 26.982, 175 14.453, 24 Jan 2008, TR Buckley R Leschen K Marske; WA, 1, ♀, Whangamai State Forest,
S W Wairarapa, NZAC04030562, 11 Apr 1954, G W Ramsay; WA, 1, Waiohine Gorge Rd, NZAC03005238, 41 1.072, 175
24.077, 25 Jan 2008, TR Buckley R Leschen K Marske; WA, 1, Waiohine Gorge Rd, NZAC03005156, 41 1.072, 175 24.077,
25 Jan 2008, TR Buckley R Leschen K Marske; WA, 1, Wairongomai Station, Rimutaka Ra., NZAC03005350, 41 15.352, 175
08.779, 23 Jan 2008, TR Buckley R Leschen K Marske; WN, 1, ♀, Jacks Bush Rd, Waikanae, NZAC03004901, 40 51.6, 175
3.0, 21 Jan 2008, TR Buckley R Leschen K Marske; WN, 1, ♂, Jacks Bush Rd, Waikanae, NZAC03004909, 40 51.6, 175 3.0,
21 Jan 2008, TR Buckley R Leschen K Marske; WN, 1, ♂, Jacks Bush Rd, Waikanae, NZAC03004916, 40 51.6, 175 3.0, 21
Jan 2008, TR Buckley R Leschen K Marske; WN, 1, ♂, Jacks Bush Rd, Waikanae, NZAC03004872, 40 51.6, 175 3.0, 21 Jan
2008, TR Buckley R Leschen K Marske; WN, 1, Jacks Bush Rd, Waikanae, NZAC03005012, 40 51.6, 175 3.0, 21 Jan 2008,
TR Buckley R Leschen K Marske; WN, 1, Jacks Bush Rd, Waikanae, NZAC03005000, 40 51.6, 175 3.0, 21 Jan 2008, TR
Buckley R Leschen K Marske; WN, 1, ♀, 145 Creswick Tce, Northland, Wellington, NZAC03000343, 41 16.477, 174 45.128,
21 Mar 2004, TR Buckley; WN, 1, ♀, 145 Creswick Tce, Northland, Wellington, NZAC03000083, 41 16.477, 174 45.128, 20
Mar 2004, TR Buckley; WN, 1, ♀, Otari-Wiltons Bush, Wellington, NZAC03000115, 41 16.029, 174 45.449, 18 Mar 2005, TR
Buckley ; WN, 1, ♀, Otari-Wiltons Bush, Wellington, NZAC03000120, 41 16.029, 174 45.449, 18 Mar 2005, TR Buckley ;
WN, 1, ♂, Otari-Wiltons Bush, Wellington, NZAC03000371, 41 16.029, 174 45.449, 18 Mar 2005, TR Buckley; WN, 1, ♀,
Otari-Wiltons Bush, Wellington, NZAC03000382, 41 16.029, 174 45.449, 18 Mar 2005, TR Buckley ; WN, 1, Otari-Wiltons
Bush, Wellington, NZAC03000394, 41 16.029, 174 45.449, 18 Mar 2005, TR Buckley ; WN, 1, ♂, Otari-Wiltons Bush,
Wellington, NZAC03000402, 41 16.029, 174 45.449, 18 Mar 2005, TR Buckley ; WN, 1, ♀, Otari-Wiltons Bush, Wellington,
NZAC03000393, 41 16.029, 174 45.449, 18 Mar 2005, TR Buckley ; WN, 1, ♂, Otari-Wiltons Bush, Wellington,
NZAC03000387, 41 16.029, 174 45.449, 18 Mar 2005, TR Buckley ; WN, 1, ♀, Otari-Wiltons Bush, Wellington,
NZAC03000061, 41 16.029, 174 45.449, 18 Mar 2005, TR Buckley; WN, 1, ♂, Otari-Wiltons Bush, Wellington,
NZAC03000054, 41 16.029, 174 45.449, 18 Mar 2005, TR Buckley; WN, 1, ♀, Otari-Wiltons Bush, Wellington,
NZAC03000086, 41 16.029N, 174 45.449, 18 Mar 2005, TR Buckley; WN, 1, ♀, Francis Bell Res, Pt Howard, Wellington,
NZAC03000337, 41 15.074, 174 54.528, 15 Mar 2006, R Hoare; WN, 1, ♀, Smiths Creek Tk, Kaitoke, Rimutaka Forest Pk,
NZAC03000088, 41 4.206, 175 14.017, 19 Mar 2005, TR Buckley E J Buckley R G Buckley; WN, 1, Dom. Museum,
NZAC04030652, 9 Sep 1947; WN, 1, Dom. Museum, NZAC04030628, 9 Sep 1947; WN, 1, Khandallah, NZAC03005378,
Apr 1962, G Ramsay; WN, 1, ♀, Khandallah, Wellington, NZAC04030825, 25 Apr 1962, C McCann; WN, 7, Eggs,
Khandallah, Wellington, NZAC04030682, 25 Apr 1962, C McCann; WN, 1, ♀, Miramar Res., NZAC04030540, 9 May 1950,
G Ramsay; WN, 5, Eggs, nymphs, Miramar Res., NZAC04030670, 9 May 1950, G Ramsay; WN, 1, ♀, Miramar Hills,
NZAC04030600, 16 Jul 1947, G Ramsay; WN, 5, Eggs, Miramar, NZAC04030619, 16 Jul 1947, G Ramsay; WN, 1, ♀, Ngaio,
NZAC04030649, 6 Sep 1948, N Thompson; WN, 1, ♀, Wainui-o-mata, Wgtn, NZAC04030640, 12 Mar 1954, S Harding (ii);
WN, 1, ♀, Wainui-o-mata, Wgtn, NZAC04030638, 12 Mar 1955, S Harding (i); WN, 7, Eggs, Wainui-o-mata,
NZAC04030545, 12 Mar 1955, S Harding (i); WN, 1, ♀, Eastbourne, NZAC04030578, 5 May 1951, G Prichard; WN, 1, ♀,
Eastbourne, NZAC04030760, 5 May 1951, G Ramsay; WN, 1, ♀, Orongorongo River Valley, Rimutaka State Forest Pk,
NZAC03000372, 41 20.364, 174 58.733, 20 Mar 2006, TR Buckley E J Buckley R G Buckley; WN, 1, ♀, Orongorongo River
Valley, Rimutaka State Forest Pk, NZAC03000357, 41 20.364, 174 58.733, 20 Mar 2006, TR Buckley E J Buckley R G
Buckley; WN, 1, ♀, Orongorongo River Valley, Rimutaka State Forest Pk, NZAC03000033, 41 20.364, 174 58.733, 20 Mar
2006, TR Buckley E J Buckley R G Buckley; WN, 1, ♀, Orongorongo River Valley, Rimutaka State Forest Pk,
NZAC03000079, 41 20.364, 174 58.733, 20 Mar 2006, TR Buckley E J Buckley R G Buckley; WN, 1, ♀, Orongorongo River
Valley, Rimutaka State Forest Pk, NZAC03000068, 41 20.364, 174 58.733, 20 Mar 2006, TR Buckley E J Buckley R G
Buckley; WN, 1, ♀, Orongorongo River Valley, Rimutaka State Forest Pk, NZAC03000100, 41 20.364, 174 58.733, 20 Mar
2006, TR Buckley E J Buckley R G Buckley; WN, 1, ♀, Orongorongo River Valley, Rimutaka State Forest Pk,
NZAC03000064, 41 20.364, 174 58.733, 20 Mar 2006, TR Buckley E J Buckley R G Buckley; WN, 1, ♀, Orongorongo River
Valley, Rimutaka State Forest Pk, NZAC03000084, 41 20.364, 174 58.733E, 20 Mar 2006, TR Buckley E J Buckley R G
Buckley; WN, 1, ♀, Orongorongo River Valley, Rimutaka State Forest Pk, NZAC03000080, 41 20.364, 174 58.733, 20 Mar
2006, TR Buckley E J Buckley R G Buckley; WN, 1, ♀, Orongorongo , NZAC04030597, 25 Mar 1921;SD, 1, Okiwi Bay,
NZAC03005381, Jun 1984, T Jones;SD, 1, ♀, Trio Is (iii), NZAC04030655, 21 Dec 1954, G Ramsay;SD, 10, Trio Is (iii),
NZAC04030652, 21 Dec 1954, G Ramsay; SD, 1, ♀, Trio Is 1, NZAC04030661, 20 Dec 1954, G Ramsay; NN?, 1, ♀, Maori
Pah, NZAC04030742, 17 Feb 1924, E S Gourlay; NN?, 1, ♀, Maori Pah, NZAC04030796, 17 Feb 1924, E S Gourlay; NN?, 1,
♀, Totara Reserve, NZAC04030841, 1 Apr 1957, J M Hoy; NN, 1, Pakawau State Forest, N W Nelson, NZAC03005386, 27
May 1971, J S Dugdale; NN, 1, ♀, Brook Motor Camp, Nelson, NZAC03000368, 41 18.7, 173 17.54, 19 Feb 2004, J S
Zootaxa 3900 (4) © 2014 Magnolia Press
·
481
REVISION OF CLITARCHUS
Dugdale; NN, 1, ♀, Brightwater, Nelson, NZAC04030709, , C Lackner; NN, 1, ♀, Brightwater, Nelson, NZAC04030843, , C
Lackner; NN, 1, ♀, Nelson, NZAC04030631, 26 Mar 1949, G Ramsay; NN, 1, ♀, Pepin Id, NZAC04030747, 2 Mar 1924, E S
Gourlay; NN, 1, Nelson, Monaco, NZAC03005380, 10 Dec 1970, R Power; NN, 1, ♀, Paremata Reserve, Delaware Bay,
Waipuka Rivermouth, NZAC03000362, 41 9.414, 173 25.133, 6 Nov 2005, J S Dugdale; NN, 1, Maiti (Maitai?) River,
NZAC03005290, 28 Jun 1921; NN, 1, ♀, Maitai R, NZAC04030783, 17 Feb 1922; KA, 1, ♀,0.5 km S of carpark, Puhipuhi
Scenic Reserve, Kaikoura, NZAC03000347, 42 17.173, 173 43.824, 5 Apr 2004, TR Buckley R A B Leschen H Harman; KA,
1, ♀,0.5 km S of carpark, Puhipuhi Scenic Reserve, Kaikoura, NZAC03000381, 42 17.173, 173 43.824, 5 Apr 2004, TR
Buckley R A B Leschen H Harman; KA, 1, ♀, 1 km S of carpark, Puhipuhi Scenic Res, Kaikoura, NZAC03000056, 42 17.173,
173 43.824, 5 Apr 2005, TR Buckley R A B Leschen H Harman; KA, 1, ♀, Kaikoura, Puhipuhi Scenic Res, NZAC03000588,
42 17.173, 173 43.824, 15 Feb 2006, TR Buckley; KA, 1, ♀, Mt Fyffe, Fyffe Palmer Scenic Reserve, NZAC03004980, 42
20.162, 173 38.438, 5 Dec 2007, K Marske D Attanayake J Allwood; KA, 1, ♀, Mt Fyffe carpark, NZAC03000015, 42 21.087,
173 34.034, 5 Apr 2004, TR Buckley R A B Leschen H Harman; KA, 1, ♀, Mt Fyffe carpark, NZAC03000082, 42 21.087, 173
34.034, 5 Apr 2004, TR Buckley R A B Leschen H Harman; KA, 1, ♀, Mt Fyffe carpark, NZAC03000067, 42 21.087, 173
34.043, 5 Apr 2004, TR Buckley R A B Leschen H Harman; KA, 1, ♀, Mt Fyffe carpark, NZAC03000062, 42 21.087, 173
34.034, 5 Apr 2004, T R Bucklet R A B Leschen H Harman; KA, 1, Kekerenga Valley Rd Site 1, Kaikoura, NZAC03009374,
41 58.087 , 173 59.015, 16 Jan 2012, L Dunning R Leschen; KA, 2, Kekerenga Valley Rd Site 2, Kaikoura, NZAC03013707,
41 58.189 , 173 59.173 , 16 Jan 2012, L Dunning R Leschen; KA, 1, Mt Fyffe Stop 1, Kaikoura, NZAC03014854, 42 20.456 ,
173 34.154, 17 Jan 2012, L Dunning R Leschen; KA, 2, Mt Fyffe Stop 1, Kaikoura, NZAC03009519, 42 20.456 , 173 34.154,
17 Jan 2012, L Dunning R Leschen; NC, 1, ♀, Napenape Scenic Reserve, NZAC03000401, 42 56.568, 173 14.821, 12 Feb
2006, TR Buckley C Simon; NC, 1, ♀, Napenape Scenic Reserve, NZAC03000478, 42 56.568, 173 14.821, 12 Feb 2006, TR
Buckley C Simon; MC, 1, ♀, Kennedys Bush Scenic Reserve, Port Hills, Banks Peninsula, NZAC03000341, 43 37.893, 172
37.486, 13 Apr 2004, TR Buckley H Buckley B Case; MC, 1, ♀, Kennedys Bush Scenic Reserve, Port Hills, Banks Peninsula,
NZAC03000388, 43 37.893, 172 37.486, 13 Apr 2004, TR Buckley H Buckley B Case; MC, 1, ♀, Kennedys Bush Scenic
Reserve, Port Hills, Banks Peninsula, NZAC03000060, 43 37.893, 172 37.486, 13 Apr 2004, TR Buckley H Buckley B Case;
MC, 1, ♀, Kennedys Bush Scenic Reserve, Port Hills, Banks Peninsula, NZAC03000070, 43 37.893, 172 37.48, 13 Apr 2004,
TR Buckley H Buckley B Case; MC, 1, ♀, Kennedys Bush Scenic Reserve, Port Hills, Banks Peninsula, NZAC03000099, 43
37.893, 172 37.486, 13 Apr 2004, TR Buckley H Buckley B Case; MC, 1, ♀, Peel Forest, NZAC03000410, 43 53.952, 171
14.042, 29 Mar 2004, TR Buckley R A B Leschen ; MC, 1, ♀, Peel Forest, NZAC03000415, 43 53.952, 171 14.042, 29 Mar
2004, TR Buckley R A B Leschen ; MC, 1, ♀, Peel Forest, NZAC03000425, 43 53.49, 171 15.69, 3 Feb 2007, TR Buckley
RAB Leschen K Marske; MC, 1, ♀, Prices Valley, Banks Peninsula, NZAC03000356, 43 45.982, 172 42.876, 19 Feb 2006, TR
Buckley H L Buckley B Case; MC, 1, ♀, Banks Peninsula, Okuti Scenic Reserve, NZAC03000462, 43 46.994, 172 50.057, 6
Feb 2007, TR Buckley RAB Leschen K Marske J Sullivan; MC, 1, ♀, Peel Forest, NZAC03000608, 43 53.49, 171 15.69, 3 Feb
2007, TR Buckley, RAB Leschen, K Marske; MC, 1, ♀, Peel Forest, NZAC03000427, 43 53.49, 171 15.69, 3 Feb 2007, TR
Buckley, RAB Leschen, K Marske; DN, 1, ♀, Dunedin City Reservoir, Dunedin, NZAC03000389, 45 53.501, 170 27.118, 28
Mar 2004, TR Buckley T Jewell, R Morris; DN, 1, ♀, Dunedin City Reservoir, Dunedin, NZAC03000046, 45 53.501, 170
27.118, 28 Mar 2004, TR Buckley T Jewell R Morris; DN, 1, ♀, Dunedin City Reservoir, Dunedin, NZAC03000091, 45
53.501, 170 27.118, 28 Mar 2004, TR Buckley T Jewell R Morris; DN, 3, ♀, Waipori Falls Rd, first picnic area,
NZAC03000422, 45 55.788, 170 3.354, 31 Jan 2007, TR Buckley RAB Leschen K Marske; DN, 1, Waipori Falls Rd, first
picnic area, NZAC03005284, 45 55.737, 170 3.260, 29 Jan 2007, TR Buckley RAB Leschen K Marske; DN, 1, Waipori Falls
Rd, first picnic area, NZAC03005335, 45 55.737, 170 3.260, 29 Jan 2007, TR Buckley RAB Leschen K Marske; DN, 1,
Waipori Falls Rd, first picnic area, NZAC03005341, 45 55.737, 170 3.260, 29 Jan 2007, TR Buckley RAB Leschen K Marske;
SL, 1, ♂, Rongahere Gorge, NZAC04030746, 15 Dec 1995, J S Dugdale; WN, 1, ♀, Otari-Wiltons Bush, Wellington,
NZAC03000098, 41 16.029, 174 45.449, 18 Mar 2005, TR Buckley.
Clitarchus rakauwhakanekeneke sp. nov.
ND, nymphs, Poor Knights Is, Aorangi, NZAC04028286, 35 28.9, 174 44, 9 Jun 2006, TR Buckley, D Seldon; ND, 1, ♂,
Aorangi, Poor Knights Islands, NZAC03000504, 35 28.9, 174 44, 9 Jun 2006, TR Buckley D Seldon; ND, 1, ♂,Aorangi, Poor
Knights Islands, NZAC03000438, 35 28.9, 174 44, 9 Jun 2006, TR Buckley D Seldon; ND, 1, ♂, Aorangi, Poor Knights
Islands, NZAC03000463, 35 28 9, 174 44, 9 Jun 2006, TR Buckley D Seldon; ND, 1, ♂, Aorangi, Poor Knights Islands,
NZAC03000459, 35 28.9, 174 44, 9 Jun 2006, TR Buckley D Seldon; ND, 1, ♀,Aorangi, Poor Knights Islands,
NZAC03000456, 35 28.9, 174 44, 9 Jun 2006, TR Buckley D Seldon; ND, 2, ♀, Aorangi, Poor Knights Islands, Urupa Point
Camp, NZAC03005355, 13 Nov 1981, J C Watt; ND, 1, ♀,Poor Knights Is, Tawhiti Rahi, NZAC03000321, 4 Dec 1980, M F
Tocker; ND, 1, ♀,Poor Knights Is, Tawhiti Rahi, NZAC03005379, 3 Dec 1980, C F Butcher; ND, 1, ♀, Poor Knights Is,
Tawhiti Rahi, South track, NZAC03005313, 4 Dec 1980, M F Tocker; ND, 1, ♀,Poor Knights Is, Aorangi, Puweto Valley,
NZAC03005288, 11 Nov 1981, J S Dugdale; ND, 4, Tawhiti Rahi, Poor Knights Is, Main Trk, top of plateau, NZAC03009467,
35 27.457, 174 44.257, 17 Dec 2009, TR Buckley R Leschen D Seldon; ND, eggs, Tawhiti Rahi, Poor Knights Is, Main Trk, top
of plateau, NZAC03014396, 35 27.457, 174 44.257, 17 Dec 2009, TR Buckley R Leschen D Seldon; ND, eggs, Tawhiti Rahi,
Poor Knights Is, Main Trk, top of plateau, NZAC03014416, 35 27.457, 174 44.257, 17 Dec 2009, TR Buckley R Leschen D
Seldon; ND, eggs, Tawhiti Rahi, Poor Knights Is, Main Trk, top of plateau, NZAC03014405, 35 27.457, 174 44.257, 17 Dec
2009, TR Buckley R Leschen D Seldon.
BUCKLEY ET AL.
482
·
Zootaxa 3900 (4) © 2014 Magnolia Press
Clitarchus tepaki sp. nov.
ND, 1, ♂, North Cape Scientific Reserve, NZAC03004892, 34 24.467, 173 0.431, 18 Dec 2007, TR Buckley D Seldon R
Hoare; ND, 1, ♂, North Cape Scientific Reserve, NZAC03005526, 34 24.467, 173 0.431, 18 Dec 2007, TR Buckley D Seldon
R Hoare; ND, 1, ♂,North Cape Scientific Reserve, NZAC03004886, 34 24.467, 173 0.431, 18 Dec 2007, TR Buckley D Seldon
R Hoare; ND, 1, ♂,North Cape Scientific Reserve, NZAC03000671, 34 24.467, 173 0.431, 18 Dec 2007, TR Buckley D Seldon
R Hoare; ND, 1, ♂, North Cape Scientific Reserve, NZAC03004774, 34 24.467, 173 0.431, 18 Dec 2007, TR Buckley D
Seldon R Hoare; ND, 1, ♀, North Cape Scientific Reserve, NZAC03004887, 34 24.467, 173 0.431, 18 Dec 2007, TR Buckley
D Seldon R Hoare; ND, 1, ♂, North Cape Scientific Reserve, NZAC03004922, 34 24.467, 173 0.431, 18 Dec 2007, TR
Buckley D Seldon R Hoare; ND, 1, eggs, North Cape Scientific Reserve, NZAC03004883, 34 24.467, 173 0.431, 18 Dec 2007,
TR Buckley D Seldon R Hoare; ND, 1, ♀, North Cape Scientific Reserve, NZAC03004769, 34 24.467, 173 0.431, 18 Dec
2007, TR Buckley D Seldon R Hoare; ND, 1, ♀,North Cape Scientific Reserve, NZAC03005018, 34 24.467, 173 0.431, 18 Dec
2007, TR Buckley D Seldon R Hoare; ND, 1, Tom Bowling Bay, Waitangi, NZAC03005307, 3 Dec 1977, L L Deitz; ND, 1,
Tom Bowling Bay, Waitangi, NZAC03005340, 3 Dec 1977,S Murray; ND, 1, Tom Bowling Bay, Waitangi, NZAC03005370, 3
Dec 1977, L L Deitz; ND, 1, Unuwhao, Spirits Bay, NZAC03005327, 18 Jan 1966,A K Walker; ND, 1, ♀,Spirits Bay,
NZAC03004967, 34 25.801, 172 52.002, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂,Spirits Bay, NZAC03004951,
34 25.801, 172 52.002, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂, Spirits Bay, NZAC04030733, 10 Jan 1957,R A
C (Cumber); ND, 1, Spirits Bay, NZAC04030730, 10 Jan 1957,R A C (Cumber); ND, 5, Waitanoni Stream, Spirits Bay,
NZAC03005358, 7 Nov 1967, J G R McBurney; ND, 1, ♂,Waikuku Flat, Tom Bowling Bay, NZAC03004957, 34 27.383, 172
59.241, 18 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, Te Paki Trig Track, NZAC03000469, 34 28.139, 172 46.675, 12
Feb 2005, D Seldon; ND, 1, Te Paki Trig Track, NZAC03005316, 22 Nov 1982, E W Valentine; ND, 1, ♂, Kauri Bush, Te Paki,
NZAC03005448, 34 28.350, 172 45.865, 9 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1, ♀, Kauri Bush, Te Paki,
NZAC03005450, 34 28.350, 172 45.865, 9 Dec 2008, TR Buckley R Leschen D Seldon; ND, 1, ♂, Kauri Bush Trk, Te Paki,
NZAC03004869, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂, Kauri Bush Trk, Te Paki,
NZAC03004870, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♀, Kauri Bush Trk, Te Paki,
NZAC03004941, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♀, eggs, Kauri Bush Trk, Te
Paki, NZAC03005524, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂, Kauri Bush Trk, Te
Paki, NZAC03005525, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂, Kauri Bush Trk, Te
Paki, NZAC03004946, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♀, Kauri Bush Trk, Te
Paki, NZAC03004910, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂, Kauri Bush Trk, Te
Paki, NZAC03004899, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, Eggs, Kauri Bush Trk, Te
Paki, NZAC03004810, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂, Kauri Bush Trk, Te
Paki, NZAC03004827, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂, Kauri Bush Trk, Te
Paki, NZAC03004961, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♀, Kauri Bush Trk, Te
Paki, NZAC03004979, 34 28.391, 172 45.842, 19 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♀, Trk from Papawiri to
Kohuranaki, Te Paki Rec Res, NZAC03005470, 34 29, 172 49, 20 Oct 2008, R Hoare; ND, 1, ♀, Kohuranaki, Te Paki,
NZAC03004940, 34 29.484, 172 50.191, 21 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂, Kohuranaki, Te Paki,
NZAC03004975, 34 29.484, 172 50.191, 21 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂, Kohuranaki, Te Paki,
NZAC03004956, 34 29.484, 172 50.191, 21 Dec 2007, TR Buckley D Seldon R Hoare; ND, 1, ♂, Karatia Rd,
NZAC03005473, 34 33.565, 172 50.263, 5 Nov 2008, TR Buckley R Leschen; ND, 1, ♂, Karatia Rd, NZAC03005449, 34
33.565, 172 50.263, 5 Nov 2008, TR Buckley R Leschen; ND, 1, ♀, Karatia Rd, NZAC03005497, 34 33.565, 172 50.263, 5
Nov 2008, TR Buckley R Leschen; ND, 1, ♂, Karatia Rd, NZAC03005444, 34 33.565, 172 50.263, 5 Nov 2008, TR Buckley R
Leschen; ND, 1, Parengarenga, Paua, NZAC03005297, 18 Jan 1974, M J Esson; ND, 6, Parengarenga, Paua, NZAC03005282,
21 Jan 1975, M J Esson; ND, 3, Parengarenga, Paua, NZAC03005281, 20 Jan 1975, M J Esson; ND, 1, Whangatupere Bay,
Karikari Peninsula, NZAC03015367, 34 50 05.8, 173 26 19.2, 18 Jan 2013, TR Buckley R Leschen D Seldon; ND, 1,
Whangatupere Bay, Karikari Peninsula, NZAC03015234, 34 50 05.8, 173 26 19.2, 18 Jan 2013, TR Buckley R Leschen D
Seldon; ND, 1, Whangatupere Bay, Karikari Peninsula, NZAC03015240, 34 50 05.8, 173 26 19.2, 18 Jan 2013, TR Buckley R
Leschen D Seldon; ND, 1, Whangatupere Bay, Karikari Peninsula, NZAC03015312, 34 50 05.8, 173 26 19.2, 18 Jan 2013, TR
Buckley R Leschen D Seldon; ND, 1, Whangatupere Bay, Karikari Peninsula, NZAC03015233, 34 50 05.8, 173 26 19.2, 18 Jan
2013, TR Buckley R Leschen D Seldon; ND, 1, Whangatupere Bay, Karikari Peninsula, NZAC03015290, 34 50 05.8, 173 26
19.2, 18 Jan 2013, TR Buckley R Leschen D Seldon; ND, 1, Whangatupere Bay, Karikari Peninsula, NZAC03015401, 34 50
05.8, 173 26 19.2, 18 Jan 2013, TR Buckley R Leschen D Seldon; ND, 1, Whangatupere Bay, Karikari Peninsula,
NZAC03015243, 34 50 05.8, 173 26 19.2, 18 Jan 2013, TR Buckley R Leschen D Seldon; ND, 20+, Eggs,Whangatupere Bay,
Karikari Peninsula, NZAC03015244, 34 50 05.8, 173 26 19.2, 18 Jan 2013, TR Buckley R Leschen D Seldon.
... Two species of stick insect from the genus Clitarchus are found on the Te Hiku o Te Ika peninsula in northern New Zealand. Clitarchus tepaki is limited to the northern tip, while Clitarchus hookeri covers the southern region of Te Hiku o Te Ika (Buckley et al. 2010b;Buckley et al. 2014;Myers et al. 2017) (Figure 1). We consider these to be valid species because they show easily diagnosable differences in the morphology of the genitalia, characters which are strongly associated with reproductive isolation in insects (Buckley et al. 2014). ...
... Clitarchus tepaki is limited to the northern tip, while Clitarchus hookeri covers the southern region of Te Hiku o Te Ika (Buckley et al. 2010b;Buckley et al. 2014;Myers et al. 2017) (Figure 1). We consider these to be valid species because they show easily diagnosable differences in the morphology of the genitalia, characters which are strongly associated with reproductive isolation in insects (Buckley et al. 2014). They also show significant genetic differentiation at mitochondrial and nuclear loci indicative of at least partially independent evolution histories (Buckley et al. 2010b;Myers et al. 2017). ...
... tepaki contain 2-4 teeth with an enlarged anterior pair that overlaps (Buckley et al. 2014). ...
Article
Documenting natural hybrid systems builds our understanding of mate choice, reproductive isolation, and speciation. The stick insect species Clitarchus hookeri and C. tepaki differ in their genital morphology and hybridize along a narrow peninsula in northern New Zealand. We utilize three lines of evidence to understand the role of premating isolation and species boundaries: (1) genetic differentiation using microsatellites and mitochondrial DNA; (2) variation in 3D surface topology of male claspers and 2D morphometrics of female opercular organs; and (3) behavioral reproductive isolation among parental and hybrid populations through mating crosses. The genetic data show introgression between the parental species and formation of a genetically variable hybrid swarm. Similarly, the male and female morphometric data show genital divergence between the parental species as well as increased variation within the hybrid populations. This genital divergence has not resulted in reproductive isolation between species, instead weak perimating isolation has enabled the formation of a hybrid swarm. Behavioral analysis demonstrate that the entire mating process influences the degree of reproductive isolation between species undergoing secondary contact. Mechanical isolation may appear strong, while perimating isolation is weak. This article is protected by copyright. All rights reserved.
... The stick insect genus Clitarchus St al is comprised of three species: Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler, found on the Poor Knights Islands (Buckley et al., 2014); C. tepaki Buckley, Myers and Bradler, which is restricted to the tips of New Zealand's most northern peninsulas (Aupouri Peninsula and Karikari Peninsula, Fig. 1) (Buckley et al., , 2014; and Clitarchus hookeri White, found throughout the North Island, south of the Aupouri and Karikari peninsulas, and parts of the South Island (Buckley et al., , 2014Morgan-Richards et al., 2010). Clitarchus tepaki was recently described on the basis of male and female genitalia, where this species has enlarged genital claspers compared with C. hookeri (Buckley et al., 2014). ...
... The stick insect genus Clitarchus St al is comprised of three species: Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler, found on the Poor Knights Islands (Buckley et al., 2014); C. tepaki Buckley, Myers and Bradler, which is restricted to the tips of New Zealand's most northern peninsulas (Aupouri Peninsula and Karikari Peninsula, Fig. 1) (Buckley et al., , 2014; and Clitarchus hookeri White, found throughout the North Island, south of the Aupouri and Karikari peninsulas, and parts of the South Island (Buckley et al., , 2014Morgan-Richards et al., 2010). Clitarchus tepaki was recently described on the basis of male and female genitalia, where this species has enlarged genital claspers compared with C. hookeri (Buckley et al., 2014). ...
... The stick insect genus Clitarchus St al is comprised of three species: Clitarchus rakauwhakanekeneke Buckley, Myers and Bradler, found on the Poor Knights Islands (Buckley et al., 2014); C. tepaki Buckley, Myers and Bradler, which is restricted to the tips of New Zealand's most northern peninsulas (Aupouri Peninsula and Karikari Peninsula, Fig. 1) (Buckley et al., , 2014; and Clitarchus hookeri White, found throughout the North Island, south of the Aupouri and Karikari peninsulas, and parts of the South Island (Buckley et al., , 2014Morgan-Richards et al., 2010). Clitarchus tepaki was recently described on the basis of male and female genitalia, where this species has enlarged genital claspers compared with C. hookeri (Buckley et al., 2014). ...
Article
Aim Comparisons of independent secondary contact events provide insights into allopatric speciation. The geographic distribution of Clitarchus stick insects across northern New Zealand corresponds to islands that existed in the Pliocene. Subsequent joining to the mainland provided multiple opportunities for secondary contact. We address the hypothesis that secondary contact has similar effects on parental species, across two peninsulas, due to the simultaneous removal of allopatric barriers. Location Northland, New Zealand Methods We explore how genetic structure relates to morphological variation and biogeographical history of Clitarchus hookeri and C. tepaki . Mitochondrial DNA sequences were used to reconstruct phylogenetical relationships. Population structure was estimated using 14 microsatellite markers. We analysed genital morphology and genetic structure within a biogeographical context to identify the consequences of independent secondary contact events. Results Geographical patterns of genital shape indicate two independent hybrid swarms occur between C. hookeri and C. tepaki . Haplotype distributions and Bayesian cluster analyses reveal C. tepaki from the Aupouri Peninsula are distinguished from C. hookeri on the basis of both genetic structure and morphology. Clitarchus tepaki on the Karikari Peninsula did not group phylogenetically or through Bayesian clustering with C. tepaki from the Aupouri Peninsula. We observed a hybrid swarm along the Aupouri Peninsula. Main conclusions Clitarchus species were separated on Pliocene islands and then subsequently made independent secondary contact as land connections formed. Clitarchus tepaki on the Aupouri Peninsula maintained genetic integrity; whereas we hypothesize genetic swamping has occurred on Karikari Peninsula. Our data demonstrate different outcomes of independent secondary contact events.
... During the summer, the endemic New Zealand stick insect species Clitarchus hookeri (White) is frequently encountered in mating pairs on Leptospermum scoparium and Kunzea sp. plants (Buckley et al., 2010a;Morgan-Richrads et al., 2010;Buckley et al. 2014). Natural populations are characterized as scramble competitors that exhibit prolonged mate guarding when female density is low (Myers et al. 2015). ...
... The genitalia of male C. hookeri comprise a genital bulge encompassing a complex internal phallus covered by an operculum (shown Fig. 1). A modification of tergite X forms a pair of claspers which curve behind the cerci, the claspers contain three to five pairs of teeth that are used for clasping onto the females operculum (Stringer 1970;Buckley et al. 2014). To initiate mating males approach females, climb onto their dorsal surface, tuck their abdomens under the female to attach their claspers onto the base of the female operculum, and remain attached for prolonged intervals well beyond the time required for spermatophore transfer (Myers et al. 2015). ...
... To initiate mating males approach females, climb onto their dorsal surface, tuck their abdomens under the female to attach their claspers onto the base of the female operculum, and remain attached for prolonged intervals well beyond the time required for spermatophore transfer (Myers et al. 2015). The opercular organ of female C. hookeri is only weakly sclerotized where the claspers attach, unlike other Clitarchus species that possess flanking lateral sclerotized patches (Buckley et al. 2014). This system is unique to the Lanceocercata whereas in other phasmids, the vomer functionally replaces the clasper (Bradler 2001(Bradler , 2009Buckley et al. 2010b). ...
Article
Full-text available
In animals with internal fertilization, male genitalia exhibit higher rates of divergence compared with other morphological trails. Recent evidence suggests sexual selection drives such as rapid and divergent trait evolution. External male genital structures which clasp or stimulate the female’s exterior are likely to be under similar selective constraints to internal genitalia; however, their function and influence on male mating success have rarely been studied in detail. Here, we modify the external genitalia of the phasmid Clitarchus hookeri (White) to assess the role of male claspers in achieving successful acceptance by females and subsequent copulation. By covering female opercular organs and abrading male claspers, we demonstrate the necessity of precise coupling between these external genitalic structures for copulation to take place. We found that modified females tolerate un-modified male clasping attempts up to four times longer than normally required for attachment. However, when un-modified females are contacted by modified male claspers, males are quickly rejected. Our results suggest that external genital structures play an important role in precopulatory mate acceptance. Here, we discuss the potential role of female choice and species, isolating hypotheses in explaining the high evolutionary rate of such structures. Significance statement Many male animals possess genital structures that allow them to grip on to females before, during, and after mating. We experimentally manipulated male claspers and the corresponding female morphology that is clasped by males for the stick insect Clitarchus hookeri, resulting in clear changes in mating behavior. We show that female mate acceptance is influenced by both the structure of male claspers and the ability of females to perceive clasping. This demonstrates that external genital structures can play an important role in precopulatory mate acceptance and that female choice is likely to influence their evolution.
... Therefore, the majority of collectors focussed on the South Island, which was deemed to have a wider range of unique habitats than the North Island. However, recent studies have concluded that there are 16 regionally endemic Mecodema species (Table 1) in Northland (Seldon and Leschen 2011;Seldon et al. 2012;Seldon and Buckley 2019) and that this high endemism matches that of other taxa, including stick insects, ground beetles, plants, etc. (de Lange et al. 2002;de Lange and Rolfe 2008;Buckley and Bradler 2010;Buckley et al. 2014;Larochelle and Larievière 2016). ...
... Although, we found evidence of deeply diverging haplotypes in the far North, we did not find evidence for reciprocally monophyletic haplotype clades or a speciation event involving the population from the former island of Te Paki, unlike studies of other invertebrate species (e.g. Morgan-Richards, Trewick & Wallis, 2001;Buckley & Leschen, 2013;Buckley et al., 2014;Myers, Holwell & Buckley, in press). Many of the invertebrate species that are endemic to this area are flightless, unlike L. barbicornis, and ongoing gene flow may have prevented population divergence here. ...
Article
Historic geological and climatic events such as tectonic activity, volcanism and glaciation have significantly impacted the distribution and genetic signature of organisms. In New Zealand, glacial cycles during the late Pliocene and throughout the Pleistocene caused the loss of large areas of forest habitat and many species are proposed to have been restricted to refugia during this period. Here we use mitochondrial DNA (COI, COII) to reconstruct phylogenetic relationships between haplotypes in association with several hypothetical biogeographical lines to determine the impact that past climatic conditions had on the giraffe weevil (Lasiorhynchus barbicornis), an endemic brentine weevil found in native forests across New Zealand. Our analyses revealed that populations in northern New Zealand have higher genetic diversity than those in the south. We compared evidence for genetic structure in relation to four biogeographical lines: Northland Line, Kauri Line, Taupo Line and Cockayne’s Line. Although, there was significant difference in genetic variation north and south of all the proposed biogeographic lines, we found most support for the Kauri Line. These results support our hypothesis that giraffe weevils had a restricted range during the Pleistocene, predominantly pushed north into refugia, which was followed by range expansion following glacial retreat.
Article
During hybrid speciation, homoeologues combine in a single genome. Homoeologue expression bias (HEB) occurs when one homoeologue has higher gene expression than another. HEB has been well characterized in plants but rarely investigated in animals, especially invertebrates. Consequently, we have little idea as to the role that HEB plays in allopolyploid invertebrate genomes. If HEB is constrained by features of the parental genomes, then we predict repeated evolution of similar HEB patterns among hybrid genomes formed from the same parental lineages. To address this, we reconstructed the history of hybridization between the New Zealand stick insect genera Acanthoxyla and Clitarchus using a high‐quality genome assembly from Clitarchus hookeri to call variants and phase alleles. These analyses revealed the formation of three independent diploid and triploid hybrid lineages between these genera. RNA sequencing revealed a similar magnitude and direction of HEB among these hybrid lineages, and we observed that many enriched functions and pathways were also shared among lineages, consistent with repeated evolution due to parental genome constraints. In most hybrid lineages, a slight majority of the genes involved in mitochondrial function showed HEB towards the maternal homoeologues, consistent with only weak effects of mitonuclear incompatibility. We also observed a proteasome functional enrichment in most lineages and hypothesize this may result from the need to maintain proteostasis in hybrid genomes. Reference bias was a pervasive problem, and we caution against relying on HEB estimates from a single parental reference genome.
Article
Full-text available
Some animal groups, such as stick insects (Phasmatodea), have repeatedly evolved alternative reproductive strategies, including parthenogenesis. Genomic studies have found modification of the genes underlying meiosis exists in some of these animals. Here we examine the evolution of copy number, evolutionary rate and gene expression in candidate meiotic genes of the New Zealand geographic parthenogenetic stick insect Clitarchus hookeri. We characterised 101 genes from a de novo transcriptome assembly from female and male gonads that have homology with meiotic genes from other arthropods. For each gene we determined copy number, the pattern of gene duplication relative to other arthropod orthologues and the potential for meiosis-specific expression. There are five genes duplicated in C. hookeri, including one also duplicated in the stick insect Timema cristinae, that are not or uncommonly duplicated in other arthropods. These included two sister chromatid cohesion associated genes (SA2 and SCC2), a recombination gene (HOP1), an RNA-silencing gene (AGO2) and a cell cycle regulation gene (WEE1). Interestingly, WEE1 and SA2 are also duplicated in the cyclical parthenogenetic aphid Acyrthosiphon pisum and Daphnia duplex, respectively, indicating possible roles in the evolution of reproductive mode. Three of these genes (SA2, SCC2 and WEE1) have one copy displaying gonad-specific expression and all genes, with the exception of WEE1, have significantly different non-synonymous/synonymous ratios between the gene duplicates, indicative of a shift in evolutionary constraints following duplication. These results suggest that stick insects may have evolved genes with novel functions in gamete production by gene duplication.
Article
Full-text available
The subfamily Platycraninae Brunner v. Wattenwyl, 1893 is polyphyletic in the traditional treatment. None of the genera attributed to the subfamily is closely related to the type-genus Platycrana Gray, 1835, which has already been suggested by morphological studies. The main key feature of Platycraninae traditionally used to characterize the subfamily, the remarkably large head and strongly enlarged genae ("cheeks"), is not true for Platycrana. Consequently, all other genera, the true "Palm Stick Insects" need to be removed from Platycraninae and are here accommodated in Megacraniinae subfam. nov.. Megacrania Kaup, 1871 is chosen as the type-genus of this new subfamily, because the name very well describes the main characteristic of this clade. Previous molecular studies have shown Megacraniinae subfam. nov. to be a member of the Old World family Phasmatidae s. str. (= Lanceocercata Bradler, 2001), which is characterized by the enlarged, laterally flattened and foliaceous or lanceolate cerci and by the male anal segment longitudinally split into two movable, interiorly dentate hemi-tergites that serve as a clasping apparatus to grasp the female abdomen during copulation and makes lacking a vomer. Neither character is true for Platycrana and Platycraninae sensu nov. respectively. Molecular studies have revealed Platycraninae sensu novo as the sister group of Phasmatidae s. str. (= Lanceocercata). The tribe Stephanacridini Günther, which has previously been without a subfamilar affiliation is transferred as a subordinate taxon of Platycraninae sensu novo. Platycraninae is characterized and distinguished from Phasmatidae s. str. (= Lanceocercata) by the small cylindrical cerci, simple anal segment and presence of a vomer in males, as well as the strongly keeled subgenital plate and remarkably elongated, filiform gonapophysis VIII of females, which greatly project over the apex of the abdomen. Characterizations are presented for Megacraniinae subfam. nov. as well as Platycraninae sensu nov. and its two tribes Platycranini and Stephanacridini. Lists of genera are provided for the three taxa. Three genera are removed from Platycraninae and Megacraniinae subfam. nov.: The Australian Echetlus Stål, 1875 is transferred to Phasmatinae: Acanthomimini and Redtenbacherus Özdikmen & Darilmaz, 2008 and Elicius Günther, 1935 are transferred to Lonchodidae: Necrosciinae. The male of Megacrania spina Hsiung, 2007 as well as the eggs of Megacrania brocki Hsiung, 2002, Erastus apalamnus Rehn, 1904, Ophicrania apterus (Redtenbacher, 1908), Ophicrania nigroplagiatus (Redtenbacher, 1908), Platycrana viridana (Olivier, 1792) and Macrophasma lyratus (Redtenbacher, 1908) are illustrated for the first time. Xenomaches moderata (Kirby, 1884) from the island of Ambon is synonymised with Platycrana viridana (Olivier, 1792) (syn. nov.).
Chapter
Full-text available
Stick and leaf insects (order Phasmatodea) are a mesodiverse lineage of large terrestrial herbivores with predominantly tropical distribution and few species inhabiting more temperate regions. The phylogenetic position of the Phasmatodea among the lower neopteran insects has been debated for many years, with basically every orthopteroid insect order proposed as the potential sister taxon. The stick and leaf insects exhibit a remarkably poor fossil record. The fascinating and variable biology of stick insects has made them excellent model systems for investigating a number of evolutionary phenomena, including speciation and reproductive isolation, evolution of parthenogenesis and alternative reproductive strategies, and more recently the evolution of cold tolerance. Evidence for monophyletic Phasmatodea is undisputed and has grown stronger in recent years, with evidence coming from various sources. The chapter lists and discusses the currently recognized, non‐encaptic monophyletic groups. The contributions of amateur taxonomists play a crucial role in describing the phasmatodean diversity.
Book
Full-text available
A synonymic catalog of species, essential for researchers. 414 pages, spiral bound [replaces a CD issued in 2003 (First Edition)] OUT OF PRINT
Article
Full-text available
The endemic New Zealand cyclophoroidean genus Cytora is revised. A total of 42 species are recognised, of which 23 are described as new. All taxa are described and illustrated with colour photographs and scanning electron microscope images.
Article
Many of the rocks of this area weather readily to clay and the steep slopes and deep subsoils cause widespread instability. Volcanic activity occurred in the late Tertiary and again in the Auckland area in the Quaternary. The coast is described, including problems of coastal instability; the effects of man on the geomorphology of the coast and on the inland area conclude the chapter.-K.Clayton
Article
Approximately 263 species of native vascular plants are listed for four forest remnants in northern-most New Zealand (c. latitude 34°30'S). The lack of tropical and endemic species in this flora may be the result of extinction in cooler Pleistocene times.
Article
Although hybridisation is common in animals, it rarely results in speciation. Yet, many examples of hybrid species have been documented in one animal group, the stick insects (Phasmida). The N ew Z ealand stick insect A canthoxyla is of particular interest as the entire genus is of hybrid origin and consists of eight morphological forms recognised as species, all of which are obligate parthenogens. Using five complementary techniques on the same individuals, our study confirms that both triploids and diploids are present in A canthoxyla populations, and further, that some individuals contain both diploid and triploid cells. Chromosome spreads and estimates of relative DNA content from flow cytometry provided contrasting information about the ploidy of this unusual parthenogenetic genus. Analysis of morphometric variation showed no correlation with ploidy level in A canthoxyla , and also mtDNA sequence networks failed to distinguish morphospecies or ploidy level. Unexpectedly, cloned sequences of a putatively single‐copy nuclear marker were also unhelpful in distinguishing ploidy, instead indicating that phosphoglucose isomerase is likely to be a multiple copy gene. We propose a mechanism for the evolution of the Acanthoxyla lineage and suggest that interpretation may be complicated by the presence of individuals that are diploid and triploid mosaics.