Fauna of New Zealand Ko te Aitanga Pepeke o Aotearoa
Hodgson, C. J.; Henderson, R. C. 2000: Coccidae (Insecta: Hemiptera: Coccoidea). Fauna of New Zealand 41, 264 pp. I E E A E SYS EMA ICS A ISO Y G OU
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SE IES E I O . K. C os y nd r r h M nt Alb rt r h C ntr r v t 2 0, A l nd, l nd Fauna of New Zealand Ko te Aitanga Pepeke o Aotearoa
Number / Nama 41
Coccidae (I sec a emi e a Coccoi ea
C. J. Hodgson Wye Co ege U i e si y o o o Wye ea As o Ke ΤΝ 5 5ΑΗ U K ( ese a ess e a me o io i e si y a Sys ema ic io ogy a io a Museum o Wa es Ca i C 1 3ΝΡ Wa es U K o gso C ca i ac uk
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Ca a ogui g i u ica io O GSO C (C is o e o Coccidae (Insecta: Hemiptera: Coccoidea) / C o gso a C Henderson
— Lincoln, Canterbury, N.Z. : Manaaki Whenua Press, 2000. (Fauna of New Zealand, ISSN 0111-5383 ; no. 41). ISBN 0-478-09335-7 I. Henderson, Rosa Constance II. Title III. Series UDC 595.752.3(931)
Prepared for publication by the series editor using computer-based text processing, layout, scanning, and printing at Landcare Research, Mt Albert Research Centre, Private Bag 92170, Auckland, New Zealand
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Published by Manaaki Whenua Press, Landcare Research, P.O. Box 40, Lincoln, Canterbury, N.Z. Website: http://www.mwpress.co.nz/
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Front cover: Cr t ll t t rn t ll Henderson & Hodgson. (Illustrator: Chris Hodgson).
u ica io o e Fauna of New ZeaLand se ies is e esu o a esea c i es me y e ou a io o esea c Scie ce a ec o ogy u e co ac um e C 9 17
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O U A SUMMA Y E W AKA Ā O O O A GA
C ass Insecta O e Hemiptera Su o e Stenorrhyncha Su e ami y Coccoidea ami y Coccidae I us a io W akaa ua: Cr t ll t t l pt p r So sca e i sec s (Maske (I us a o Kaiw akaa ua: C is o gso . The soft scale family Coccidae is one of ten families of plant-sucking scale insects present in New Zealand. The Ko e w ā au Cocci ae e w ā au "u a i mo e" Ko ia 57 known species can be divided into two distinct groups: ē a i o gā w ā au 1 o gā gā a a u a i e go go 43 indigenous species in 11 genera that are restricted to i u ā ā ou ma i e o o a a i Ao ea oa Ka o o mai New Zealand, and 14 cosmopolitan (or adventive) species gā momo e 57 e mō io ia a a ki ē a i ka a ga a ga w ā ui in six genera that are generally of world-wide distribution, e ua E 3 gā momo kei e ka a ga a ga ua a i ō gā and probably arrived here accidentally on imported plant u i ga 11 ā ko Ao ea oa a ake e wā i e ki ea ai ē ei material. Most of the indigenous species are quite closely ai a ga a u ga Kei e ui ga ua ua ko ē a i momo 1 related and are covered in a glassy wax cover ō gā u i ga e ka ki ea i a w e ua ma a o u o e characteristically divided into polygonally-shaped plates. ao kua ae ma e e mai ā ei ki ko ei o o ai E No Maori names are known for any of these insects. w aka ae ia a a i au oke e ū mai ē ei ki Ao ea oa mā The native species of soft scales are associated with u ga ί ē a i i u i uku a mai i āwā i Κo e ui ga o gā native trees and shrubs and there is only a single record of momo ō ko ei ake ake ake e āka ga ki i a i ā ka oa a native species found on any of the numerous introduced ka oa kua ka i ί ē a i ki i mō i u i u a gaia ai e ki i plant species. Many of these native scale species have ei ki ē a i wā a ga i i e a a au e ā ua Kāo e e mō io ia quite wide host-ranges whilst others appear to be more a a gā i goa Māo i mō ē ei mo oi i host-specific. The closeness of the association between Κo gā momo ō Ao ea oa ū u u ki ea ai e i i a i the indigenous species and the New Zealand flora, on a a ki gā ākau me gā mauw a ū u u o Ao ea oa E ai which there are no definite records of serious damage, ki gā kō e o kua ko ia mō ē ei u a i mo e ko a i a ake suggests that the association between the scale and its host e ki e ga o ē a i e i i a a ki ē a i ākau i akea mai i plants has had a long evolutionary history. None of these āwā i Κo ē a i o gā u a i mo e ake ake ake o o a i native species are of economic importance, and none has ai ki e u ua o gā momo ākau ko ē a i a ō ka i i a i been recorded outside New Zealand. ki e momo ākau ko a i ei e o oka a o u gā o o i Most of the introduced or adventive species are of wae ga ui gā u a i mo e ake ake ake me gā akau some economic importance, mainly on introduced crops ake ake ake e ga i kāo e a ō kia ki ea ē a i ā ga i o and cultivated garden plants, both indoors and outdoors. ki o o e o o a u a gā u a i mo e ki aua ākau ō ei a Two species of wax scales require control on citrus, whilst e w aka ae ia a a ō ua w aka e e e o o a i me e other species are occasionally significant on plums, kuku e a i a ē ei o gā ama iki a ā e Kāo e e ā ga apricots, and cherries. Biological control agents were ki o a a mo i ei o gā momo o Ao ea oa ā kāo e a imported earlier to control two of the pest scales. One kia ki ea ē a i e o o a a i ē a i a u w e ua o e ao hymenopterous parasitoid species was introduced in 1921 Ko e ui ga o gā momo kua kawea mai i āwā i e to control brown soft scale (Coccus hesperidum ā ga ki o ō ā ou ki gā i u i mau ia mai i āwā i ki Linnaeus), and four species have been imported against ē a i i u oki o o o i ā ā ou mā a ae a u ki gā i u black scale (Saissetia oleae (Olivier)). Generally the w aka ai ai ka o o ki ō w a e A ā ē a i momo u a i
( nt n d v rl f (h r t n
6 d n & nd r n (2000 : C d (In t : pt r : C d cosmopolitan scales in New Zealand are each attacked by a a e e ua eā au o o a i a a ki gā ākau o e w several species of parasitoids. ēma a ka mā ua a ua e e a ga a kia ko e e a u aua Like many other related sap-sucking insects, the soft ākau A ā a ō ē a i ka w aka a u i gā ākau a amu scales eliminate honeydew from the anus. This is a sugary ā e eko i ae a u ki e e e e mea iki a u oki ē a i solution produced in the gut after feeding on the plant sap. gā a a i āwā i ei a u i ē a i o gā u a i ei ē ā ē a i A black sooty mould fungus often grows on the honeydew. momo gā a a ō e ū oi yme o e a ko e i i oa ō a In most parts of the world, this honeydew is attractive to i e i mau ia mai ki Ao ea oa i e au 19 1 ko a a ma i ants, but in New Zealand only a single native species is e a u i e u a i mo e a au i (C h p r d known to be ant-attended — the reasons for this are i aeus āa ā a ō ē a i momo e w ā kua mau ia mai unknown. However, soft scales are rarely as abundant as ei ā āmi i e u a i a go (S t l (O i ie Mō some other honeydew-producing scale insect families, e wā i ki gā gā a a u a i o Ao ea oa ka ki ea i gā such as the margarodids and eriococcids, and so are not the wā i u ua o e ao e ma a a ō gā gā a a i i oa ka main cause of the sooty moulds so abundant on beech, ā āmi i ē ā momo i ē ā momo manuka, and kanuka. ē ā a ō i ē a i a u gā a a go go ia ka u a ake e The life cycle and appearance of male and female soft o ey ew i e e o o e gā a a u a i mo e e wai eka e scales are quite different. The stages that are most often o ey ew ka a gaia i o o i e uku ma kai e gā a a i e seen are those of the female, which passes through two or ia o ē a i ākau Ka i i a u e wai eka ki e ākau ā ka three immature stages before becoming sexually mature, i u ia e ē a i momo ō u u u u a go I e i o ui ga despite still looking immature. The males, on the other o gā w e ua e ki ea ai e o ey ew kei ei a a ō ē a i hand, pass through four stages before giving rise to the momo ō oko ua ā ko e o ey ew ē a i o a a i o kai very small, fly-like, adult male. Quite unlike the adult E ga i i Ao ea oa ko a i a ake e momo u a i w aka u a female, the adult male has no functional mouthparts and o ey ew kua o o mai e ō oko ua ei oa o o mō a only lives for a few days. Its sole function is to locate and Kāo e oki e mō io ia a a e a a i ē ei ai eoi a ō fertilise the female. Some species, which live under the kāo e e ē ā awa e mā i i i i o gā u a i mo e ē ā i bark of trees, are thought to have dispensed with males and ē a i a u o gā gā a a u a i w aka u a o ey ew ( ei to reproduce asexually. Most of the native species seem to aui a gā ma ga o i me gā e iococci Wai oki e a a have only one or two generations a year. ā gā u a i mo e e ō u u u u a go ka ki ea e i u The main stage used to identify and describe a species a a i e aw ai e mā uka me e kā uka of scale insect is the adult female and this revision deals e e ekē e ā ua me e aka i ae e o gā u a me gā only with this stage. Mature females rarely reach 10 mm ā e Ko e aka i ae e o e u a e mea e ki ea ui ia a a long and many are only 3-4 mm when fully grown and, E ua e o u ā ā ei gā u i ga o ō ae ā ua i e w although many New Zealand species can be identified by kō u ga u ga a a kāa i a ō ka u i ei akeke eoi a ō the form of the glassy cover or test, it is best to check the e i e o u ō a a ga i ē ā wā ki ō e kō u ga u ga identification with a microscope by mounting the E ga i ko e mea ā e e w ā gā u i ga kā a i ka u i specimen on a glass slide. The characters that are most ei akeke me e i e o ō a ā ua akeke ki ō ē a i ga o critical for identification are the numerous minute wax- i i eoi a ō kāo e e wa a o e a e ā i o o i gā ā secreting pores found over most of the body, and these o u o u ei ka emo Ko a i a ō e kau a a a e ā e ko require a high-power microscope for examination. e kimi i e u a ka uku ā ea ki e kāka o A ā oki ē a i momo ka o o ki a o i e e a o ē a i ākau e w aka ae ia a a kua ko e kē e mea ā e; ka w aka u a u i i u ga i e ōkai-ko e Ko e ui ga o gā momo ake ake ake o Ao ea oa ko a i oa i o e ua ā ei gā wā e w aka u a u i ai i e au Contributor C is o gso graduated from Kings Col- Ko e ua ā o e gā a a u a i ka i o ia ei au u i lege, London University, in 1960 and taught in a small ei w akaa ua i ē ā i ē ā momo ko ō e u a akeke ā public school for two years. In 1962 he emigrated to what kua w āia ē ei ika ga i ko ei Me uaua ka oa ake i e 1 was then Rhodesia where he was a Research Officer for mm gā u a ā ko e ui ga kei e ā ua 3- mm kē Ko e the Research and Specialist Services, Ministry of Agricul- okoma a o gā momo o ko ei ake ake ake ka aea e ture, until 1967. It was during this period that he became au u i i u g i e ā ua a ō o ō ā ou ki i kōa aa a e ga i fascinated by scale insects and published some 16 papers ka w ai ua o u e ā a ā ae a u ei i o mā e ka u
( nt n d v rl f (h r t n n f l nd 4 on them, mainly soft whakarahi, kia tino tika ai e ma i Ko gā mea ei ā a scales, many of these i o i o māu i a koe ka ma i ki e au u i i ē a i u a i publications covering mo e ko gā ko o u a a a e mo oi i e mau mai a a ki e the whole of the Ethio- i a a eoi a ō me mā ua w akama i ē a i ka u pian Region. In 1967 w aka a i ka a o u e a ai e ki e a u he returned to England and became a Lecturer in Agricultural Ento- mology at Wye Col- I puta mai a C is o gso i e Kā e i o Ki gs W a e lege, University of Wā a ga o ā a a i e au 19 ā ka ua au e w akaako London, where he was a a i ē a i ku a ūma a ui aku ei ō e au 19 ka until August 1999. For eke ki e w e ua i ka a ga ia ko o esia i ē ā wā Ko the first 20 years at a a ū a ga i ei a ā ae awa ki e au 19 7 ko e Ā i a Wye, Chris worked a ga au mō e a o ga a ga au Kau a a W āi i ί e mainly on aphids, par- Ma a ū A uw e ua I a ia i ei a ka wa ea ia ki gā gā a a ticularly on apterous dispersal and aphid-plant-virus in- u a i ekau mā o o gā u i ga ka w aka u ai a e ia teractions on which he did his doctoral research. Chris mō g gā a a u a i ko e ui ga e ā a a ki gā u a i started working on scale insects again in 1990, since when mo e Ko e ma a o aua u i ga i i i o w ā ui ki gā he has published a further 14 papers, three of them on momo o e akiwā o E io ia ō Ν e au 19 7 ka oki ki New Zealand scale insects. Chris has written or edited three I ga a gi ū ei ūke gaka mō e Μā ai e eke i o o i books, and is the author or co-author of 6 book chapters gā Ma i A uw e ua i e Kā e i o Wye W a e Wā a ga and 59 Scientific papers. Last year (1998) he organised o ā a a I ei a o u a ia e ma i a a ā aka awa ki e the VIIIth International Symposium on Scale Insects Stud- e e- u i-kōkā 1999 Ι ō a au ua a i i Wye ko gā ies, held at Wye, and is currently editing the Proceedings. a i ā a a o ga ui ā ko gā kau a a w i i ko e w akaw i iw i i e e-ko e a gā a i me gā o o i Contributor Rosa wae ga ui i e a i e i u me e w eo i - koi ā a ō oki Henderson graduated e kau a a o a a ma i a ga au āku a a ga Ka ae awa from the University of ki e au 199 ka a u i a ō a C is ki e ā a i o i o i gā Canterbury, New Zea- gā a a u a i Mai ί ē ā wā 1 gā u i ga kua u a i a land, in 1965, and was ia e 3 e ā a a ki gā gā a a u a i o Ao ea oa E o u gā a Research Fellow in- uka uka kua u ia e ia kua o o ā ei ko ia e ē i a Ι tua vestigating chromo- a u i ē ā e gā wā a ga uka uka e 59 a ō oki gā some abnormalities in u i ga ū aiao o o kua u ia eia ko ia ā ei ē a i o gā leukaemia for 5 years kai u i Ι e au 199 a i w aka i e e W aka auika a ga at the Cytogenetics uawa uū o e Αo mō gā ā ai Μ gā a aki U a i i Unit, Christchurch Wye ā ko ia a ō e ē i a o gā kō e o o e ui e Hospital. After a 15- w aka uka uka gia a a year break from sci- ence bringing up her Νō e au 19 5 i w iw i ai a osa e e so i a a o u family, she began an mā au a ga mai i e W a e Wā a ga o Wai a a i Ao ea oa entomological career Ka ima au a aia ei aewaie ū a ga au i e Wā a ga in 1985, rearing insects for DSIR at Mt Albert Research i o i o Ā ua a ga ukuō i e a Ι o o gā ū au i e o Centre. From moths and crickets, this progressed to rear- Ō au a i ko ā a e ū u a i e wā i kiūii a ē a i ko okē ing predatory ladybird beetles and mites (both feeding on i o o i e ma e u u o o ekau mā ima au ia e w aka au scale insects) in a biological control project for kiwifruit. ka a a a ki e w aka i u ama iki ā ō e au 19 5 ka When the DSIR was broken up into ten Crown Research oki mai ki aiao e aoka īma a ū ki e w āw ā ae e i Institutes in 1992, Rosa became a science technician for gā ma i mi ai e eke Ko a a ma i ua a i e ā a Landcare Research working on scale insects in the N.Z. w aka i u i ē a i o gā ai a ga a u ga mā e SI i e Arthropod Collection. She is responsible for the curation oka ū a ga au o M A e e ū ē ua e i a ei ga of the specimens in that part of NZAC. Her interest in the gā mea ua a i ka w aka i u ia ka w ai mai ko ē a i
( nt n d v rl f (h r t n
8 d n & nd r n (2000 : C d (In t : pt r : C d soft scales had its origins with the collection of many e Ku u Āw i a - G ossa y undescribed species during the Insect Survey of the East Cape Region, 1992-1994, organised by John Dugdale. ā ua a ga uku i o o gā ū au — cy oge e ics John was leader of the Systematics team at that time, and a gawa o — a o o strongly supported the revision of the N.Z. soft scales. In here — cherry 1995, Rosa gained a Queen Elizabeth II Technicians Study huakiwi — kiwifruit Award and travelled to U.K. for a short period, to work kaihangarau — technician with Chris Hodgson at Wye College on this revision. Rosa ko okē — a o ma is author or co-author of 14 scientific papers. kuku e — e o e mohe — soft harare — wax karu whakarahi — microscope goiku a me ē a i a u mo oi i e ua e ua kai ai i e gā a a kōa aa a — g assy u a i Ko ē ei ma i w akamu u ga e kau a a i ko o u a — o e w aka ū ia ei a u ā-koio a ί gā a a w aka a u i e mā ai — s u y e ami e c ose y uakiwi ō e wāwā a ga o e SI i e au 199 kia 1 mate ruru toto — leukaemia awa gā ū a i a ga au Ka au a ka o o a osa ei mō i u i u — g ossy wa y kai a ga au a a ū aiao mā Ma aaki W e ua Ko a a ngoikura — ladybird kau a a w āi i ko e mā ai i gā gā a a u a i kei e ngongo pia — sap-sucking Ko i ga A gawa o o Ao ea oa Ko ia oki kei e āka ga ki i a i — c ose e a i e w aka ae e i gā ma i iaki i ē ā wā a ga o e Ko i ga Paewai — Fellow Ko a a gākau ui ki gā u a i mo e i akea mai i e pirinoa — parasite ko iko i ga o ē a i momo ma a o u kāo e a ō kia ā a ūi a — c omosome w akaa ua ia ί e i o a ga gā a a ki e aki o e ū ē ua — mo ai āw i i 199 -9 ί w aka ae e ia i a o i e ma u o taparau — polygon o ug a e Ko o e kai au ū o e kā ui ū a a tautuhi — identify W aka ō ū i ē ā wā ā a i akiaki kia ā a i o ia a ōkai-ko e — ase ua gā w aka a a o gā u a i mo e o Ao ea oa I e au 1995 u a i — sca e (i sec ka w akaw iw ia a osa ki ē a i o u Quee E i a e w eo i — i us ΙΙ mā e Kai a ga au i ā ei ai ia ki te haere ki Ingarangi mō ē a i wā ki e ma i a i ki a C is o gso i e Kā e i o Wye ā ko ei i u uki ai ē ei i i o ua ua ki gā u a i mo e ei ekau mā w ā gā u i ga ū aiao ko osa e Translation by aco kai u i ē a i ā ei o gā kai u i Huatau Consultants, Levin
The authors shared the work of this revision as follows: the 11 indigenous genera were divided between us, with the senior author (CJH) being responsible for about two-thirds. Draft generic diagnoses and species descriptions were written and then critically appraised by the other author. The introduction was a shared contribution. CJH was responsible for the section on adventive species, and for the sections popular summary and morphology. To all descriptions RCH added comments on biology, distribution, and host plants, collated the information in appendices A-D, and plotted the distribution maps. RCH also did most of the collecting and curation of the speci- mens, photographed the live insects, and was responsible for the scanning electron microscope studies. CJH drew all the figures, except for some of the adventive species as acknowledged. n f l nd 4
A S AC
The adult females of all Coccidae (Hemiptera: Coccoidea) known from New Zealand are described and illustrated. In addition to the four indigenous genera previously known (Ct n h t n, In l , n h t n and n , seven new genera are described (Aph n h t n, Cr t ll t t , Εp l d h t n, K l r , l h t n, r p z and U b n h t n , all of which are considered to be fairly closely related to Ct n h t n and all endemic to New Zealand. Within these eleven indigenous genera, 25 new species are described and 12 species are transferred to new genera, bringing the total to 43 species. Ct n h t n l n t Maskell is designated a n n d b . None of the species currently included in the genera Ct n h t n and In l described from outside New Zealand are here considered to be congeneric. In addition, all known adventive (exotic) species found in New Zealand are reviewed and briefly described. n (E l n p n Brittin is synonymised with rth n l n p r (Fabricius), and the synonymy of n n Maskell with S t l (Olivier) is upheld. lv n r p d (Maskell) has not been recorded in New Zealand and its inclusion in previous checklists is erroneous. Keys to adult females are included for the separation of all genera and species, and also for preliminary instar separation. Earlier work done on the superfamily Coccoidea in New Zealand is briefly reviewed and the present classification of the Coccidae and closely related families is outlined. Their biology, distribution, host-plant interactions, parasites and predators, and economic importance are also described. The current methods for collecting and mounting specimens on slides for the New Zealand Arthropod Collection are outlined, followed by a description of the morphology of adult female Coccidae, introducing the terminology used. A list of plant hosts is included in an appendix.
Keywords. Hemiptera, Sternorrhyncha, Coccoidea, Coccidae, new genera, new species, adult females, taxonomy, keys, distribution, host plants, natural enemies, fauna.
Hodgson, C. J.; Henderson, R. C. 2000. Coccidae (Insecta: Hemiptera: Coccoidea). n f l nd 4 , 264 pp.
v d: br r . A pt d: 2 Μ .
C ECK IS O A A
Family COCCIDAE Genus Cr t ll t t Henderson & Hodgson, n. gen. 79 Indigenous species [Not yet allocated to subfamily or f (Maskell, 1891), n. comb. 87 tribal levels of classification]. f (Maskell, 1884), n. comb. 89 Genus Aph n h t n Henderson & Hodgson, n. gen. 57 l pt p r (Maskell, 1882), n. comb. 90 h n hl Henderson & Hodgson, n. sp. 68 n f Henderson & Hodgson, n. sp. 92 d r Henderson & Hodgson, n. sp. 69 rn t (Maskell, 1885), n. comb. 94 r Henderson & Hodgson, n. sp. 70 rn t ll Henderson & Hodgson, n. sp. 95 n n p (Maskell, 1892), n. comb. 71 Genus Ct n h t n Maske Ι 79 97 h Henderson & Hodgson, n. sp. 72 h l n Henderson & Hodgson, n. sp. 103 t Henderson & Hodgson, n. sp 74 p r v r d Henderson & Hodgson, n. sp. 105 pr n Henderson & Hodgson, n. sp. 75 t r Henderson & Hodgson, n. sp. 107 p b n Henderson & Hodgson, n. sp. 76 v r d Maskell, 1879 108 bt l Henderson & Hodgson, n. sp. 78 [ l n t Maskell, nomen dubium] 184
1 d n & nd r n (2000 : C d (In t : pt r : C d
Genus Ep l d h t n Henderson & Hodgson, . gen.111 rn (Bouché, 1944) 206 p p r (Maskell, 1882), n. comb. 112 p r (Fabricius, 1776) 207 Genus In l Maskell, 1879 114 Genus S t Déplanche, 1859 208 p t ll Maskell, 1879 117 ff (Walker, 1852) 208 Genus K l r Henderson & Hodgson, . ge . 119 l (Olivier, 1791) 210 d pr (Maskell, 1884), . com . 24 Tribe Pulvinariini Targioni Tozzetti 212 p r d pr Henderson & Hodgson, . s . 126 Genus lv n r Targioni Tozzetti, 1867 212 p rf r t (Maskell, 1879), . com 127 fl f r (Westwood, 1870) 212 Genus n h t n Maskell, 1882 130 h dr n Steinweden, 1946 217 t t Henderson & Hodgson, . s . 132 br nth (Vallot, 1829) 218 tr d r Maskell, 1882 139 v t (Linnaeus, 1758) 219 n r Maskell, 1891 140 t ll r Henderson & Hodgson, . s . 141 CO E S Genus l h t n Henderson & Hodgson, . ge 142 I o uc io 11 d d Henderson & Hodgson, . s . 144 Classification 12 l rp (Maskell, 1885), . com . 151 Biology and life cycle 13 fl v (Maskell, 1884), . com . 153 Distribution and host-plant associations 14 n Henderson & Hodgson, . s . 155 Parasites and predators 16 156 p ll n Henderson & Hodgson, . s . Economic importance 17 p n t t Henderson & Hodgson, . s . 158 Collecting and mounting methods 18 Genus r p z Henderson & Hodgson, . ge . 159 Conventions 20 l b t Henderson & Hodgson, . s . 160 Key to instars of Coccidae in New Zealand 21 d r d (Maskell, 1892), . com . 164 Key to adult female Coccidae in New Zealand 22 Genus n Henderson & Hodgson, 1998 ... Black and white plates of SEMs (morphology) 29 Ν 166 Colour plates of female Coccidae in life 33 n t Henderson & Hodgson, 1998 X169 Morphology 49 t b l Henderson & Hodgson, 1998 170 Taxa of indigenous species in New Zealand 57 Genus U b n h t n Henderson & Hodgson. . ge ..... Taxa of adventive species in New Zealand 184 171 References 221 d l Henderson & Hodgson, . s . 173 Appendix A: Alphabetical listing of host plants includ- b ll t Henderson & Hodgson, n. s . 179 ing family, with their associated soft scale species in h n nth r (Maskell, 1885), . com . 180 New Zealand 231 j b t Henderson & Hodgson, . s . 181 Appendix B: Alphabetical listing of endemic soft scale p ll p Henderson & Hodgson, . s . 182 genera and species with their associated host plants and families 239 Adventive s ecies: Appendix C: Hymenopterous parasitoids recorded from Subfamily CEROPLASTINAE Atkinson 184 Coccidae in New Zealand 242 Genus C r pl t Gray, 1828 185 Appendix D: Geographical coordinates of collection r f r (Fabricius, 1798) Ι 5 localities with area codes 243 191 d tr t r Newstead, 1917 Distribution maps 246 [r b n 192 Maskell, 1893, ou u eco ] Taxonomic Index 257 [r (Linnaeus, 1758), ou u eco ] 192 n n Del Guercio, 1900 193 ACK OW E GEME S Subfamily COCCINAE Fallén 194 Tribe: Coccini Fallén 194 We are especially grateful to colleagues in Landcare Genus C Linnaeus, 1758 195 Research (all at Auckland unless noted at Lincoln): Jo h p r d Linnaeus, 1758 195 Berry for the identification of reared parasitoids and l n l (Douglas, 1887) 198 information about earlier records; Leonie Clunie and Tribe Saissetiini Hodgson 199 Grace Hall for their company on many enjoyable field Genus r t Takahashi, 1955 205 trips; Des Helmore who did the final layout of the n r (Nietner, 1861) 205 morphology figures; Robert Hoare for advice on the Genus rth n l n S l , 1908 206 construction of the scientific names for new genera and n f l nd 4
species; Peter Johnston and Eric McKenzie for the Technology, under research investment Contracts C09617 identification of entomogenous fungi; Birgit Rhode who and C09812. Chris Hodgson thanks Landcare Research prepared the scanning electron micrographs and paved the for a generous travel grant (also from FRST Contracts path towards digital versions of these; Dominic Thoreau C09617 and C09812) to visit New Zealand, January– for digital preparation of the maps; and Ian Payton and February 1998, to finalise this revision, and also the Royal Peter Wardle (Lincoln) for advice regarding the longevity Society, London, for a further grant towards his travel of leaves on indigenous evergreens. For valuable expenses. information on adventive species we thank the following: Chris Hodgson would especially like to thank his Karyn Beal for providing records from PPIN (source of colleagues at Wye, particularly Mike Copland, for taking data — Plant Pest Information Network (PPIN) Database, on many of the activities Chris should have been Ministry of Agriculture and Forestry, New Zealand); John responsible for whilst this work was in progress and during Charles (HortResearch, Auckland), Jill McLaren his visits to New Zealand and Australia. In addition, Chris (HortResearch, Alexandra), and David Steven (IPM offers his sincere thanks to his wife Charlotte and his Research Ltd, Auckland) for information about economic daughter Jessica for their patience, support, and importance of adventive species; Maurice O'Donnell of understanding — with the hope that the visit to New the National Plant Pest Reference Laboratory, MAF, Zealand was some small compensation! Lincoln, for providing verified records of adventive scales in New Zealand — most of these were identified by Lindsay Hawke at Lincoln, with additional records from I O UC IO Levin identified by Dave Manson. We are also grateful to the following for the loan of The earliest record of a soft scale insect from New Zealand material in their care: Jon Martin and the Keeper of is from Cook's First Voyage, 1768-71. During this Entomology for specimens from The Natural History voyage, the botanists Banks and Solander collected much Museum, London, U.K.; The Curator of Entomology, herbarium material, and parts of that collection have been Forest Research, Rotorua, N.Z.; the Collections Manager deposited in various museums, including the United States and Dr. Dug Miller of the Systematics Entomology National Museum of Natural History (USNM). At some Laboratory, USDA, Beltsville, U.S.A., for specimens stage, a single adult female soft scale, along with the piece from The National Museum of Natural History, of leaf beneath it, was cut out from a herbarium specimen Smithsonian Institution, Washington, U.S.A. and preserved dry in the USNM (accession number We thank editors and authors for kind permission to 1276401); this is labelled Ct n h t n on C pr use their figures as follows: Figs 140, 142, 144, M ll r . There is some doubt about the accuracy of the plant n bl t n 84 f th A r lt r l Exp r nt host record, as C. r has very narrow leaves, far St t n, Un v r t f M r l nd, and W.F. Gimpel; Fig. narrower than the breadth of a fully mature female 141, ll t n f th r t h M ( t r l t r Ct n h t n lt is uncertain whether the specimen is Ent l , l 6, ©Natural History Museum, Ct n h t n p r v r d n. sp. or Ct n h t n v r d London, UK. (G. De Lotto); Fig. 143, Ann l d l Maskell; the latter has been recorded from C pr S été Ent l d r n ( .S.], l 0, and whereas C. p r v r d has not, but without slide mounting G. Pellizzari; Figs 146, 149, 151, 153, hn l S r n and therefore altering a unique sample, the benefit of the A r lt r l t t nd l nt th l . , doubt concludes that it is possibly C. v r d . California Department of Food and Agriculture, and R.J. New Zealand Coccoidea were first studied between Gill; Figs 145, 147, 148, 150, 152, 154, 155, h S l 1879 and 1898 by W.M. Maskell, recognised then as one In t l C d , I nternational Institute of of the World's leading authorities on scale insects. He Entomology, CABI International (C.J. Hodgson). described about 300 species of Sternorrhyncha from The authors wish to thank the following referees for around the world, including many scale insects from New their helpful comments and suggestions for improving the Zealand. Maskell's interest in the then new science of manuscript: Penny Gullan, Australia; Doug Williams, microscopy meant that he was well ahead of his U.K.; John Dugdale and Rowan Emberson, New Zealand. contemporaries in describing species from slide-mounted Rosa Henderson thanks her husband Graeme for his specimens. Although they were unstained and had other encouragement, for his company on various field deficiencies, these slides can still be used today. Maskell adventures, and for his help with digital image processing. also kept dried material both of the specimens he collected Funds for the work by Rosa Henderson were provided and of those he was sent. However, these cannot always be from the Foundation for Research, Science and considered to be syntype material as he tended to put
1 d n & nd r n (2000 : C d (In t : pt r : C d
material collected on different days or from different None of the species currently included in Ct n h t n localities in the same box if he thought they were the same and In l described from outside New Zealand are here species (see Deitz & Tocker, 1980). Maskell was before considered to be congeneric. Thus ln l becomes his time in recognising that stages other than the adult monotypic (I. p t ll Maskell) with the four other New female might be important in the taxonomy of coccoids Zealand species originally placed in In l transferred and his dry material collection can be useful as it often to th rAph n h t n n. gen. —A. n n p (Maskell) contains nymphs and males. — or Cr t ll t t n. gen. — C. f (Maskell), C.
Since Maskell's time, there have been a number of l pt p r (Maskell), and C. rn t (Maskell). The ge- other studies on New Zealand Coccoidea. The ten families nus Ct n h t n now includes only C.v r d Maskell and of Coccoidea now known from New Zealand are as three new species — C. h l n, C. p r v r d , and C. follows: Asterolecaniidae: Maskell's species were re- t r . Ct n h t n d pr Maskell and C. p rf r t described by Morrison & Morrison (1927) and the World Maskell are transferred to K l r n. gen.; C. l rp fauna was revised by Russell (1941); Cerococcidae: 3 Maskell and C. fl v Maskell to l h t n n. gen.; C. species; 2 redescribed by Lambdin & Kosztarab (1976, d r d Maskell to r p z n. gen.; C. f to 1977) and 1 new species (Lambdin, 1998); Coccidae: Cr t ll t t n. gen.; C. h n nth r Maskell to Morrison & Morrison (1922) and Hodgson (1994a) U b n h t n n. gen.; and C. p p r Maskell to redescribed the type species of Maskell's three New Ep l d h t n n. gen. Ct n h t n l n t Maskell is Zealand genera; Hodgson & Henderson (1998) described designated a nomen dubium for reasons explained in full a new genus and 2 new species; Diaspididae: some on page 184. The genus n h t n retains its original species of the tribe Leucaspidinae by Britten (1937) and de two species . tr d r Maskell and . n r Maskell. Boer & Valentine (1977); an unpublished field study of In addition, n (E l n p n Brittin is the same taxon by Emms (1985); Eriococcidae: a synonymised with rth n l n p r (Fabricius). complete revision by Hoy (1962) and more recently a revision of the genus Er h t n by Hodgson (1994b) and The known adult males and immature stages will be de- Hodgson & Henderson (1996); Halimococcidae: a scribed in future volumes. C l b p species described by Deitz (1979); Margarodidae: completely revised by Morales (1991); earlier papers covering aspects of this family are Brittin The Classification of the Lecanoid Coccoidea. The use (1935), Dumbleton (1967), Morrison & Morrison's of cladistic analyses to determine possible relationships (1923) redescription of Maskell's species, and Morrison's within the Coccoidea have been relatively few: Miller, (1928) world monograph; Ortheziidae: comments by 1984 (many taxa of the Coccoidea, based on a range of Green (1929); Phenacoleachiidae: Beardsley (1964); and characters); Miller and Miller, 1993a, 1993b (affiliations Pseudococcidae: completely revised by Cox (1987) with of t and Er r Foldi, 1995 (affiliations of an earlier paper by Williams & de Boer (1973). Miller and Williams, 1995 (affiliations of Thus, the soft scales or Coccidae have not received the Micrococcidae); Qin and Gullan, 1995 (relationships much attention this century since Maskell described 17 within the Ceroplastinae); Hodgson and Henderson, 1996 species of indigenous soft scales in three genera (affiliations of Er h t n Miller & Hodgson, 1997 (Ct n h t n, In l , and n h t n over a period of (relationships within the lecanoid coccids), and Foldi, about 20 years, his last paper published in 1898. Exactly 1997 (many taxa, considered from the viewpoint of the 100 years later Hodgson and Henderson (1998) described evolution of their feeding sites and protective structures). a fourth genus, n , containing two new In addition, several non-cladistic phylogenies have been species. suggested, i.e.. Borchsenius, 1958 (based mainly on adult This volume provides a complete revision of the adult ema e c a ac e s ; o a yński & a ies 1971 ( ase o females currently known, adding descriptions of seven adult male characters); Koteja, 1974b (based on the new genera — Aph n h t n, Cr t ll t t , structure of the adult female mouthparts); Miller & Ep l d h t n, K l r , l h t n, r p z , and Kos a a 1979 (a e sio o o a yński & a ies (1971 U b n h t n — and 25 new species, bringing the total to modified on the basis of more recent descriptions of 11 indigenous genera and 43 species in New Zealand, males); Danzig, 1986 (using the morphology of the adult although it is clear that there are still further species female, adult male, and crawler and also life history awaiting description. In addition, 14 adventive species are characters); and Koteja (reviewers' remarks in Kosztarab illustrated and briefly described. & Κo á 19 n f l nd 4
Aclerdidae Cissococcinae Eulecaniinae i i ii ae Cyphococcinae Pseudopulvinariinae E io e i ae Myzolecaniinae Cardiococcinae Paralecaniini ig. . C a og am o uce y successi e weig i g, wi e Ac e i ae as e ou g ou , usi g 0 c a ac e s om e Ce o as i ae a u ema e, a u ma e, a s i s a ym s, c a ac e s a es u o e e ( e g 22 CI0.4 I 0. ( om Saisse i i Mi e & o gso , . Ct n h t n a ge e a c ose o i u i a ii i a e oug o e o g o e Ca iococci ae w i e n may e o g o e a a eca ii i. Cocci i
Few classifications have been suggested for the family paurametabolous type (Fig. 2, p. 21). On the other hand, Coccidae, the most recent being that of Tang t l. (1990), the development of the male resembles that of a Tang (1991) and Hodgson (1994a). The two papers by holometabolous type, producing a fully-winged adult Tang proposed a rather complex classification based stage, quite different from that of the female. The 1st- entirely on female characters but most of the generic instar stage or crawler of both males and females usually groupings were very different from those suggested by the are indistinguishable. This is the main dispersal stage studies of males (Giliomee, 1967). Hodgson (1994a) (Greathead, 1997) and is a feeding stage. The 2nd instar is introduced a classification based on the relationships also a feeding stage for both sexes but the nymphs are suggested by the morphology of both adult males and usually easily separable because they are sexually females. He divided the Coccidae into 10 subfamilies, dimorphic, those of the male secreting a more elongate with the Coccinae divided into four tribes, although he glassy test under which all further development takes considered that the status of these groupings needed place. In the male, the 2nd-instar nymph then moults three further study. This classification was the basis of a times giving rise to the non-feeding, sessile prepupal and cladistic study done by Miller & Hodgson (1997), the pupal stages and finally the winged, non-feeding adult results of which are shown in Fig. 1. The New Zealand male. These moults take place beneath the protective coccid fauna with a glassy test (those related to glassy test secreted by the 2nd-instar male nymphs. The Ct n h t n are here believed to be close to the glassy adult males of indigenous species emerge backwards from scales (Cardiococcinae); the placement of genera such as beneath their test, wingtips first, by means of the upwardly n uncertain but they appear to be close to flexing plate on the posterior end of the test. the Paralecaniini. It is interesting to note that in all the The crawler disperses to locate a suitable feeding site, cladograms of Miller & Hodgson (1997) the Cardiococcinae which may be on the same plant as the natal mother or a a a eca ii i we e sis e a a Α e ai e y oge e ic elsewhere. The test covering a neonate crawler is analysis will be undertaken once the description of all composed of soft wax that is less protective than that stages, i.e., including the adult males and immatures, has secreted once feeding has started. Crawler mortality is been completed. high as many do not find a suitable food source before succumbing to environmental factors, such as high Biology and life cycle. Male and female coccoids go rainfall, desiccation by wind or high temperatures, through very different post-embryonic development. The overcrowding on the natal leaf, and losses through wind females are considered to be neotenic and reach the non- dispersal. Surviving nymphs apparently often move away winged adult stage after two or three moults. The changes from the first settlement site and disperse further. Male that take place at each moult are relatively small and the nymphs can move to non-specific host plants once they metamorphosis is of the heterometabolous- have ceased feeding (or, perhaps, in the late feeding stages
14 d n & nd r n (2000 : C d (In t : pt r : C d before becoming completely sessile), settle and secrete the nymphs, there are also two groups of tubular ducts on the wax which fixes the test firmly to the substrate, and then dorsum on about the 4th abdominal segment. These ducts begin metamorphosis. This ability to settle on other plants secrete a type of wax that appears to act as the hinge for the makes determining the relationships of an adult male to a posterior test plate, so that it can be raised during the particular female difficult; thus, plant species on which emergence of the adult males. The 2nd-instar males moult only male stages are known are not here considered to be into an elongate prepupal stage that shows the first signs of true host plants, as the female stages appear to have a more wing-buds, dorsal and ventral eyes, and a genial sheath; restricted host range. they lack functional mouthparts. The pupa is similar but In the female, subsequent nymphal instars change the legs and wing-buds are better developed and the genial little other than in size, and the only safe way to be sure that sheath is longer. The adult male remains beneath the male you are looking at an adult is to locate the pregenital disc- test for several days until fully developed and the long wax pores or the vulva (more difficult). Usually on adult caudal filaments have been secreted (not present in all females, other pores such as the dorsal macropores are species), when it emerges backwards, and then actively obvious and the ventral tubular ducts, when present, are flies looking for females. As it has no functional more numerous than on younger instars. When most mouthparts, its life is rather short, usually only a few days. female stages are available, the difference in size of the It is likely that some species of soft scale lack males anal plates and/or the clypeolabral shield makes instar completely and are therefore parthenogenetic; for identification reasonably easy (see key to stages, p. 21). example, those of Poropeza, where the adult females are The female life cycle may have either two or three hidden beneath the bark of the host trees. moults after the 1st-instar. Most indigenous species have The adult females of all soft scales have functional three nymphal stages before the last moult; the third mouthparts and feed on sap from the phloem. Before the nymphal stage is sometimes very brief and so it is possible development of their ovaries, young adult females are that a third instar is more widespread than records suggest. only about the size of the last nymphal stage (Fig. 121 With few exceptions, New Zealand native plants are under Lecanochiton). They then undergo a very consid- predominantly evergreen (e.g., Wardle 1991, p. 39) and erable increase in size, generally at least two to three times there are few records of coccids from the deciduous native their post-moult size, although this may be as much as ten trees. Amongst the evergreens recorded as hosts for scale times for species of Ctenochiton and Poropeza. Once this insects in New Zealand are Hedycarya arborea, Griselinia swelling period is complete, the females are ready to spp., Mida salicifolia, Myrsine australis, Pseudopanax reproduce. In many species (such as those in the genus spp., and Schefflera digitata. The leaves of these lowland Lecanochiton), the venter develops a distinct concavity to broadleaf trees may not senesce for several years, although form a brood chamber. In species of Lecanochiton, the individual leaves may be shed at any time of the year. vulva is towards the middle of the abdomen, so that the Thus, a scale insect species with an annual life cycle can eggs can be deposited directly into the chamber. In some reliably overwinter on the previous summer's (old) leaves. of the other New Zealand species, such as those in the In some univoltine species, e.g., Ctenochiton paraviridis genera Kalasiris and Pounamococcus, the posterior half and C. viridis, it is the 2nd-instar nymphs that overwinter; of the glassy test is used as the brood chamber, the adult in this case, the female nymphs migrate in the spring from female shrinking into the anterior half of the test as she the old leaves and resettle on the new, fast growing shoots, oviposits; in these species the vulva is close to the where the rising sap supplies nutrients for rapid growth to anogenital fold at the posterior end of the abdomen. Quite maturity. Other univoltine species overwinter as the a lot of species appear to contain fully developed nymphs immature adult female on the leaves (e.g., Aphenochiton and, therefore, are either viviparous or ovoviviparous (see subtilis) or on the stems (e.g., Plumichiton pollicinus), Tremblay, 1997). whilst others, such as Aphenochiton pubens, are mature In the adventive Pulvinaria species, the ventral and reproduce on leaves from late winter. Sessile male tubular ducts, of which there are three or four types, nymphs remain on the old leaves and adult males emerge secrete a long white ovisac from beneath the venter (Figs from there. C84, C85, C87). The eggs are then laid within this. In the male, the crawler and the 2nd-instar nymphs are the only feeding stages. The test is usually glued to the is i u io a os a associa io s. Fourteen plant surface, apparently by waxes secreted from a cosmopolitan species of soft scale insects are established submarginal band of tubular ducts, the presence of which in New Zealand. Most of these adventive soft scales can be can separate most 2nd-instar males from 2nd-instar found in citrus orchards as well as on other host plants. The females. In all known male New Zealand 2nd-instar species found on citrus include the three Ceroplastes n f l nd 4
s ecies i ew ea a —C. r f r ( a icius , C. e e a ea s o e o is i c a e o e a io s i s d tr t r ews ea , a C. n n e Gue cio— amo g e e emic so sca e s ecies a ei associa e w ic a e es ic e o e wa me a eas, ge e a y om os a s ecies. e eg ee o o y agy a ges om o a o Gis o e, a wo s ecies f C , wo e ig f Ct n h t n p r v r d .s ., Ep l d h t n s ecies f S t , a o e r t p . e p p r (Maske , a l h t n fl v (Maske wi s ecies o eco e om ci us a e rth n l n , 6, a os a s ecies es ec i e y (i 2 4 rn ( ouc é a e ou lv n r s ecies eco e a ge e a a a ami ies o ows o 2 o i ew ea a : . fl f r (Wes woo , . h dr n os a s ecies (e.g., Aph n h t n pr n .s . a S ei we e , . br nth ( a o , a . v t l h t n d d .s . . A o e o y agous ( i aeus . s ecies is K l r p rf r t (Maske wi 8 os a I ew ea a , e mos commo a wi es ea o s ecies i c u i g 6 C pr a 6 tt p r s ecies. e a e i e so sca es is C h p r d i aeus O e e emic cocci s ecies a a e e wee 4 a i as ee ou o e Ke ma ec Is a s o e o a 8 associa e s ecies o a os s, (i 4 o 4 a C a am Is a s o e sou eas , as we as oug ou ge e a , r p z d r d (Maske is e o y o e mos o e o a Sou Is a s. C. h p r d is es ic e o a si g e a ami y, e o oca aceae (see mai y a es o o ame a s, o ou oo s a i oo s, A e i . a i a so occu s i ea y a y es o ui o c a s— e um e o e emic so sca e s ecies eco e eco s a e om a o a o e o ic a s ecies a om om a icu a os a s ecies is equa y i e se, a 22 s ecies o a i e a s. o o e S t s ecies i i is d r rb r is y a e mos a ou e ew ea a a e a so commo a wi es ea S. l o e os a s, wi e emic cocci s ecies i (O i ie is eco e om 2 e o ic a 2 a i e a ge e a (a o a e i e cocci s ecies eco e om i . s ecies w i e S. ff (Wa ke is eco e om 8 C pr s ., tt p r s ., a d rp t t r e o ic a 4 a i e a s ecies. a e es ec i e y , , a 6 cocci e emics eco e A oge e 6 s ecies o a e i e so sca es a e ee om em (see A e i A . ou o i ige ous a i e a s as we as e o ic a s, Amo gs some e emic so sca e ge e a a a e a oug o y wo, C. h p r d a S. ff , a e ecome associa e wi c ose y e a e o si g e a ee eco e om wi i a i e o es s e o e ou ge e a, e mos o ious a e n h t n, es ic e o s ecies, C r pl t n n , C l n l ( oug as , e a ge us M tr d r , a e a u ema es o S t l , a rth n l n rn we e ei e r p z , es ic e o c ose y e a e ge e a i eco e om ga e a i es, a o es ma gi s a a o oca aceae. A g ou o ee s ecies Cr t ll t t ee is u e y uma i ue ce, o o e a u a l pt p r (Maske , C. rn t ll .s ., nd l h t n ecosys ems, e.g., o ma g o es a acke . A se e p ll n .s . a e i e ec mo o agous o e c ose y s ecies, lv n r br nth , is ou o ice e a e K nz r d a pt p r p r . a s a may e co si e e a i e u o i ige ous. O e mo o age e am es o w ic e e a e ai y e o e s ecies a e o ee eco e om a i e e e si e eco s a e Cr t ll t t f (Maske a a s a a e ge e a y o i mi o sig i ica ce o e o ic Cr t ll t t n f .s . eco e om th f a s. Aph n h t n h .s . om W n nn l h t n I ega o is i u io , a e emic s ecies cu e y n .s . fr h p l t l p d , nd n k ow om ew ea a e o g o ge e a a a e e e n t nd r n & d n fr f rn ( l hn co si e e o e e i e y e emic. E e e ge e a fraseri i a icu a . Ct n h t n a In l (w ic co ai s ecies es Some is i u io a e s o e emic so sca es ca c i e om e sew e e a e co si e e e e o e e emic, pp r d j n t, . ., Aph n h t n bt l n. p th so a e o ew ea a s ecies wi ee o e oca i ies eco e om e ee Ki gs Is a s a e eassig e o o e ge e a. is a ies e e o e o e a o e o Is a , e io a a Aus a ia s ecies, w ic mig a e ee e ec e o e Sou a (ma , a Kalasiris depressa (Maske a so c ose y e a e . is s o g e emism is a so em asise wi wi e y se a a e eco e oca i ies om o a y e os a s o e ew ea a so sca e au a, as o io a (ma 22 . e seemi g y a a a a u e o a mos o e emic cocci s a e ee eco e om e a associa io s wi i sec s i ew ea a was o e ma y i o uce a s ecies (e e i e case o e o e y ug a e ( , a may a e e o e o e e as e ce io , e co ec o o e a i was c ose o i s o ma mi io s o yea s i es o se o e some imes io e os . As some o e e emic s ecies a ea o e ai y e e sa s i o e c ima e a e s a e a e e o o y agous, is is su isi g. e a mass si ce e C e aceous. I ea i y, ma as
1 d n & nd r n (2000 : C d (In t : pt r : C d probably induced many of the relic populations of scale (C ll n r was recorded feeding on the large insects over the last approximate 150 years, with the huge Ct n h t n species known as `sixpenny scales' loss in indigenous forest cover when much of New (Leathwick t l., 1983). Hudson (1891) noted an instance Zealand was cleared for farming by European settlers (see of control of C h p r d (as n h p d Miller, 1925, for maps showing total area of indigenous by a species of h z b (Coccinellidae) and Koebele forest in 1840 and in 1924). Thus, the survival of remnant (1893) mentioned the coccinellid beetle r n forest patches may have a greater bearing on the modern nt p d (White, 1846) as a predator of Ct n h t n distribution of indigenous soft scales than past geologic or v r d . Predatory mites and Cecidomyiidae have not been tectonic events. recorded feeding on indigenous soft scale species, Some of the plant-host genera on which the endemic however, the following have been recorded attacking the soft scales are known have a much wider geographic introduced C r pl t d tr t r: the mite r ph distribution. For instance, r ph ll is also known p rn , which is thought to prey on the eggs (Lo, from Australia and New Caledonia; pt p r is 1995); the steel-blue ladybird ( l (=Or known from Australia, Malaya, and New Caledonia; h l b which is considered to be a useful biocontrol th f is known from Australia, New Caledonia, agent in citrus orchards in Northland (Lo t l., 1992; Lo New Guinea, and temperate South America, and yet the & Blank, 1992) and a cecidomyiid belonging to the genus New Zealand coccid genera, as understood in this r p (Crosby, 1986). Indigenous species are revision, have not been found in these countries. There are commonly attacked by entomogenous fungi; indeed, a few records in the literature of indigenous species Hosking & Kershaw (1985, p. 208) noted that "the elsewhere in the world but these are here considered to be collapse of the epidemic [of Cr t ll t t f ] appeared misidentifications (see under Ct n h t n l n t , to be caused by an entomogenous fungus, probably K l r p rf r t , and l h t n fl v . p r ll d pl x". Three species of parasitic fungi are Like many other New Zealand Sternorrhyncha currently known (see list below) but there are probably (Morales, 1991), the adult females of both species of more species still to be identified that attack coccids in r p z are cryptozoic, living hidden beneath the bark of New Zealand. The list of hymenopterous parasites so far their host trees, and may have an unusual life cycle identified (see Appendix C; Valentine & Walker, 1991) is associated with this habit, with the immature stages also probably very incomplete and again many more feeding on other plants (see nd r r p z nd probably species remain to be discovered. Most instances of reproducing parthenogenetically. Other species, despite parasitism are observed by the round exit hole in tests, and living on the leaf, are extremely well camouflaged. Thus, thus parasitoids are seldom reared through for most Aph n h t n species are very flat and are either a identification. The number of parasitoids emerging from a very similar green to that of the leaf or are more or less scale appears to be partly related to the size of the scale, transparent — the light reflected off the glassy test being with many larvae contained in large-bodied soft scales the main clue to their presence (Figs C32–C36). such as Ct n h t n p r v r d (Fig. C56) and with only Most leaf-living species are found on the lower leaf l-2 from small scales, such as male nymphs. As a defence surface, but n h t n t ll r , l h t n against parasitism, coccids are often able to encapsulate d d , . p ll n , . p n t t , and n the invading parasitoid eggs, thus preventing their t b l all settle on the upper surface. With regard to development (see Blumberg, 1997); in mounted n t b l , we have postulated (Hodgson & specimens, examples of encapsulation are frequently seen Henderson, 1998) that this is a mechanism for remaining as brown, stalked oval bodies, usually near the margin. clean of sooty mould and debris in an area of very high rainfall. In these areas, the lower leaf surfaces of the host Entomogenous fungi. 1. "Brown blobs": A r t plants of . t b l acquire a thick burden of other bb r H.S. Fawc. (Fig. C69): particularly infesting invertebrates, lichens, fungi, and detritus that could Ct n h t n p r v r d , probably during the early smother this insect, while the upper surfaces are nymphal stages; when completely infested, there is no maintained clean and shiny by the almost daily washing, remaining evidence of the scale insect (see Myers (1928)). which also removes the honeydew. The former four 2. "Orange puffs": p r ll d pl x (Berk.) Petch (Fig. species inhabit less extreme climates and are either heavily C70): bright orange. Commonly infesting Aph n h t n sclerotised ( . t ll r , or have very thick tests (the h , l h t n fl v , and . n . In early infest- l h t n species). ations, the scale insect's shape and patterns are visible; in older infestations, the insects become puffed up, either at Parasites and predators. Birds have been observed one end or all over, with a softer, yellow centre on top, feeding on soft scales in the native forest, e.g., the kokako sometimes exuding a ribbon-like part. n f l nd 4
3. "White fungus": rt ll l n (Zimm.) Viégas onto these plants. For the same reasons, it would appear (Fig. C71): commonly seen on adult female coccids, that they do not have great potential as pests outside New particularly C. p r v r d , where body outlines remain Zealand, other than on closely related host species or visible. Note: rt ll has a looser texture than the genera. other two fungal species — the mycelium strands have However, a number of other coccids were known from whorls of small branches with spores on their tips, which New Zealand from about the time of Maskell. As these are shine under the light of a binocular microscope. all geographically widespread, cosmopolitan species, it is assumed that all are adventive (i.e., have been accidentally a asi oi s. All the adventive coccid species are introduced from outside), and it is likely that most of them parasitised by non-native hymenopterous parasitoids, and came with plants introduced by the early European only one of these parasitoid species has also been recorded settlers. Fourteen species have been definitely recorded from an indigenous soft scale, namely E x nth ll plus two others that were apparently misidentified and, as ph l pp (Hymenoptera: Aphelinidae) from K l r there are no voucher specimens available, their possible p rf r t ( Ct n h t n p rf r t . Otherwise the earlier presence cannot be confirmed. The species that are parasitoids recorded from indigenous species are all currently of some importance are: C r pl t d tr t r endemic hymenopterous species. A full list is given in and, to a lesser extent C. n n , both of which are serious Appendix C. pests of citrus in both of the major citrus orchard areas of See Hill (1989) and Morales (1989) for discussion of Kerikeri (Northland) and Gisborne (D. Steven pers. comm.); parasitoids introduced to New Zealand for biological C. n n also is a pest on feijoa ( j ll n . control of C h p r d and S t l . C h p r d is a serious pest on ornamental plants, shrubs and ferns, both those grown indoors and outdoors, and can cause death when the infestations are heavy (D. ECO OMIC IM O A CE Steven, pers. comm.); C. h p r d also is found in low density populations on a wide range of introduced It seems that none of the indigenous soft scales are perennials, shrubs, and trees including n r d t and economically important. They produce much less r n pp., and has been recorded on about 20 honeydew than the Margarodidae and are, therefore, less indigenous plant species as well as on subtropical crops important both in terms of providing honeydew as a food such as citrus, avocado ( r r n , and tamarillo source (that from Margarodidae supports not only native (C ph ndr b t . rth n l n rn occurs birds, geckos, and insects but also honey bees and the at low levels in Central Otago stonefruit orchards, recently adventive European wasps) and as a substrate for especially unsprayed orchards, and in particular on sooty moulds. Cr t ll t t [as In l ] f was blamed apricots ( r n r n and plums ( r n spp.) (G. for the death of red beech trees ( th f f in an McLaren, pers. comm.). lv n r fl f r has been outbreak in the Maruia Valley, South Island, between recorded on an experimental tea (C ll n n r p 1976 and 1978 but there is little direct evidence, and in Nelson, as well as on ornamental camellias. lv n r Hosking & Kershaw (1985) considered that the trees had v t occurs in sporadic outbreaks on apricots in Central been stressed by several years of drought and that this Otago orchards, records being clustered around 1951-52, could have accounted for the higher scale populations. 1984-85, and (at present) in 1998-1999. lv n r v t Hoy (1958) studied the scale insects on rata has also been recorded on grapevines ( t v n f r and, (M tr d r and kamahi (W n nn in an attempt to although not a serious pest by itself, has been shown to find the causes of reported large areas of die-back in these transmit viruses associated with grapevine leafroll forest trees. As well as species belonging to other scale closterovirus disease (Belli t l., 1994). Other coccids insect families, he found n h t n tr d r and . recorded in low numbers on grapevines are C n r on rata, and l h t n Ct n h t n] fl v on h p r d , rth n l n rn , . p r , kamahi; he concluded that there was no indication that S t ff , and S. l . S t ff a minor these coccids occurred in high enough numbers to have an pest of indoor plants and outdoor ornamental shrubs in the adverse effect. It is relatively unlikely that any of the warmer north (e.g., Auckland) but has also been found in indigenous species will become an economic pest as they remnant native forest on indigenous plants including appear to be well adapted to their hosts, on which it is ferns; S. l is a pest of citrus in the Gisborne orchard likely that they evolved. As they do not appear readily to area (D. Steven, pers. comm.); both S t species have accept any of the introduced plants, including all current been found on indigenous trees in native bush margins commercial crops, it is also unlikely that they will move near human habitation. Soft scales are seldom found on
1 d n & nd r n (2000 : C d (In t : pt r : C d kiwifruit (A t n d d l , although C r pl t 1 Remove one or more specimens from the ethanol d tr t r, C h p r d , and S t l , each preservative and place in a small flat dish. Under the a primary association, are recorded in the Plant Pest binocular microscope, make a small cut or slit in the side of Information Network ( I Database. the abdomen, avoiding damage to the legs. [Note: if separating the body into dorsum and venter for any reason, this is best done at this stage.] CO EC I G A MOU I G ME O S Transfer the specimens to a heat resistant glass tube (small test tube) which has about 1 cm depth of 95% While many species of soft scale insects cannot be ethanol; put a bung of cotton wool in the top and place the identified by looking at either their general live tube in a water bath (at about 90-95°C) and heat for 2 appearance, their host plant, or their infestation site, this is minutes. not quite so true of most of the indigenous New Zealand 3 Remove from heat and carefully tip the contents of the coccids and it is hoped that the accompanying colour tube into a clean small dish; transfer the specimens to plates will help identify many of these species. However, another small test tube with about l cm depth of 10% the identity of even these and the majority of other species KOH. Return to the water bath and heat for about 3-7 is best determined or checked by microscopic study of minutes, or until the specimens appear to be mostly cleared carefully processed, slide-mounted specimens. or until the body contents appear liquid (avoid boiling or overclearing at this stage). i i g a co ec io Soft scales may occur on any part Remove from heat, allow to cool slightly (to avoid the of their host plants, from the roots to the fruit, but most hot fumes of KOH affecting one's eyes when working at often they are found on stems and the undersides of leaves. the microscope) and again tip contents into a small dish, In other parts of the world, a collector may increase his such as a 5 cm Petri-dish, to allow easy manipulation with chances of success by looking for intense ant activity, bent pins or other flattened tools; gently pump and tease honeydew droplets, and/or sooty mould. In New Zealand, out the remaining body contents, including any eggs and/ the only known association between an ant species and a or unborn nymphs (sometimes it can be useful to have one coccid is that of the ant r l dv n with r p z specimen with unborn nymphs remaining inside, for d r d when the latter is under the bark of podocarp confirmation of ovoviviparity or viviparity). [Note (i): the trees. While sooty mould can be an excellent indicator, above maceration method is extremely difficult for very again this is often less helpful than in other parts of the small specimens such as crawlers as they are easily lost world due to the presence of large numbers of other after becoming transparent with heating in KOH. This honeydew producers, particularly margarodids and method can still be used if there are many specimens as eriococcids. some will undoubtedly be retained but, alternatively, it Once found, the specimens need to be collected and may be best to macerate them in 10% KO at room stored either in 70-95% ethanol or as dried specimens. temperature for about 24 h instead. Note (ii): heavily However, the latter method requires continuous dry sclerotised specimens may need extended maceration storage thereafter, with careful protection from such time; the limits for hot KOH may be <15 minutes, `museum pests' as dermestid beetles. All stages of the alternatively leave in KO at room temperature for scale insect should be collected if possible, including the McKenzie's stain in Essig's Solution may be re-used 1 Add a printed label to each slide with full collection many times (so long as you ensure that you remove all details. At NZAC, we use typed labels printed on a laser specimens each time!)] printer. The labels are cured on the sheet after printing, to Transfer specimens to a clean staining well with 75% anneal the flakes of ink, by heating in a small oven at 150- ethanol, leave for about 5 minutes. If the specimen(s) are 160°C for 1 minute. large and very full of stain, repeat this step to wash out excess stain. eci es 7 Transfer as above to 95% ethanol, leave for about 5 1 Acetic-acid-alcohol: minutes (= dehydration). ethanol, 95% — 50 ml Transfer as above to a de-waxing mixture, made up fresh distilled water— 45 ml thus: mix (1:1) about 0.5 ml xylene and 0.5 ml absolute glacial acetic acid — 20 ml isopropanol (2-Propanol) (or, if isopropanol unavailable, 95-100% ethanol will do instead), leave for 2-10 minutes McKenzie's Stain: to clear any remaining wax; check under the microscope equal parts of acid fuchsin, erythrosin, and lignin pink, that all wax is gone and, if not, repeat this step, or gently each made up into a 2% aqueous solution. knock off any waxy pieces adhering to the outer body with a pin. 3 Essig's Solution (Essig's Aphid Fluid): 9 Transfer as above to absolute isopropanol or absolute lactic acid (reagent grade 85%) — 20 parts ethanol; leave for 10 minutes minimum (= washing and phenol (liquified) — 2 parts final dehydration). Repeat this step if specimens large. glacial acetic acid — 4 parts 1 Transfer as above to 1-3 drops of clove oil; leave for water, distilled — 1 part from 2 minutes up to 2 days (longer than this, the Heat at 56-60°C for 30-60 minutes. Store in a brown specimens may become brittle). bottle. The suggested stock staining solution is made up of Slide mounting: take a clean glass slide and put a small 11 30 ml Essig's Solution with 60 drops of Mckenzie's stain drop of clove oil in the centre; transfer a specimen from the added; this has a shelf life of several years under normal staining well of clove oil to the drop of oil on the slide; storage conditions. under the microscope arrange the specimen so that it is spread out straight; then, with a small piece of paper tissue, Im o a The slide-numbering system for Coccoidea very carefully blot away the excess clove oil from the operating in NZAC before 1992 used the Julian calendar specimen, gently touching the sides only. to select the (accession) number, which represented the Add a very small drop of Canada balsam beside the day on which the slide(s) were made, e.g.,10 April 1990 specimen on the slide, just enough to cover the specimen would be the 100th day of the year. If two or more with a thin film and then, with a pin, gently move the unrelated samples were mounted that day, each sample balsam over the specimen; this sticks it in place on the was given a unique lower case letter, e.g., three slides of a slide; it is very useful to do this when mounting a series of mealybug sample might be all numbered #90-100a, and crawlers or nymphs in a row on one slide, as they should four slides of another sample from a different locality not move from their position after the coverslip is applied mounted the same day would be all #90-100b, and so on. (see next step); write a code label or number on the slide A sample is defined as a unique collection of material in and leave to set for a few minutes. terms of species, host plant, locality, date, and collector. 1 Add a large drop of Canada balsam by gently dropping Thus, the number on the slide referred to the date on which on top of the specimen; with forceps, carefully place a the slides were made, and the lower case letter referred to clean cover slip on it, by lowering at one edge first; there the sample or collection data. A difficulty with this system should be enough Canada balsam to infill under the compared with the system used now is highlighted by two coverslip without unduly flattening the specimen, so that slides of Poropeza cologabata n. sp. numbered #83-326c, the dorsal and ventral surfaces are clearly separated; a but having no collection data, whereas a slide f r p z generous amount of Canada balsam at this stage is dacrydii (Maskell) is numbered #83-326f and has particularly important for mounting adult males. collection data from Fiordland. The assumption of shared 13 Store the slide flat in a drying oven at about 40°C for 2 collection data cannot be made about these slides, or other weeks, or at room temperature for 6 weeks. The slide must slides with the same accession number but with different be fully dried before storing on its side, but it may be taken lower case letters, if they were mounted at NZAC before out of the oven earlier if stored flat. 1992. 20 d n & nd r n (2000 : C d (In t : pt r : C d However, since 1992 the (accession) numbering imm. — immature system has been changed and slides with the same km — kilometre(s) accession number, but with different lower case letters, are L — Lake all from the same sample. The numbering now relates to lvs — leaves the order in which samples are slide-mounted: the m - metre(s) assigned number consists of a prefix with the year's last Mt — Mount, Mountain two digits (as before), followed by the sample number— no./nos — number(s) simply in a running order starting with 001 at the nr — near beginning of any year, the next sample handled would be NZFS — New Zealand Forest Service 002 and so on, regardless of the actual date collected. Pt—Point Thus, the first sample for 1998 was #98-001a–c (three R — River slides were made) and a later sample was #98-021 (the Ra — Range twenty-first coccoid sample for 1998 and only one slide Rd—Road was made). The change was necessary to allow for Res — Reserve databasing of records, with different instars or adult male/ SF — State Forest females mounted on separate slides each requiring a Stm — Stream unique code number, and also because much more Stn — Station material has been mounted since 1992. Tk, tk — Track, track V —Valley CO E IO S Co ec o s. RCH = R.C. Henderson; W.M.M. = W.M. e osi o ies Maskell (all other collectors not abbreviated). BMNH The Natural History Museum, London, U.K. CMNZ Canterbury Museum, Christchurch, N.Z. I us a io s. Text Fig. 1 is a cladogram from a cladistic FRNZ Forest Research, Rotorua, N.Z. study of coccid families. Fig. 2 shows the life stages of a NZAC New Zealand Arthropod Collection, Auckland, typical indigenous coccid. Figs 3-7 are labelled for N.Z.; [note: NZAC should be assumed where convenience, with Fig. 3 showing the structures most no depository is mentioned]. commonly noted on the indigenous species, whereas Fig. USNM United States National Museum of Natural 4 also includes structures found only on some adventive History, Smithsonian Institution, Washington species. Figs 5-7 show each of the main structures in more D.C., U.S.A. detail. The size of many structures are given in the text; details of how they were measured are shown in Fig. 7. Ma e ia s u ie . The data given under each species are Figs M8–M31 are Scanning Electron Micrographs those on the slide label, categorised under the area codes of showing detail of various structures. Figs C32–C95 are 64 Crosby et al. (1998), and arranged approximately from colour photographs of females of most of the species north to south in the New Zealand subregion. After the described or discussed. Figs 96-155 are presented in the collection data, the depository is given if different from usual way for Coccoidea, with a central 'map' of the NZAC, followed by the number of slides and specimens insect, the left half representing the dorsum and the right studied, including their life stages, thus: "6/3 [ff] ad, 2 the venter, with the approximate number and position of 3rd, 1 [m]2nd" indicates six slides with 3 adult females, two most structures indicated by symbols in the body of the female 3rd-instars, and one male 2nd-instar. map. This central map is surrounded by vignettes showing some of the important features, enlarged as seen under phase-contrast. Phase-contrast emphasises differences in A e ia io s use i co ec io a a the degree of sclerotisation of a given structure and the asl — above sea level appearance of the latter may be different using normal cnr — corner direct light (and even more so under the electron DSIR — Department of Scientific and Industrial Research microscope!). FP — Forest Park Geographic distribution maps for each indigenous FRI — Forest Research Institute species were prepared by plotting the latitude and I — Island; Is — Islands longitude point for each collection site. n f l nd 4 2 Key o e imi a y se a a io o g ow s age o i s a i ew ea a s ecies ema e Ma e Wings or wingbuds present; anal plates and mouthparts absent 2 c aw e —Wings or wingbuds absent; anal plates and mouthparts s i s a clearly present 4 2 Wings fully developed; body clearly divided into head, thorax, and abdomen; head with distinct eyes; antennae 10-segmented, bead-like and setose adult male —Wings present as wingbuds; demarcation between head, thorax, and abdomen indistinct; eyes absent; antennae with indistinct segmentation and without setae 3 Wingbuds barely extending past metacoxae; length of leg segments subequal; prothoracic legs shorter than length of head; genial sheath lobe shorter than or about same length as lateral anal lobes prepupa —Wingbuds clearly extending past metacoxae; coxae and trochanter significantly shorter than femur, tibia, and tarsus; prothoracic legs longer than length of head; genial sheath lobe clearly longer than lateral anal lobes pupa 4 Anal plates usually with one pair of apical setae longer than length of anal plates (all apical setae short in 1st- instar nymphs of n h t n t t and some Aph n h t n spp.); claw and tarsal digitules all fine, with bases of tarsal digitules clearly offset 1st-instar nymph —Apical setae on anal plates much less than half length of ig. 2. iag am s owi g e i e s ages o Cr t ll t t l pt p r (Maske . e e co um s ows e ema e anal plates; one or more claw and/or tarsal digitules s ages a e ig co um e ma e s ages. is s ecies often broad; bases of tarsal digitules either not offset or as ee ym a s ages io o e a u ema e. only very slightly so 5 Ventral multilocular disc-pores present in anogenital [*Note: there appears to be no character for consistently area; dorsal macropores or preopercular pores present; separating the 2nd- and 3rd- instar female nymphs. One useful claw digitules generally both broad adult female attribute is the proportional increase in size with growth. Thus, at least with indigenous species, if more than one stage is available —Ventral multilocular disc-pores almost never present (preferably including the 2nd- instar male and adult female), then posteriorly on abdomen; dorsal macropores or the approximate proportional increase in the length of the anal preopercular pores absent; claw digitules never both plates of the different instars can be summarised as follows: lst broad 6 instar = 1; 2nd instar = 1.5; 3rd instar = 2 to 3, and adult female = 4 to 5. If the eyepiece graticule being used to measure the 6 Dorsal tubular ducts present in at least two groups specimen is approximately calibrated to these ratios, then a quick medially on about 4th abdominal segment, and/or also measure is sufficient to identify the instar of the specimen in most in a reticulate pattern and/or marginally on venter, cases, i.e. if the anal plates measure "2.5" then it is almost certainly a 3rd-instar female (as no males have anal plates that particularly on head and thorax. 2nd- instar male long). In addition, the 3rd- instar female tends to have more —Dorsal tubular ducts absent; ventral tubular ducts, if marginal spines and more abundant dorsal pores than the 2nd- present, in a different arrangement instar female and the 3rd instar of a few species have multilocular 2nd- or 3rd-instar females* disc-pores near the anal area]. 22 d n & nd r n (2000 : C d (In t : pt r : C d SO SCA ES O EW EA A spinose setae along each inner margin; ventral tubular ducts forming a distinct submarginal band one duct Key o a u ema es wide (p. 114) ... In l —Characters not in this combination 5 Adult females either without an apparent test or secreting a glassy wax test; dorsal setae never present; with Ventral tubular ducts absent (including Ct n h t n either no stigmatic spines or only one in each stigmatic v r d in part) 6 area (except several present on n —Ventral tubular ducts present (including dorsal tubular ducts either absent (or in two diverging Ct n h t n v r d in part) 8 lines just anterior to anal plates on P. n t dorsal tubercles never present; tibio-tarsal articulatory Ge e a acki g e a u u a uc s ( sclerosis never present Group 1: Indigenous genera 6 Body more or less pyriform, widest about 2nd or 3rd —Adult females secreting a variety of tests: thick soft abdominal segment; mouthparts generally displaced wax, woolly wax or just a thin colourless wax to one side; ventral microducts most abundant covering, but not a glassy test; dorsal setae present, medially on thorax and abdomen; pregenital disc- usually fairly frequent; with three or more stigmatic pores with mainly 8-10 loculi, distributed across all spines per stigmatic area; tubular ducts occasionally abdominal segments (p. 108) ... present on dorsum; dorsal tubercles often present; Ct n h t n v r d (in part - cal/2 have 1-2 ventral tibio-tarsal articulatory scleroses usually present tubular ducts) Group 2: Adventive (or exotic) genera —Body more or less oval, widest about metathorax; mouthparts rarely displaced to one side; other characters not in above combination 7 Each reticulation area on dorsum with a sclerotisation, G OU . I IGE OUS GE E A small on young adults but complete on mature 2 Marginal setae absent; true dorsum of mature adult specimens; with a distinct anal sclerotisation on significantly smaller than dorsal surface of insect, dorsum around anal plates; stigmatic spines, when lateral margins composed of an expanded venter; legs present, only slightly larger than marginal spines absent; antennae much reduced (p. 111) ... Ep l d h t n (p. 130) ... n h t n —Each reticulation area on dorsum without a —Marginal setae present; dorsum always forming sclerotisation; anal sclerotisation on dorsum around complete dorsal surface of insect (if dorsum slightly anal plates absent; stigmatic spines at least 1.5 x and smaller than venter, then ventral surface lying in a leaf generally 2-3 x length of marginal spines depression (Ct n h t n spp.)); legs present, usually (p. 57) ... Aph n h t n well developed; antennae well developed 3 Each stigmatic cleft with more than one stigmatic spine; Ge e a i w ic u u a uc s a e ese e a y ( dorsal pores not forming a reticulate pattern; ventral 8 With a distinct band of ventral multilocular disc-pores tubular ducts restricted to medial and mediolateral around margin of body and along anal cleft margins; areas of abdomen (p. 166) ... n distinct anal sclerotisation present on dorsum around —Each stigmatic area either lacking stigmatic spines, or anal plates (p. 159) ... r p z with only one spine, or with stigmatic spines —Multilocular disc-pores never forming a complete undifferentiated from margin spinose setae; dorsal marginal band; anal sclerotisation absent 9 pores forming a distinct reticulate pattern (except on In l p t ll and r p z l b t ventral Pregenital disc-pores abundant and forming distinct tubular ducts, when present, not restricted to medial bands medially across most or all abdominal segments and mediolateral areas of abdomen 4 10 4 Marginal setae spinose and of two distinctly different — Pregenital disc-pores not abundant, either restricted to shapes, one type sharply spinose and other clubbed, groups on either side of ano-genital area or forming a the two types often alternating; posterior band of line between anal cleft and posterior spiracles on spiracular disc-pores absent; anal plates with about 6 medio-lateral lobes of abdominal segments 12 n f l nd 4 2 Ge e a i w ic e ege i a isc o es a e ese G OU 2. A E I E GE E A I EW EA A me ia y ac oss mos a omi a segme s ( 1 Dorsum covered in a thick wax test; anal plates 0 Ventral tubular ducts, in addition to forming a broad surrounded by and supported upon a heavily submarginal band, also present medially on all sclerotised caudal process or anal sclerotisation; thoracic segments, even if only near coxae stigmatic spines present in large groups in each (p. 79) ... Crystallotesta stigmatic area, distinctly differentiated from marginal —Ventral tubular ducts present as a submarginal band or setae; dorsum with specialised uni-, bi-, and tri-locular very sparsely scattered submarginally, never medially pores of Ceroplastes-type, each with a long, inner, on all thoracic segments; sometimes with a group near much divided, filamentous duct mouthparts and procoxae 11 (p. 185)... Ceroplastes —Dorsum not covered in a thick wax test; if anal Ventral tubular ducts of two types, a distinctly different sclerotisation present, relatively small and usually smaller type present on either side of anal plates and poorly sclerotised; with only three stigmatic spines per anterior end of anal cleft; normal ventral tubular ducts stigmatic area; dorsal pores never f C r pl t t p , forming a dense band close to margin; anal plates and only uni-locular 2 without a broad sclerotised internal plate; mouthparts usually not displaced to one side (p. 119) ... Kalasiris 2 Pregenital disc-pores present medially on thorax and —Ventral tubular ducts of one type (except a few head as well as on abdomen; ventral tubular ducts degenerate ducts scattered on median abdomen on C. more or less restricted to a submarginal ring, although chelyon), not in above distribution; when present a few may be present medially marginally or submarginally, not forming a dense (p. 206) ... Parthenolecanium band close to margin; anal plates each with an internal —Pregenital disc-pores restricted to abdominal segments sclerotised plate; mouthparts often displaced to one and metathorax; ventral tubular ducts either absent, side (p. 97) ... Ctenochiton restricted to medial area of thorax, or almost as abundant medially on abdomen as submarginally .. 3 Ge e a i w ic e ege i a isc o es o m is i c i es e wee e a a a ea a e os e io s i ac es ( 3 Mature adult female secreting a woolly or felted ovisac 2 Dorsal macropores strongly conical, either `cone-like' posteriorly from beneath abdomen; venter with 3 or 4 or `bollard-like'; pregenital disc-pores with mainly 5- types of ventral tubular duct; ventral tubular ducts 8 loculi; without a distinctly folded area along inner abundant medially on thorax and abdomen; dorsum margins of anal plates; anal plate setae sharply with infrequent small tubular ducts spinose; with fewer than 15 marginal spinose setae (p. 212) ... Pulvinaria laterally between stigmatic areas —Mature adult female not secreting a woolly or felted (p. 171) ... Umbonichiton ovisac; venter with 0, 1, or 2 types of ventral tubular —Dorsal macropores at most slightly convex; pregenital duct, never abundant both medially and submarginally disc-pores with mainly 8-10 loculi (except Plumichiton on thorax and abdomen; tubular ducts usually absent diadema); each anal plate with a strongly folded area from dorsum (but present submarginally on Coccus dorsally along their inner margins; anal plate setae hesperidum) 4 rather spinose and often distinctly blunt; with more than 15 marginal spinose setae laterally between 4 Dorsum often convex at maturity and becoming heavily stigmatic clefts (p. 142) ... Plumichiton sclerotised, generally with dermal areolations; dorsal tubercles and pocket-like sclerotisations present; pregenital disc-pores often present on metathorax as well as abdomen (p. 205, 208)... Parasaissetia & Saissetia —Dorsum usually not highly convex and generally not heavily sclerotised; without distinct dermal areolations; dorsal tubercles present but pocket-like sclerotisations absent; pregenital disc-pores almost restricted to abdominal segment VII (p. 195) ... Coccus 24 d n & nd r n (2000 : C d (In t : pt r : C d o sa e a i e a e a se ae ea e a o e s igma ic a ea s igma ic s i e s i acu a e i eme c y eo a a s ie o. ma gi a se ae e wee s igma ic a eas e a mic o uc s , s i acu a u ow wi isc o es su ma gi a o e su ma gi a se a ma gi a se ae me a o acic eg u u a uc s e a se ae e icu a io i e me io a e a o es u a e icu a io a ea o g ege i a se a mac o o e a oge i a o mic o uc u es & sim e o es ege i a isc o es e icu a io oi se a a a c e se ae os e io me ia mac o o e i e e a a a o e se s ig. . iag amma ic e ese a io o a i ige ous s ecies, wi e si e s owi g o sa s uc u es a ig si e s owi g e a s uc u es. n f l nd 4 2 o sa e a i e a e a se ae su ma gi a se a a e a s igma ic sc e o isa io c y eo a a s ie s igma ic c e s i acu a u ow wi isc o es s i acu a e i eme a ium u u a o sa u u a uc s uc s, y es s igma ic s i e i io a sa a icu a o y sc e osis ma gi a se a e a mic o uc s o sa se ae me a o acic eg e ma a eo a io o g ege i a eo e cu a se ae o es su o i g a o sa u e c e ege i a isc o es ocke ike sc e o isa io a a sc e o isa io a oge i a o a a c e ma gi a a a o e se ae ig. 4. iag amma ic e ese a io o a a e i e s ecies, wi e si e s owi g o sa s uc u es a ig si e s owi g e a s uc u es. 26 d n & nd r n (2000 : C d (In t : pt r : C d ε G , - - ‚ .uΡu, . cy i ica , u Ε. oi e . c a a e G. s i e ike . cu e o sa se ae Κ asa age a e s ou socke i e a se ae Β. mo e a e e g Ο Ρ Α -λ , Ί ι _ Α. e y o g C. g ou o (as i mos a e i e s . n λ S igma ic s i es . im ia e M. s i ose . oa im ia e 0. oi e s i ose . se ose Ma gi a se ae eo e cu a o es ocke ike sc e o isa io Ce o as es y e o es o sa u e c es Fig. 5. iag amma ic e ai ( ig e es o se ae a o sa o es ou o a e i e s ecies. o e: se a e g was measu e mi us ei asa socke s. n f l nd 4 2 Οo Α. sim e Β. sma C. g a u a . c im ey u o s a e um s a e u o s a e owιoΜΙΙwawsyis o sa mac o o es Ε ocke ike mac o o e Co e s a e G. i e e Η. o a s a e ocu a o a 6 ocu a o se i e ocu us isc o es a oo — i ame ous — 1 ocu a 8 ocu a , ocu a , o a i e ocu us a ow i e ocu us o sa mic o uc u es e mi a g a c ose open Ν i e uc cu s a e i agi a io Sim e o es g a mic o uc o e— ou e uc — e a mic o uc s u u a uc s ig. 6. iag amma ic e ai ( ig e es o o sa mac o o es ou o i ige ous s ecies, a o o e o sa a e a o es a uc s. o e: e a mic o uc s a e a i io a y ee aw i a u ig osi io ei o ie a io is e e se e e o s ow a ey o e e a y. 28 d n & nd r n (2000 : C d (In t : pt r : C d eg o g co a — age a e ea e a oc a e emu a ica — se a o e se a es y i ia se ae a ica segme i io a sa a ium a icu a o y sc e o isa io e i eme Ct n h t n s . c aw a sus e ic e — cam a i o m se si a c aw e uce a e a, c aw e ice In l p t ll n h t n s . n sc e o ise n t s i acu a a e sca e A e a S i ac e c aw igi u es a sa igi u es qua a e a a a es y i o m a a a es mi u e o es isca se a i e ma gi se a ou e ma gi se a a ica se ae i e ma gi se a 2 a ica se a ln l p t ll A a a e s uc u es: u e su ace c y eo a a a a i g o e u a o g ege i a se ae s ie i e su o i g a e a a i g se ae a oge i a o a e io ma gi se ae su o i g A a i g a e a ma gi se ae a a ium A. a e i e s ecies Β. i ige ous s ecies Mou a s A a a e s uc u es: u e su ace ig. . iag amma ic e ai ( ig e es o a e a, eg, s i ac e, mou a s, a a a a es. a i es i i ca e ow e g ea was measu e o eg, s i ac e, c y eo a a s ie , a a a a es. n f l nd 4 2 e icu a io ε Ρ" , .. • — i e .. .Υ r ι. , { [Μ8] Ct n h t n p r v r d [Μ ] n h t n t t e icu a io a ea: [Μ1 ] Ct n h t n p r v r d [Μ ] n hlt n t t [Μ 2] Ct n h t n h l n [Μ ] n h t n t t Μ8 . Sca i g e ec o mic og a s (SEMs . Μ8. Ct n h t n p r v r d : o sa iew, e icu a io a e o i es o o sa o es o i ges (sca e a mm . Μ . n h t n t t : o sa iew, ue o sum su ou e y a e e o o sa su ace (sca e a 0. mm . M 0. Ct n h t n p r v r d : d r l iew, se e a e icu a io i es a a eas (sca e a 0. mm . Μ . n h t n t t : o sa iew, i e " ocke ike mac o o es" (sca e a 0 μm . Μ 2. Ct n h t n h l n: i e a iew o o sum, g a u a s uc u es o o sa o es i e icu a io i es (sca e a 0. mm . Μ . n h t n t t :i e a iew o o sum, i e " ocke ike mac o o es" (g a u a s uc u e o e i e (sca e a 0 μm . 3 d n & nd r n (2000 : C d (In t : pt r : C d [Μ 4] U b n h t n j b t [Μ ] n t b l [Μ 6] Aph n h t n n n p [Μ ] U b n h t n p ll p [Μ 8] Ep l d h t n p p r [Μ ] Cr t ll t t rn t Μ 4—Μ . o sa iews o mac o o es. Μ 4. U b n h t n j b t : wo co e s a e mac o o es (sca e a μm . Μ . n t b l : c im ey s a e mac o o e (sca e a μm M16. Aph n h t n n n p : o a s a e mac o o e (sca e a μm Μ . U b n h t n p ll p : s a ke , mus oom s a e mac o o e (sca e a μm . Μ 8. Ep l d h t n p p r :ou e a o ask s a e mac o o e (sca e a μm . Μ . Cr t ll t t rn t : "p pp r o " mac o o e (sca e a μm . n f l nd 4 31 [Μ20] K l r p rf r t [Μ2 ] l h t n p ll n [Μ22] l h t n d d [Μ2 ] Aph n h t n h [Μ24] Ct n h t n h l n [Μ2 ] Aph n h t n p b n Μ20—Μ2 . o sa iews o mac o o es. Μ20. K l r p rf r t : wo i ocu a mac o o es (sca e a μm . Μ2 . l h t n p ll n : e o e i gs o ee i e e , co ca e mac o o es (sca e a μm . Μ22. l h t n d d : Ia ge, sim e o e y e mac o o e (sca e a μm . Μ2 . Aph n h t n h : u o s a e mac o o e (sca e a μm . Μ24. Ct n h t n h l n: g a u a , co e u o s a e mac o o e (sca e a μm . Μ2 . Aph n h t n p b n : mac o o e, um s a e u e ig mic osco e (sca e a μm . 2 d n & nd r n (2000 : C d (In t : pt r : C d [Μ26] Ct n h t n v r d [Μ2 ] l h t n fl v a oo s a e [Μ28] l h t n fl v [Μ2 ] In l p t ll [Μ 0] Ct n h t n v r d [Μ ] n n t Μ26. Ct n h t n v r d : o sa iew, o e sim e o e a wo mic o uc u e o es (sca e a μm . Μ2 . l h t n fl v : o sa iew, ee sim e o es a wo mic o uc u e o es (sca e a μm . Μ28. l h t n fl v : o sum i e a iew, mic o uc u e wi a oo s a e o ima u u e a i ame ous is a u u e (sca e a μm . Μ2 . In l p t ll : c oss sec io o wa es , i e a iew, a es cu away (sca e a 0. mm . Μ 0. Ct n h t n v r d :c oss sec io o wa es , so i wa s uc u e o a wa a e, u i o m ick ess (sca e a μm . Μ . n n t : c oss sec io o es , aye e s uc u e o a sma a e, ickes a o e si e (sca e a μm . n f l nd 4 [C 2] Aph n h t n n n p , 2 [ ] [C ] Aph n h t n h , [f] wi u o ym s isi e [C 4] Aph n h t n p b n , [f] [C ] Aph n h t n bt l , [f] [ o o: M. e e ] 4 d n & nd r n (2000 : C d (In t : pt r : C d [0 6] Aph n h t n pr n , [ff] [0 ] C r pl t d tr t r, [ff] a em y [m] es [ o o: SI ] [C 8] C r pl t n n , 2 i s a um s [0 ] C r pl t n n a u , [ ] (w i e & i s a ym s ( i k n f l nd 4 35 [C40] C h p r d , [ff] [04 ] C h p r d , [ff] a ym s, o ea a ym s, o s em [C42] C l n l , [f] [C4 ] Cr t ll t t f , you g [ ] [ o o: SI ] 6 d n & nd r n (2000 : C d (In t : pt r : C d [044] Cr t ll t t f , o [ ] , s u k o [C4 ] Cr t ll t t n f , o [ ], o e i o a e io o es ( owe e [ o o: SI ] a e io a o es ( owe e [ o o: SI ] [046] Cr t ll t t l pt p r , you g [ ], [C4 ] Cr t ll t t l pt p r , o [ ], (o e , mimics ea sca es o ig s u k o a e io o es [ o o: M. e e ] n f l nd 4 [C48] Cr t ll t t rn t , you g [ ] [C49] Cr t ll t t rn t , ma u e [ ] [C 0] Cr t ll t t rn t ll , you g [ ] [051] Cr t ll t t rn t ll , o [ ] 8 d n & nd r n (2000 : C d (In t : pt r : C d [C 2] Ct n h t n h l n, 2 ma u e [ ] [C ] Ct n h t n p r vlr d , you g [ ] wi c aw e s wi wa i ge [C 4] Ct n h t n p r v r d , ma u e [ ] [0 ] Ct n h t n v r d , [f] a c aw e n f l nd 4 39 [C 6] Ct n h t n p r v r d , [f] a asi ise [C ] " um s" o u e si e ea associa e wi y yme o e ous a ae Ct n h t n p r v r d , [ff] o u e si e [C 8] Ep l d h t n p p n , you g [ ], ma u e [C ] ln l p t ll , you g [ ] [ ] ( a k g ey , a em y [m] es (w i e 40 d n & nd r n (2000 : C d (In t : pt r : C d [C60] K l r d pr , you g [ ] [C6 ] K l r d pr , ma u e [ ] , oo c am e u e os e io a o es [C62] K l r p rf r t , you g [ ] [C6 ] K l r p rf r t , [f] sec e i g wa o e icu a io i es n f l nd 4 41 [C64] n h t n t t , you g [ ] [C6 ] n h t n tr d r , [f] , a e io s i acu a u ows wi w i e wa [C66] n h t n t ll r , [f] a c aw e [C6 ] r t n r , 2 [ ] a ym s o e: co ou a ies wi os , is o e ow [ o o: SI ] 42 d n & nd r n (2000 : C d (In t : pt r : C d [C68] rth n l n rn , [ff] [C6 ] a oge A r t bb r , i es i g o s em o c e y Ct n h t n p r v r d ym s [C 0] a oge p r ll d pl x, i es i g [0 ] a oge rt ll l n , i es i g Ct n h t n p r v r d ym s Ct n h t n p r v r d [ff] a ym s n f l nd 4 4 [C 2] l h t n d d . you g [ ] [C ] l h t n fl v , [f] a em y [m] es [C 4] l h t n l rp , you g [ ] [C ] l h t n l rp , ma u e [ ] 44 d n & nd r n (2000 : C d (In t : pt r : C d [0 6] l h t n p ll n , n [f] [C ] l h t n p ll n , o [ ] [0 8] l h t n n , [f] [C ] r p z d r d , ma u e [ ] a c aw e n f l nd 4 4 [C80] n n t , you g [ ] [08 ] n n t , ma u e [ ] wi eo a e i oo c am e [C82] r p z d r d , you g a u [ ] [C8 ] lv n r fl f r , imma u e [ ] a i s a 4 (ye ow 46 d n & nd r n (2000 : C d (In t : pt r : C d [C84] lv n r h dr n , [f] [C8 ] lv n r br nth , [ff] wi wi o isac [ o o: SI ] o isacs, you g [ ] (g ee , & ym s [C86] lv n r v t , you g [ ] [C8 ] lv n r v t , ma u e [ ] wi o isac n f l nd 4 4 [C88] S t ff , [ff] a ym s [C8 ] S t ff , ma u e [ ] [C 0] S t l , imma u e [ ] [C ] S t l , ma u e [ ] 48 d n & nd r n (2000 : C d (In t : pt r : C d [C 2] U b n h t n d l , you g [C ] U b n h t n b ll t , you g [ ] [C 4] U b n h t n h n nth r , you g [C ] U b n h t n p ll p , n [f] n f l nd 4 4 MO O OGY these genera, it is likely that each reticulation plate grows as the insect expands. Studies of a section through a reticulation plate of Ctenochiton viridis ( ig Μ3 EMA E ( igs A U showed that the wax was of uniform thickness and density The following applies to those species (both adventive and across its width; this suggests that the wax was laid down indigenous) currently known from New Zealand. at a uniform thickness at the edge of the expanding reticulation plate. When sufficiently thin and hyaline, the Basic shape: young teneral females of most Coccidae are colour of the underlying female shows through and, as broadly oval and flat but a few tend to be long and narrow many species are green or brownish, they can harmonise (e.g., Aphenochiton grammicus). A few remain more or with their backgrounds very closely; however, adult less flat as they age (e.g., Coccus hesperidum, Ctenochiton female Ctenochiton chelyon are relatively brightly col- viridis, and all Aphenochiton species) but some expand oured. The test may appear less hyaline and even whitish, laterally, becoming nearly round in outline (e.g., due to striations and bubble-like inclusions (e.g., Kalasiris Epelidochiton piperis), while others often become highly perforata) or brownish, as on Crystallotesta ornata. convex (e.g., Crystallotesta species). The venter of Some tests may be very flat, as on species of species of Lecanochiton expands to form part of the dorsal Aphenochiton, moderately convex, as on Plumichiton surface. species, or highly conical, as on species of Crystallotesta. The tests of Epelidochiton piperis become dull and non- Size: most adult female Coccidae are between about 2 and hyaline at maturity, while those of Poropeza dacrydii (the 6 mm long, but some species may grow to a large size (10 only species of Poropeza for which a test is known) have mm in length), as with Poropeza species. distinct rows of Ctenochiton-like plates but, at maturity, the plates become slightly separated, possibly due to a Tests: the term `test' (derived from Latin testa – a shell) is cessation of wax secretion as the female swells. used to describe the protective waxy covering over the The second group contains species in the genera dorsum, whose function is to reduce both harmful Inglisia and Pounamococcus; these have a glassy test but environmental effects such as dehydration and also the of a different structure to that of the first group. The test on chances of attack by parasites and predators. Although all Inglisia patella is of one piece, 10-sided and shaped like coccids in New Zealand have some form of wax coating the shell of a limpet (Fig. C59); as the insect grows, the over their dorsum, not all these wax coats can be easily height of this test increases due to more wax being added separated from the derm in mature adults and the term test to its base—and the growth-rings produced at each moult is used only for those where the coat is more or less are often distinct. Studies of the test in cross section (Fig. removable. Species without a test include all the adventive Μ 9 s owe a e so i a e o e es is ma e u o species (except the Ceroplastinae) and the indigenous four transverse layers of wax, whereas the walls apparently species of Lecanochiton and Plumichiton elaeocarpi (in have a honeycomb internal section between solid outer part), although nymphs of all species generally have some wax layers. The test of Pounamococcus tubulus is also of form of waxy plates on the dorsum at some stage in their one piece, while that of P. cuneatus is formed from seven development. Most of the other indigenous taxa have a plates, four on the head and three long ones covering the distinct, close fitting, glassy test over the dorsum which is rest of the body (Figs C80, C81). A study of one lateral secreted by dorsal pores, but these tests are not part of the plate of the test of P. cuneatus ( ig Μ31 s owe a i is dorsum and can be easily broken away, leaving the derm made up of layers that are most numerous at the edge intact. nearest the neighbouring median plate (where the lateral The tests on indigenous species fall into two groups: plate is thickest) and decrease to very few layers at the those with a distinctly reticulated, glassy wax cover, and outer margin, thus the plate expands due to secretion of those with some other form of test. new broader layers of wax beneath it, and also The first group have a test divided into numerous, asymetrically from one side. Each of these plates appears more or less 5- to 6-sided glassy plates that are bounded by to be of one piece, without much external ornamentation. distinct sutures; these reticulation plates are usually in 5 The only group amongst the adventive species to have or 7 longitudinal rows, counted across the middle of the a true test is the Ceroplastinae. They have an extremely test (Figs C34, C , Αph n h t n p b n and A. subtilis). thick white soft wax test that not only protects them from This group contains species in the following genera: harmful environmental effects, such as dehydration, but Aphenochiton, Ctenochiton, Crystallotesta, Epelidochiton, also prevents easy control of pest species by insecticidal Kalasiris, Poropeza, Plumichiton, and Umbonichiton. In sprays. 5 d n & nd r n (2000 : C d (In t : pt r : C d O isacs a egg o ec io a true ovisac — a separate margins are referred to as the a e io ma gi s and the cover secreted to protect the eggs—is only produced by os e io ma gi s although the latter is here mainly the Pulvinariini (as far as the New Zealand fauna is referred to as the ou e ma gi to clearly distinguish it concerned) (Figs C84, C85, C87; lv n r fl f r , . from the posterior apex. On most indigenous species, the h dr n , . br nth , and . v t , and is combined plates are not obviously quadrate but are more probably secreted mainly by the ventral tubular ducts. It often pyriform (e.g., those of Ct n h t n species) or oval, takes the form of a white woolly `sac', which protrudes with the anterior margin shorter than the posterior margin. from under the posterior end of the adult female and is On some genera, the inner margins diverge posteriorly and often characteristically ridged, into which the eggs are then most of their spinose setae are along the inner margins laid. It can be quite long—up to several times the length of rather than apically or on the outer margin (e.g., n In l the female — and sticks to the substrate. It is likely that p t ll . On most species, there are two setae along the females of Pulvinariini withdraw their stylets once they inner margins (i e ma gi se ae one or more setae on start egg-laying and move forwards at the speed the ovisac the posterior apex (a ica se ae and another either along is secreted; once egg laying is complete, the females die the posterior outer margin (ou e ma gi se ae or on the and may fall away, leaving the ovisac behind. dorsal surface. On species in some genera (e.g., S t , Protection for the eggs and/or nymphs in the other the dorsal setae are in the middle of the posterior half of subfamilies is provided in other ways. In some taxa, the each plate and, when they are distinctly enlarged, are eggs and/or nymphs are protected beneath the venter, referred to as the isca se ae in the isca osi io which becomes a concave brood chamber, as with species Another feature that may be of taxonomic significance is of S t and Ct n h t n, the females of both genera the surface of the plates, which may be characteristically producing large numbers of eggs beneath their abdomens. ridged or nodulated (e.g., on species of l h t n . Although all species f n h t n have relatively large brood chambers, these seem to contain few young. An A a i g (Fig. 7): is a sclerotised ring, composed of two alternative strategy is found with species of Cr t ll t t lateral crescents, that surrounds the anal opening. (Figs C44, C45, C47), K l r (fig. C61) and Typically, it has numerous wax-exuding pores (which may n (fig. C81) where the ovipositing females be in one or more rows) and 3, 4, or more pairs of long shrink beneath the glassy test so that the shrivelled body setae, the a a i g se ae The anal ring is at the inner end comes to lie in the anterior half and the neonate nymphs lie of the a a u e; when withdrawn and inverted, this tube in the posterior half. lies within the body cavity and usually extends well anterior to the anal plates; rarely the tube is short (e.g., as on r p z species). The following sections deal with the structures found on the dorsum, margin, and venter. A a sc e o isa io (Fig. 4): is a horseshoe-shaped sclerotisation around the anterior margin of the anal plates, O SA S UC U ES e.g., on species of r p z . On members of the Ceroplastinae, the anal sclerotisations have become A a c e (Figs 3, 4): is the cleft between the a a o es at greatly enlarged to form thecaudal or a a ocess which the posterior end of the body, and is usually about 1/7th to carries the anal plates above the thick wax test. 1/10th of the total body length but may be very short in some species (e.g., In l p t ll . A oge i a o (Fig. 7): this fold lies across the anterior end of the anal cleft, more or less at right angles to the long A a a es (Fig. 7): are the paired, approximately axis of the body, beneath the anal plates. It separates the triangular plates, that lie at the anterior end of the anal anus on the dorsum from the vulva on the venter. The cleft, and are sometimes referred to as anal opercula; they anogenital fold usually has setae at either corner and along are a major recognition character of Coccidae as similar the anterior margin and these are here referred to as plates are otherwise found only on a few species of a e io ma gi se ae (also called fringe setae by other Eriococcidae. These plates are `hinged' along their authors). Setae are also generally present along the lateral anterior margin and open anterolaterally to allow the a a margins and these are referred to as a e a ma gi se ae u e to evert, aiding the elimination of honeydew from the On either side of the anogenital fold, there is frequently a a a i g On most adventive soft scales, they are sclerotised supporting bar that appears to be part of the approximately quadrate when combined, with the two underside of each anal plate, and which is probably for the i e ma gi s lying more or less parallel. The outer attachment of the muscles used in opening the anal plates; n f Ν Ζ l nd 4 51 these bars extend anteriorly beneath the derm, where they reticulations being approximately six-sided and underly- frequently expand and may even meet medially; they may ing similarly-shaped e icu a io a es in the test. In the be narrow, as on C h p r d or broad, as on most descriptions below, the following terminology is used: the New Zealand species. In addition, there are sometimes lines of pores are e icu a io i es, the area of derm groups of setae (called y o ygia se ae on the ventral bounded by reticulation lines is the e icu a io a ea, and derm just anterior to the anogenital fold and posterior to the reticulation areas are in e icu a io ows along the the pair of long pregenital setae on abdominal segment II length of the insect; the junction of a reticulation line with (present on several species of Aph n h t n . the margin is a e icu a io oi , often associated with an enlarged marginal seta, the e icu a io oi se a. Most e m: that of teneral females tends to be thin and dorsal pores on the species with a reticulate pattern lie unsclerotised and this is the best stage for making slide within the reticulation lines but their arrangement within preparations for identification. Species with dense each line is also species specific; for instance, each a eo a io s (Fig. 4) (small clear areas in the derm), such as reticulation line in Ct n h t n p r v r d is composed S t ff , may have quite a thick derm, even just of three parallel rows of pores (Fig. M10). On those after the final moult but, even then, the derm becomes species without reticulations, the dorsal pores are more or much thicker with age (up to 10 times thicker than the less randomly distributed. venter), with the area within each areolation remaining There are three main types of dorsal pores on the quite thin, usually with a microduct. The size and indigenous New Zealand species (Fig. 6): distribution of the dorsal areolations may be useful in (a)Dorsal mic o uc u es ( igs Μ —Μ eac co sis s separating species. In the Ceroplastinae, the derm of a small sclerotised pore, round or slightly oval in shape, becomes heavily sclerotised quite soon after the final usua y a ou -3μm i iame e se a e ase o a s o moult and only young females can be easily identified to ductule and with a minute pore or slit-like opening, when species. viewed from above. The true structure is best seen when On indigenous species protected by a glassy test, the viewed from the side (for a discussion of gland structure, derm is thin and membranous, but may become slightly see Foldi, 1997). Each microductule has a non-staining sclerotised with age; exceptions are Ep l d h t n p p r membranous i e uc u e or i ame , that is often (Fig. 115), on which sclerotised patches grow and balloon-shaped proximally and quite long distally. coalesce in each reticulation area, and r p z species, Because the inner filament is non-staining, it can be which live beneath the bark of trees and become difficult to detect in many preparations. In adventive sclerotised throughout, possibly for added protection species, a microductule is usually present in each dermal against pressure from the confined space. areolation, when the latter are present. The derm may also show other characteristic markings (b)Simple o es ( igs Μ Μ 7 sma o es wi ou a and structures. On most indigenous species, the derm has inner filament; represented by at least two types, although distinct polygonal reticulations throughout, these being they may appear similar under the light microscope: (i) partly thickened ridges along the lines of dorsal pores `open' pores that have a distinct pore opening and are flat ( igs Μ M1 O u y-g ow a u s o s ecies suc as (and should not be confused with closed pores that have U b n h t n b ll t and U. j b t , each reticulation lost their structure through being overcleared during slide area on the dorsum becomes convex, forming a lobe, preparation), and (ii) `closed' pores that have no apparent underlying each knob-like projection of the test. aperture under the light microscope but generally have a granulate surface (through which minute ducts emerge) o sa se ae (Figs 4, 5): these are absent from all known and may be either flat or convex (see Foldi, 1997). Most indigenous species but present on all adventive species. a e a ou - μm wi e u ey ca e qui e a o a ge ey a e s o ( μm o 1 μm o g a s i ose a isi g ( 1 μm w i e some a e s ig y icke ma gi s a from a distinct basal socket and are randomly distributed may then appear `dark-rimmed'. They are usually round throughout the dorsum. but may be oval. They have been referred to by previous workers as `dark-rimmed', `disc', or `discoidal'pores. o sa o es (Figs 3-6): most species have at least 3 or 4 (c) Mac o o es ( igs Μ1 Μ1 —Μ 5 is e m is different types of dorsal pores, whose function is probably introduced here to describe some large pores, with a to secrete different types of wax. glandular internal structure (Fig. M12), which appear to be On most of the indigenous species, dorsal pores are highly characteristic of the indigenous Coccidae. They are is i u e i a is i c y e icu a e a e ( igs 3 Μ usually at least twice the size of simple pores but are highly Μ1 e e icu a io s i 5 o 7 o gi u i a ows mos variable between species, both in size and shape, and are 5 d n & nd r n (2000 : C d (In t : pt r : C d therefore very useful diagnostic characters. They may be pores' by De Lotto (1971a). C r pl t t p pores are flattish button-shaped (e.g., Ct n h t n species) or more abundant throughout the dorsum except on the lateral convex and bollard- or cone-shaped (e.g., U b n h t n lobes or clear areas (areas on the dorsum without visible species), or concave as on species of l h t n. When o es a eac o e i g is —5μm wi e ey a e a mos convex, they may extend above the derm surface through certainly involved in the production of the thick, soft, holes in the test (as in U b n h t n p ll p and have waxy test typical of the Ceroplastinae. heavily sclerotised margins and/or basal rings; when concave, they usually have membranous margins and a Dorsal tubercles: also called sub-marginal tubercles. sclerotised base. Macropores are restricted to the They are frequently present on adventive species but reticulation lines when both these are present, and are most absent from all indigenous species. They are rather abundant medially on the abdomen. An indication of variable in structure but those on species currently known frequency may be the number of macropores in the most from New Zealand are convex, usually wider than tall and posterior medial transverse reticulation line with rather sclerotised, with a central duct which has a small macropores, here referred to as the posterior medial swelling/thickening at its inner end, with an inner macropore line (Fig. 3), which is anterior to the anal filamentous ductule on one side, rather similar in structure plates and to an intervening narrow medial line of small to the cup-shaped invagination and inner ductule of a dorsal pores. Macropores are absent from all adventive tubular duct. species, but they may be homologous with the preopercular pores. Pocket-like sclerotisations: sclerotisations which have a (d) Multilocular disc-pores: these are rare on the dorsum pocket-like invagination and mark the site of the dorsal but are present on r p z species on the lateral margins tubercles on the previous instar. These are unknown on of the anal cleft. Occasionally, the spiracular disc-pore indigenous species although the large pores (here referred bands extend onto the dorsum (as on Cr t ll t t f . to as pocket-like macropores) in the two median lines on For further comment, see under ventral pores. n h t n t t and . tr d r have a very The following dorsal pore-types are not present on similar structure. However, pocket-like sclerotisations are indigenous species but are found on one or more of the found on some adventive species of S t and adventive species (Figs 4, 5): rth n l n . When present, pocket-like sclerotis- (e) Preopercular pores: these pores are found in loose ations are generally found in association with a dorsal groups, typically just anterior to the anal plates, although tubercle, either between it and the margin or close by, and they may be much more widespread in some genera and only in a submarginal ring. Their function is unknown. species. They are `closed' pores (see under simple pores Eac sc e o isa io is usua y a ou 5μm wi e above) and are rather variable in shape. They may be small, flat, round to oval, relatively unsclerotised and may Tubular ducts (Fig. 6): within the indigenous fauna, have a granular surface, as on C h p r d , when dorsal tubular ducts are known only from n they look very similar to closed simple pores. In other n t (as two diverging lines anterior to the anal plates) genera, they are large, strongly convex, and heavily but dorsal tubular ducts are often common on some of the sclerotised (as on S t species); these pores can have adventive species, on which they are apparently randomly quite deep, vertical margins, which give them a strongly distributed. Each duct consists of four parts: (a) an outer `dark-rimmed' appearance when viewed from above, and ductule, thin-walled, barely sclerotised, round in cross- have a rough granular surface; they have also been called sec io a ge e a y a eas 1 μm o g w ic o e s `discoidal pores' and `paraopercular pores'. Typically, through the dorsum by a small inconspicuous pore that eac o e is 3-5 μm wi e e um e a s a e may e may occasionally be mildly sclerotised; at its inner end is of both specific and generic importance. Their function is (b) a characteristic structure, here referred to as the cup- unknown. s haped invagination because the outer ductule terminates (f) C r pl t t p pores: these are possibly the only in a thick-walled structure that is bowl- or cup-shaped, and sclerotised pores present on the dorsum of the usually slightly asymmetrical; from one side of the cup Ceroplastinae, to which they are restricted. They are arises (c) the inner ductule which is usually narrower and heavily sclerotised, with a large central pore and 0-4 shorter than the outer ductule; this terminates in (d) a smaller (satellite) pores. C r pl t t p pores generally `flower-head'-like structure, here referred to as the have a long inner filament arising from the base of the terminal gland. Tubular ducts can vary in the relative central pore, which is much branched distally and, because lengths and widths of the inner and outer ductules, in the of the presence of these ductules, were termed `dendritic form (particularly the depth) of the cup-shaped n f l nd 4 invagination, and in the size of the terminal gland. pore band meets the margin, or they may be absent (as on Including those types of tubular duct found on the venter, Epelidochiton piperis and species of Poropeza). On those some Pulvinaria species have four or five types and the species in which stigmatic clefts are absent, each of these structure of each type and their distribution within a points on the margin is here referred to as the s igma ic species are good diagnostic characters. Usually the a ea. dorsum has only one type of tubular duct. S igma ic s i es (Figs 3-5): are one or more marginal MA GI A S UC U ES setae that are usually differentiated from the other setae in each stigmatic cleft or stigmatic area. On almost all Ma gi : on most Coccidae, the margin is distinct and indigenous species except Pounamococcus species, there marked by the presence of marginal setae. On is only one stigmatic spine per stigmatic area, although Lecanochiton, where marginal setae are absent and where rarely one or more of the lateral marginal spines may the lateral margins of the venter spread outwards to form become differentiated, as on Plumichiton pollicinus. On part of the dorsal surface, the demarcation between most other Coccidae, including Pounamococcus species, dorsum and venter is indicated by differences in there are usually three or more stigmatic spines and, when sclerotisation and ridges on the true dorsum (Fig. M9). three, the middle (median) spine is generally longer than the laterals. Most stigmatic spines tend to be set slightly Ma gi a se ae (Fig. 3): these form a marginal line one onto the dorsum but, on Crystallotesta fusca, they are seta wide on almost all New Zealand species (but are in a displaced some distance onto the dorsum. On the band 1-3 setae wide on Poropeza cologabata). They are Ceroplastinae, the stigmatic spines are in a large group in usually of one shape, are distinctly differentiated from each stigmatic area, and may extend medially onto the other setae, and are frequently abundant; however, on dorsum. When stigmatic clefts are present, the stigmatic Inglisia patella two types of setae are present and they spines are located at their base. The number, shape, and usually alternate, while species of Lecanochiton lack relative lengths of stigmatic spines are of taxonomic marginal setae altogether. The shape and structure of these importance. setae are highly variable but are usually constant for a given species. As a result, marginal setae are significant Eyespots (Figs 3, 4): in the Coccidae, the eyespots are taxonomic features at all levels and the number of setae placed on the dorsum, very close to the margin, each laterally between the anterior and posterior stigmatic clefts consisting of a single lens. These can be hard to detect on (or occasionally round the head between the anterior slide-mounted specimens but are often clear on fresh stigmatic clefts) is given in most descriptions as an specimens. indication of their frequency. On most genera, marginal setae are absent from the margins of the anal cleft, but E A S UC U ES occasionally they extend along the entire cleft margin (e.g., on species o Poropeza) or at least part (as on some A e ae (Fig. 7): the basic structure of most normally species of Kalasiris). Also, as on Saissetia coffeae, one or developed antennae is as follows. The basal segment or more marginal setae on the anal lobes may be significantly scape (segment I) is well developed and has three setae; longer than normal marginal setae (ma gi a a a o e segment II (pedicel) has one long and one short seta on its se ae (Fig. 4); these are in addition to the (often long) pair ventral surface and a cam a i o m se si um on the of e a a a o e se ae (Fig. 3). dorsal surface. Between the basal two segments and the Each seta is set in a asa socke (Fig. 5) that is usually terminal three segments there are normally one to four well developed and may be narrow or shallow. On those further segments, although usually it is only the segment New Zealand species which have a reticulate pattern of nearest the apical three segments (or the distal end of dorsal pores on the dorsum, the marginal setae at the point segment III) which has setae and then there are three where the reticulation lines reach the margin are often flagellate setae. On many indigenous species with only six differentiated and are larger. These are here referred to as segments on the antennae, the third can be as long as or e icu a io oi se ae they are particularly obvious on longer than the other five combined. At the apex of each Crystallotesta neofagi. antenna are three segments which are remarkably constant in structure. The a ica segme has about five normal S igma ic c e s (Figs 3, 4): may be distinct clefts, with setae and probably three or four es y se ae (these can parallel sides, as with Aphenochiton pronus, or only appear rather flagellate on some species), two or three on shallow indentations at the point where the spiracular disc- the ventral surface and one on the dorsal; on most species, 5 d n & nd r n (2000 : C d (In t : pt r : C d the terminal seta (apical seta) is rather straight, non- proportion to the rest of the body. The coxae are attached flagellate, and its length appears to be of taxonomic to the venter along their width and articulate with a significance; in addition, one of the flagellate setae on the sclerotisation at the lateral corner; this is often most dorsal side is usually very long, but is less useful obvious on legs that are much reduced. The structure of taxonomically as it is often broken. The subapical the coxae appears to be very similar throughout the segment has a single fleshy seta on the ventral surface and Coccidae; nor is there much variation in the trochanter and a flagellate seta laterally (two setae on n femur. Each trochanter has a pair of large pores on each species), whilst the third segment from the apex has only a side (whose function is unknown) and one or two long single fleshy seta ventrally (plus a flagellate seta on setae on its ventral surface. The setae on the femur have n species). The number of segments on the not been found to vary much. The tibia is always longer antennae can be very constant, as on C h p r d , than the single tarsus and usually the proportions are where it is always seven, or it can be rather variable. similar on all three pairs of legs. The tibia generally lacks Species f n h t n have much reduced antennae and the campaniform sensillum (here referred to as a cam- the segmentation is rather obscure. The structure of the paniform pore) typically present at its base on most other antennae is of taxonomic importance. coccoid families (Koteja, 1974b) but this pore-like structure is present on species of n the Derm: thin and membranous on all indigenous New absence of this pore is otherwise a major taxonomic Zealand species, except on species of n h t n, character of the Coccidae. The tibia and tarsus are usually where the dorsum and the outer expanded margins of the separate but without any articulation on most indigenous venter that form the lateral margins of the dorsal surface species, however they are fused on Aph n h t n bt l , become heavily sclerotised; the ventral surface also In l p t ll , and r p z l b t . Ο o e becomes sclerotised but less than the more dorsal parts. coccids, there is clearly a true tibio-tarsal articulation with The venter may have two or three pairs of shallow, an articulatory sclerosis (as on lv n r species); the radial grooves. In the species known from New Zealand, presence or absence of this sclerosis is of taxonomic there is usually one groove from each peritreme to the importance. The frequency, distribution, and length of the margin, the stigmatic groove or stigmatic furrow (Fig. setae on the tibia and tarsus may also be of some 3), in which most of the spiracular disc-pores lie. A third significance. At the distal end of the tarsus is a pair of thin pair is sometimes present from the base of each antenna to digitules, the tarsal digitules; these tend to be slightly the point on the margin where the eyespots are located, but dissimilar on most New Zealand species, one being this is indistinct or absent on indigenous species. slightly shorter and slimmer than the other; they tend to be shortest when the leg shows signs of reduction. Labium and mouthparts (Fig 7.): the structure of the The structure of the claw and claw digitules show mouthparts is reasonably constant throughout the several features of taxonomic importance. The claw may Coccidae, varying mainly in size—here usually indicated be short and broad or long and thin and may or may not by the length of the clypeolabral shield. The labium is 1- have a small denticle near the apex. The claw digitules segmented (occasionally indistinctly 2-segmented on may both be broad, both be narrow, or of distinctly some nymphs) and cone-shaped, usually with four pairs of dissimilar width. Fine claw digitules are generally setae (Koteja, 1974a), although these are difficult to see on associated with a reduction in the size of the legs. some specimens. The position of the mouthparts is usually central between the anterior legs but on species that lie Segmentation (Figs 3, 4): this is usually reasonably very flat on the leaf lamina and yet feed in the vascular obvious medially on the abdomen and thorax. There are bundles in the main leaf veins, the mouthparts can be six visible segments between the vulva and the displaced asymmetrically as the insects expand (as on metathoracic coxa and these are here numbered segments n and most specimens of Ct n h t n II to VII, following the system of previous authors who species). This is an environmentally induced effect considered that the first visible segment ventrally on the because, when a female settles by a flatter subsidiary leaf abdomen is the 2nd (segment II), the 1st being represented vein, she expands equally on either side of the stylet by an area laterad to the metathoracic legs; thus the attachment point. pregenital segment is the 7th (segment VII). The segmentation on the thorax also usually can be seen, Legs (Fig. : when present (they are absent on including the demarcation between the thorax and head, n h t n , the legs are normal insect legs, each with where the line runs posteriorly to the labium from near five segments, although they are generally slightly small in each procoxa. No segmentation is visible on the head. On n f l nd 4 55 many genera, each abdominal segment has a pair of lobes only five loculi; other taxa, such as some Pulvinaria mediolaterally, so that there is a line of mediolateral lobes species, tend to have seven or eight loculi, whereas the from the anal area to each metacoxa; this line of lobes often most frequent number of loculi per disc-pore is 10; they divides the segmental bands of pregenital disc-pores into a a e usua y 5- μm i iame e ege i a isc- o es median group and/or two lateral groups and this is have been shown to exude short, curved wax filaments considered to be an important character of New Zealand which are hydrophobic and prevent the eggs from sticking Coccidae. These lobes are large and very distinct on together (Foldi, 1997; Hamon et al., 1975); they are species of Ceroplastes and Lecanochiton. therefore least frequent on viviparous species, particularly in the Coccinae. The number of loculi and the number and Spiracles (Fig. 7): each spiracle is composed of a distribution of these pores on the venter are important sclerotised, funnel-shaped outer peritreme, which opens taxonomic characters. through the spiracular opening or atrium into the tracheae. The size of each peritreme is important and can (b) Spiracular disc-pores: are in bands in the stigmatic be useful in placing a species at the generic level; for furrows between the stigmatic area on the margin and the instance, the peritremes of Ctenochiton species are much spiracles. On most species, each pore has five loculi and wi e a e e g o e co ae i e 1 μm o mo e most authors refer to them as quinquelocular pores. wi e w e eas mos e i emes a e e wee a μm However, on species in a few genera, as in Poropeza, the wide (those of Lecanochiton are particularly small, less pores have more than five loculi or have a variable number a μm O some s ecies e e is a sc e o isa io of loculi and so these pores are here referred to as around each spiracle mesad to the peritreme and this is spiracular disc-pores. Most spiracular disc-pores are here referred to as the sclerotised spiracular plate, as on a ou - μm wi e As wi ege i a isc- o es e Inglisia patella. distribution and frequency of spiracular disc-pores and the number of loculi/pore are important taxonomic characters. Ventral pores Spiracular disc-pores exude short, curved, wax filaments (i) Disc-pores (Fig. 6): each disc-pore has a central loculus that are hydrophobic and help in gas diffusion along the which is usually round but may be oval, surrounded by a stigmatic furrows (Foldi, 1997) and this wax can be seen number of similarly-shaped loculi or pores, the complete even in the thick wax covering the Ceroplastinae (Tamaki disc-pore looking rather like a wheel with spokes. Each et al., 1975). disc-pore usually has 5-10 loculi in the outer ring and the Each band of disc-pores is referred to as the pores are therefore known as multilocular disc-pores. spiracular disc-pore band and, on species with narrow Multilocular disc-pores can be divided into two bands, each band is often more or less of equal width groups: the pregenital disc-pores and the spiracular throughout its length but, on other species, the pore bands disc-pores. widen near the peritremes and margins (as on several (a) Pregenital disc-pores: are primarily located on the Ctenochiton species). On Inglisia patella and Lecanochiton pregenital segment VII, thus the name. However, on most species, the posterior band is reduced to a few pores near genera, they are also found across some of the more each peritreme, although the anterior bands are complete. anterior abdominal segments and, less frequently, On most species, the disc-pore bands end medially at or medially on the thorax and head (where they are referred to close to the peritremes of each spiracle but on Poropeza as multilocular disc-pores). Thus, on Coccus hesperidum cologabata, they extend medially well past the anterior for instance, pregenital disc-pores are almost entirely coxae. restricted to the pregenital segment, while on members of the Saissetiini and species of Crystallotesta, Ctenochiton, (ii) Ventral microducts (Fig. 6): the oval, sclerotised pore and Kalasiris, they are typically abundant across all the of each ventral microduct is located at the base of a short, abdominal segments; on Crystallotesta, they are also often outer ductule (longer than that of the dorsal microductules). present laterad to the metacoxae. On species of The pore opening is across the widest part of the pore and Plumichiton and Umbonichiton the pregenital disc-pores the non-staining inner ductule is usually broad and skirt- are mainly on the mediolateral lobes and so form a line ike o i ame ous Eac o e is usua y a ou -3 μm from near the anterior margins of the anal cleft to near each wide. Generally, ventral microducts are present more or posterior spiracle. On some taxa (e.g., Inglisia patella and less throughout the venter, although they may be more species of Aphenochiton), the pregenital disc-pores are frequent submarginally; occasionally they have distinctive restricted to either side of the genital opening or near the distributions. On Kalasiris depressa and Epelidochiton margins of the anal cleft and then each pore tends to have piperis, those nearest the margins are distinctly larger. On 5 d n & nd r n (2000 : C d (In t : pt r : C d the Ceroplastinae, the pore opening appears to be Pulvinariini, which have three or four different types of cruciform. The function of the ventral microducts is ventral tubular duct, each with its own distinctive uncertain. distribution. These ducts are most frequently distributed in one of three patterns: (i) restricted to a more or less (iii e a e a o es (Fig. 7): these are small, convex complete submarginal band, as on Saissetia oleae; (ii) pores that are present just anterior to each scape. Their more or less throughout, as on species of Crystallotesta function is unknown. and Poropeza; or (iii) in a group medially on the abdomen, as on Pounamococcus. (i O e e a o es: other types of pore are infrequent on the venter and are therefore good taxonomic u a (Figs 3, 7): this is the female genital opening and is characters. Sim e o es (Fig. 6), similar to those on the found on abdominal segment VII. Whilst the derm dorsum, are occasionally present; they are most frequently surrounding the opening of the u a is thin and associated with the margin (as on Crystallotesta ornata membranous and obscured by the anal tube, it can and C. ornatella) but may have a wider distribution, as on occasionally be found just anterior to the anogenital fold. Poropeza cologabata, or be present with spiracular disc- On Inglisia patella and species of Lecanochiton, the pores, as in the anterior stigmatic furrows of Lecanochiton anterior margin of segment II and the vulva lie some minor and L. scutellaris. distance anterior to the anal area, near the centre of the abdomen. e a se ae (Figs 3-5): most ventral setae are short and flagellate, but a few are longer and these (and their For a detailed description of the structure of the Coccidae, frequency) can be of some taxonomic significance. The see Hodgson (1994a). most common distribution of the longer setae is: a pair medially on the pregenital segment (segment II (the ege i a se ae and often with additional pairs on the preceding two segments (segments VI and V); in addition, there may be long setae elsewhere, such as just mesad to each coxa and between the antennae (i e a e a se ae . On Inglisia patella and species of Lecanochiton, the pairs of long setae on the pregenital segments are replaced by groups or segmental rows of short, rather spinose, setae. Some other groupings of setae are now thought to be of significance and are here referred to as follows: the su ma gi a se ae are a single row of setae just mesad to the margin and their frequency is given as the number laterally between the stigmatic clefts; e a e io a a c e se ae are a group of 1 to several setae that occur on either side towards the anterior end of the anal cleft; a pair of ventral a a o e se ae occur on the anal lobes near the margin and their length appears to be taxonomically significant; and the a omi a se ae are those found medially on each abdominal segment and their frequency are also thought to be significant. Details of these setae (and others on the antennae and legs) are mentioned in most of the descriptions below. e a u u a uc s (Fig. 6): the structure of the tubular ducts on the venter is similar to that of those on the dorsum and, like them, their structure and distribution are important taxonomic characters. When present, most species have only one type of dorsal and one type of ventral tubular duct and then they are usually identical but some species may have several types, as on members of the n f l nd 4 57 Ge us A E OC I O e e so & reticulation setae rarely differentiated from marginal o gso setae; marginal setae on anal lobes not differentiated. ew ge us Stigmatic clefts absent on A. grammicus, otherwise distinctly invaginated and narrow, without stigmatic y e s ecies Inglisia inconspicua Maskell (here sclerotisations; each cleft with a single stout, spinose designated). stigmatic spine, never longer than about 4 x length of Diagnosis. Adult female. Test: glassy, flat to slightly larger marginal setae. Eyespot present or absent. convex, hyaline, thin and brittle, showing a reticulate Venter: pregenital disc-pores with 5-9 (mainly 5-6) pattern, reticulation plates all subequal in size. outer loculi and a single round central loculus; distributed Shape: mature adults of moderate size, less than 7 mm on either side of anterior end of anal cleft and occasionally in length, elongate oval to almost round, flat to slightly with a few on mediolateral lobes of preceding segments convex, with a shallow anal cleft; colourless to light green. (across most posterior segments on A. inconspicuus). Dorsum: derm membranous. Dorsal setae absent. Spiracular disc-pores with mainly 5 loculi, in narrow Dorsal pores distributed in a distinctly reticulate pattern, bands l-4 pores wide between spiracles and margin, each delineating a reticulation area underlying each wax plate band generally broadening near spiracle but with a few of test; these reticulation areas in? longitudinal rows, with extending medially past peritreme on some species. 7 to 10 areas in median row between anal plates and Preantennal pores: 0-2 present. Ventral microducts anterior margin and 27-29 areas around margin; each present in a broad marginal band and medially on head, reticulation area with few discernable pores. Dorsal pores thorax, and abdominal segments II — I Ventral tubular of 3 or 4 main types: (i) small to minute, dark ducts absent. Ventral setae: with 0-7 pairs of small microductules, with a long inner filament, consisting of a anterior anal cleft setae, forming a distinct line along narrow balloon-like proximal end and a filamentous distal margins of anal cleft on some species; hypopygial setae end: restricted to lines of reticulation; (ii) small simple present or absent; with lto several pairs of long pregenital pores, often of 2 sizes: mostly present just laterad to lines setae on segment VII and usually also on segment VI and of reticulation but also occasionally present medially in frequently with medium-sized setae on these and more reticulation areas, particularly in more lateral areas and anterior segments; other setae distributed as for family. sometimes forming a distinct band just dorsad to marginal Antennae well developed, 6-to 8-segmented, 3rd segment spines; and (iii) macropores, of 3 basic shapes: either (a) much longest; when 6-segmented, with 0-3 `button-shaped', quite large (at least twice width of pseudosegments; setal distribution as for family. pregenital disc-pores) and rather flat, with a distinctly Mouthparts occasionally displaced to one side. Spiracles granulate surface, smaller on A. pronus; or (b) `drum- typical of family. Legs well developed, generally with a shaped' much smaller (subequal in size to pregenital disc- separate tibia and tarsus (fused on A. subtilis) but no tibio- pores), distinctly convex with a heavily sclerotised border; tarsal articulatory sclerosis; distribution of leg setae as for or (c) very large and convex, `bollard-like', heavily family; claws small and short, without a denticle; claw sclerotised, often with an inner duct (on A. inconspicuus); digitules similar and broad; tarsal digitules dissimilar, one all 3 types restricted to reticulation lines. Preopercular shorter and narrower than other; tarsal campaniform pores pores, dorsal tubercles, and dorsal tubular ducts absent. absent. Vulva opening on posterior of segment II Anal plates together broadest in anterior third, tapering to apex, each plate with 2 finely spinose setae along inner Remarks. This genus contains nine species: Aphenochiton margin near apex, 1 longer seta ± apically and another on chionochloae Henderson & Hodgson, n. sp., A. dierama outer margin or upper surface near apex; with no anal Henderson & Hodgson, s A. grammicus Henderson sclerotisation on dorsum around anterior margins of anal & Hodgson, s A. inconspicuus (Maskell), com plates. Anogenital fold with 2 large sclerotised plates A. kamahi Henderson & Hodgson, s A. matai arising internally and extending anteriorly from anterior Henderson & Hodgson, s A. pronus Henderson & margin; with 3-5 pairs of long setae along anterior margin Hodgson, s A. pubens Henderson & Hodgson, s and a single seta on each lateral margin. Anal tube and A. subtilis Henderson & Hodgson, s This group moderately long; anal ring with 3 pairs of setae. is characterised by: Margin: marginal setae in a single line extending (i) the complete absence of ventral tubular ducts; around margin to anal cleft (but not along anal cleft (ii) pregenital disc-pores with 5-9 (generally 5-6) loculi; ma gi s ; i e y s i ose ( oa y s i ose o Α (iii) rather few marginal setae, with 3-16 between the inconspicuus), with a fairly sharp apex and a narrow basal lateral stigmatic clefts, each usually finely spinose; socket; with 3-16 between lateral stigmatic areas; (iv) dorsal reticulations in 7 longitudinal rows; 5 d n & nd r n (2000 : C d (In t : pt r : C d (v) stigmatic clefts usually distinct; 4 Body of mature female large, at least 4 mm long; with (vi)dorsal macropores either bollard-like (A. n n p , more than 50 spiracular disc-pores in each disc-pore or flat-convex and quite large, or smaller and more a ; a e ae mo e a μm o g convex. h n hl Aph n h t n is one of the New Zealand genera in —Body of mature female small, less than 3 mm long; with which the species have pregenital disc-pores which are less than 30 spiracular disc-pores in each disc-pore mainly 5-locular. They differ from all similar species a ; a e ae ess a 3 μm o g t except Ep l d h t n p p r in lacking ventral tubular ducts. Species in the genus Aph n h t n differ from E. p p r in: Dorsal macropores abundant and widespread, extending (i) lacking sclerotisations in each reticulation area; laterally to near margin 6 (ii) the absence of a horseshoe-shaped anal sclerotisation —Dorsal macropores few and restricted to reticulation on dorsum anterior to the anal plates; lines around median 3 rows of reticulation areas 7 (iii) the position and length of the anal plate setae; (iv) having 1 type of ventral microduct, and 6 Body distinctly elongate, length usually at least 2.5 x (v) the absence of multilocular disc-pores near the width; abdominal segment VI with only short mouthparts. pregenital setae; with only a single pair of anterior anal The species currently placed in Aph n h t n are all cleft setae; anterior margin of anogenital fold with 3 endemic to New Zealand. pairs of setae r Generic name derivation: the name refers to the —Body oval, length usually less than 2 0 x width; inconspicuousness of these species: ph n (Gr.) abdominal segment VI with at least l pair of long meaning unseen, invisible, secret, obscure, and h t n pregenital setae; with 3+ pairs of anterior anal cleft (Gr.) tunic or garment worn next to the skin. setae; anterior margin of anogenital fold with 4 pairs of setae d r Legs with a separate tibia and tarsus; apical segment on Key o a u ema e Aph n h t n antenna short, length about 2.5 x width; anterior anal cleft setae not resembling a pair of inwardly pointed Marginal setae strongly spinose, rather lanceolate in combs. 8 shape, tapering to a fine point; pregenital disc-pores —Legs with tibia and tarsus fused; apical segment on tending to spread across most abdominal segments.. a e a o g e g a eas Χ wi ; a e io a a inconspicuus cleft setae rather long and stiff, resembling a pair of —Marginal setae finely spinose, narrow at base; inwardly pointed combs bt l pregenital disc-pores restricted to laterad to anogenital fold and to mediolateral folds of most abdominal segments 2 8 Dorsal macropores quite large, at least 2 x width of pregenital disc-pores, mainly in reticulation lines along margins of median row of reticulation areas, 2 With more than 4 marginal setae on each side between e ce o a ome ; egs s o 15 -1 5 μm stigmatic areas; hypopygial setae present 3 h —With 3 marginal setae on each side between stigmatic —Dorsal macropores small, about size of pregenital disc- areas; hypopygial setae absent 5 pores, and present around margins of median 3 rows of e icu a io a eas; egs o g 1 5- 9 μm pr n 3 Pregenital disc-pores restricted to pregenital segment only p b n —Pregenital disc-pores extending onto mediolateral folds of most or all abdominal segments 4 n f l nd 4 ig. 6. Aph n h t n h n hl e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . 60 d n & nd r n (2000 : C d (In t : pt r : C d . . Aph n h t n d r nd r n & o gso , . s ., a u ema e. n f l nd 4 6 ig. 8. Aph n h t n r e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . 62 d n & nd r n (2000 : C d (In t : lpt r : C d ig. . Aph n h t n n n p (Maske , . com ., a u ema e. n f l nd 4 6 . 00. Aph n h t n h e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . 64 d n & nd r n (2000 : C d (In t : pt r : C d ig. 0 . Aph n h t n t e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . n f l nd 4 6 . 02. Aph n h t n pr n e e so & o gso , . s ., a u ema e. 66 d n & nd r n (2000 : C d (In t : pt r : C d . 0 . Aph n h t n p b n e e so & o gso , . s ., a u ema e. n f l nd 4 6 . 04. Aph n h t n bt l e e so & o gso , . s ., a u ema e. 68 d n & nd r n (2000 : C d (In t : pt r : C d Aph n h t n h n hl e e so & o gso setae; with pairs of long pregenital setae restricted to ew s ecies segment VII (one specimen had 1 long seta on VI); number of setae medially on abdomen: VII, 2 long, 0-3 medium + Fig. 96 l-5 short; VI, 0-1 long, 6-10 shorter; V, 7-14 short; IV, Unmounted material: only known material is of dry adult 10-17; IlI, 9-13; and II, 7-9; with 9-13 setae medially on females removed from host plant; females elongate-oval metathorax; 6-9 medially on mesothorax and l-2 setae and rather flat. near each procoxa, all rather short, length of setae associa e wi eac oco a 5-7 μm; wi 3- ai s o Mounted material: body elongate-oval, with shallow inter–antennal setae; with 5-11 submarginal setae on each stigmatic clefts; anal cleft quite deep, about 1/6th body side between stigmatic areas. Antennae 6- to 8- length; length 4.2-6.3 mm; breadth 2.0-2.7 mm. segmented, with 0-2 pseudosegments in 3rd segment w e -o 7-segme e ; o a e g 71-5 7 μm; e g Dorsum: dorsal pores in a reticulate pattern, probably o a ica segme 3 -53 μm; e g o a ica se a 5- with 7 rows of reticulation areas across dorsum, with about μm Mou a s o a a e y is ace o o e si e; 8-9 reticulation areas between anal plates and anterior e g o c y eo a a s ie 1 -195 μm o g Wi o margin and perhaps 29 areas around margin. Dorsal pores s i acu a e i emes a e io - 3 μm os e io 55- 1 of 3 or 4 types: (i) microductules: frequent within μm egs wi a se a a e i ia a a sus; e g s reticulation lines and along margins of anal cleft, where (me a o acic co a 1 7- 3 μm oc a e + emu perhaps slightly larger; (ii) simple pores of 2 sizes: smaller 39-317 μm i ia 19 - 5 μm a sus 1 -1 μm c aw pores subequal or slightly larger than pore of 1 - μm microductule, frequent in all reticulation lines and throughout more marginal reticulation areas; larger pores: Material examined: HOLOTYPE [f]: NEW ZEALAND: perhaps more convex, scarce near reticulation lines and in NC: Mt Oxford, [3,500 ft] 1060 m, 8 Feb 1955, W.R. a sparse submarginal band; and (iii) relatively small Boyce, Ch n hl fl v n , #90-214d, NZAC: 1/ mac o o es (3-5 μm iame e — su equa i wi o 19 ad. ege i a isc- o es ea i y sc e o ise a ig y PARATYPES [ff]: NEW ZEALAND: NC: as for co e mos a u a i me ia a su me ia holotype, #90-214a–c, e–i + 3 slides, W. Cottier e icu a io i es a se ea ma gi a a e io y; wi Collection, as previous except [as nth n fl v n ]: - mac o o es i e os e io me ia mac o o e i e 11/11 [ff] ad. A a a es 15 -17 μm o g com i e wi s 153-1 9 μm; eac wi -7 mi u e o es o o sa su ace; e g Other material: none known. o se ae i e ma gi 1 7-9 μm; i e ma gi 9-11 μm; a ica - μm; ou e ma gi 1 - μm A oge i a Remarks. A. h n hl is very similar to A. p b n . o wi ai s o se ae a o g a e io ma gi o a a o However, A. h n hl has: a 1 ai a e a y; o ges 5 - 1 μm (i) more pregenital disc-pores (15-35 as compared with 5- Margin: marginal setae finely spinose, with 8-13 on 21 in A. p b n , which are also present mediolaterally each side between stigmatic clefts; setae of 2 sizes: most on some of the preceding abdominal segments; se ae sma a i e 1 - μm o g u ose us (ii) fewer (3-6) hypopygial setae than A. p b n (which os e io o s igma ic c e s o ge a mo e s i ose 3- has 5-16); 31 μm o g; e icu a io se ae o o icea y a ge (iii) s o e a a o e se ae (37-51 μm o A h n hl S igma ic s i es o a e u i o m ick ess ge e a y as agai s 7 -1 μm o A p b n e wi a u a e ; e g 57-7 μm (iv) legs, antennae, and spiracular peritremes all 20-25% Venter: pregenital disc-pores with mainly 5-6 loculi larger than on A. p b n (range 4-8): with (on each side of segment): VII, 15-35; (v) twice as many disc-pores in each spiracular disc-pore VI, 2-5; V, 1-4; IV, 0-2; III, 0-2; and II, 0-1. Spiracular band (usually more than 60 on A. h n hl but disc–pores: with 47-98 in each anterior band and 61-97 in less than 45 on A. p b n . each posterior band; each band becoming 5-6 pores wide near spiracle and margin; anterior bands with 0-3 pores Biology. Unknown. present a short distance mesad to peritremes. Ventral microducts as for genus. Ventral setae: ventral anal lobe Distribution. Currently only known from the original se ae mo e a e y Io g eac 37-51 μm; wi 1- ai s o collection off Ch n hl from Mt. Oxford in the South a e io a a c e se ae; wi a g ou o 3- y o ygia Island (Map 1). n f l nd 4 6 Name derivation. This species has been named after the e a se ae e a a a o e se ae 5 - 1 μm o g; wi genus of its host plant, Ch n hl which in turn means 4-5 pairs of anterior anal cleft setae, not in a distinct line; `snow grass'; C. fl v n is the broad-leaved snow with 2-3 long setae present medially on segment VII only tussock. but with medium length setae sometimes present on more anterior segments; number of short and medium length setae medially on abdomen: VII, 5-10; VI, 4-13; V, 5-10; IV, 14-16; III, 16-22; and II, 8-15; with 11-22 setae medially on metathorax, 8-11 medially on mesothorax and 3-5 near each procoxa; length of setae associated with Aph n h t n d r e e so & o gso ew oco ae 5- μm o g; wi 3-5 ai s o i e –a e a s ecies setae; with 3-6 submarginal setae on each side between stigmatic areas. Antennae: occasionally with a pseudo- ig. segment on segment ΙΙΙ o a e g -51 μm; e g Unmounted material: unknown. o a ica segme 53-57 μm; e g o a ica se a 5 - 1 μm e g o c y eo a a s ie 1 -1 μm Wi o Mounted material: body broadly elongate-oval; with s i acu a e i emes a e io 3 - 5 μm os e io 3 - 9 distinct, moderately deep stigmatic clefts; anal cleft about μm egs wi a se a a e i ia a a sus; e g s 1/6th-1/7th body length; length 3.0-6.6 mm; breadth 1.8- (me a o acic co a 17 - 11 μm oc a e + emu 4.7 mm. 11- 9 μm i ia 1 - 3 μm a sus 99-1 μm c aw 1 -19 μm ooke a o c aw e y s o Dorsum: dorsal pores in a reticulated pattern, with 7 longitudinal rows of reticulation areas across dorsum, with Material examined. HOLOTYPE [f]: NEW ZEALAND: 8 areas between anal plates and anterior margin and with NC: Arthur's Pass, 23 Jan 1983, J.M. Cox, under bark 27 areas around margin. Dorsal pores of 3 types: (i) r ph ll , #161, NZAC: 1/3 [ff] (holotype is nearest microductules: frequent within lines of reticulation, each the locality label and is clearly marked). often in an areolation; also in a line on either side of anal PARATYPES: as for holotype, the other 2 females on cleft; (ii) flat, simple pores, rather variable in size but slide. mainly a little larger than microductules: frequent along margins of all reticulation areas; and (iii) large macropores Other material: New Zealand: NN: Mount Lodestone, ( -11 μm wi e—a eas wi o ege i a isc- 1160 m [as 3800 ft], r ph ll tr v r , 17 Nov o es ea i y sc e o ise a o s ig y co e wi a 1969, J.A. de Boer, #604: 3/4 [ff] ad. Mt. Arthur, Flora granulate surface: abundant in all reticulation lines except Track, r ph ll sp., 8 Feb 1982, C.F. Butcher, #83- near margin; with 7-12 macropores in posterior medial 300g: 1/1 [f] ad.OL:Makarora d rp [as Makaroa], mac o o e i e A a a es 17 - 1 μm o g com i e h ll , 1 Feb 1983, J.M. Cox, #227: 2/2 [ff] ad. wi s 1 - μm; eac wi -1 mi u e o es o o sa su ace; e g o se ae i e ma gi 1 7- μm; Remarks. A. d r is one of the Aph n h t n species i e ma gi 9-13 μm; a ica 13-1 μm; ou e ma gi (along with A. r and A. h which have 1 -1 μm; a ea a a a e a e A oge i a o wi large, rather flat, dorsal macropores. It can be separated 4 pairs of setae along anterior margins and a single pair from the other two species by the following combination a e a y; o ges 5- 3 μm of characters: Margin: marginal setae finely spinose, with 3 on each (i) dorsal macropores abundant in all reticulation lines si e e wee s igma ic c e s; e g 1 -3 μm ose us except near margin; posterior to stigmatic clefts usually much larger and (ii) with 2-3 long pregenital setae, restricted to segment distinctly spinose. Stigmatic spines of rather uniform VII; thickness, often rather bent with a blunt apex; length 36- 7 μm (iii) with 27 reticulation areas around the margin; Venter: pregenital disc-pores: restricted to segment (iv) marginal setae just posterior to each stigmatic cleft VII, in groups of 11-21 on either side of anterior end of rather large and spinose; anal cleft. Spiracular disc-pores: in bands 1-2 pores wide; (v) body much broader than A. r . with 38-62 in each anterior band and 37-64 in each posterior band; with 0-2 pores present mesad to each Biology. Likely to have only one generation a year, as all anterior peritreme. Ventral microducts as for genus. collections are of adult females around midsummer. 7 d n & nd r n (2000 : C d (In t : pt r : C d Distribution. Only known from montane localities in the anterior band and 14-34 in each posterior band; anterior South Island (Map 2). bands with 1-3 pores present just mesad to peritremes. Ventral microducts present throughout except in a narrow Name derivation. The name of this species is taken from marginal band and medially on abdominal segments VI the Greek dierama, meaning a sieve or strainer, referring and VII. e a se ae e a a a o e se ae 5-5 μm to the large number of dorsal macropores, and is used as a long; with 2-4 pairs of anterior anal cleft setae rather noun in apposition. randomly distributed; hypopygial setae absent; with a pair of long pregenital setae restricted to segment VII; number of short setae medially on abdomen: VII, 4-6; VI, 3-7; V, 4; IV, 9-14; III, 9-11; and II, 9-13; with 6-7 setae medially on metathorax, 2 near each mesocoxa and 2-3 Aph n h t n r e e so & o gso ew near each procoxa, length of setae associated with each s ecies oco a 9-11 μm; wi 3- ai s o i e —a e a se ae plus I seta medially; with 7 submarginal setae on each side Fig. 98 between stigmatic areas. Antennae: total length 279-340 Unmounted material: adult female very thin and μm; e g o a ica segme 3 - μm; e g o a ica transparent and almost covering the width but not the full se a 37- 7 μm e g o c y eo a a s ie 1 7-155 length of the host plant leaf. μm Wi o s i acu a e i emes a e io 3-33 μm os e io 5-3 μm egs wi a se a a e i ia a a sus; Mounted material: body rather elongate-oval, anterior e g s (me a o acic co a 7 -117 μm oc a e + end acuminate and posterior end narrow; without emu 131-19 μm i ia 1 -1 μm a sus 73-1 μm stigmatic clefts; anal cleft quite deep, about 1/6th body c aw 13-1 μm length; length 3.3-4.2 mm; breadth l.0-1.3 mm. Material examined: HOLOTYPE [f]:NEW ZEALAND: Dorsum: dorsal pores in a reticulate pattern, with 7 TO: Pureora Forest Lodge, 10 Jan 1995, R.C. Henderson, rows of reticulation areas across dorsum, with 8-9 Dracophyllum subulatum lvs, NZAC #95-012: 1/1 [f] ad. reticulation areas between anal plates and anterior margin and with 29 areas around margin. Dorsal pores of 4 types: Other material: NEW ZEALAND: GB: Kakanui, 300 m, (i) microductules: within reticulation lines; (ii) simple Dracophyllum sinclairii, 1 Feb 1993, RCH, #93-206a—b: pores, smaller than pore of microductule and flattish, 2/2 [ff] ad, 1 [m]2nd, 1 1st. scattered within reticulation lines and within marginal reticulation areas; (iii) slightly larger, rather convex, Remarks. A. grammicus is closest to A. dierama a Α simple pores, occasionally within reticulation lines and kamahi in having moderately large, rather flat dorsal near margin; and (iv) moderately large macropores, macropores, but is separated from the other two species by ea i y sc e o ise a a o s ig y co e 5-7 μm the following combination of characters: wide: most abundant medially, becoming less frequent (i) its very elongate shape (the other species tend to be near margin and anteriorly; with about 4 macropores in more oval); posterior medial macropore line. Anal plates: 156-189 (ii) dorsal macropores frequent to abundant in all μm o g com i e wi s 133-171 μm; eac wi 5- reticulation lines except near margin; minute pores on dorsal surface; length of setae: inner (iii) antennae without pseudosegments; ma gi 1 7-9 μm; i e ma gi 10-13 μm; a ica 1 - 1 μm; ou e ma gi 1 -1 μm A oge i a o wi 3 (iv) a single pair of long pregenital setae on segment VII, pairs of setae along anterior margin and a single pair all other setae medially on abdomen short; a e a y o ges 7- 7 μm (v) 3 pairs of setae along anterior margin of anogenital Margin: marginal setae finely spinose, with 3 on each fold. si e e wee s igma ic c e s e g 1 -13 μm wi ose us os e io o s igma ic s i es o ges u o 3 μm Biology. Unknown. Stigmatic spines of rather uniform thickness, often rather e wi a u a e ; e g 39- 5 μm Distribution. Restricted to narrow-leaved Dracophyllum Venter: pregenital disc-pores with mainly 5 outer spp. and collected in the East Cape region near e Araroa loculi; in groups of 5-7 on either side of anterior end of and on the frost flats at Pureora in central North Island anal cleft only. Spiracular disc-pores: with 16-28 in each (Map 3). n f Ν Ζ l nd 4 71 Name derivation. This species is particularly long and μm; a ica (o a e 13-31 μm a ou e ma gi (o narrow and the specific name is taken from the Greek o sa su ace o a es 1 - μm A oge i a o wi grammikos meaning linear. ai s o se ae o a e io ma gi a 1 ai a e a y; o ges 5 - 1 μm Margin: marginal setae moderately strongly spinose, with 3-16 setae on each side between stigmatic clefts; Aph n h t n n n p (Maske ew setae usually curved and tapering to a fine point, of widely com i a io varying size, but with poorly developed basal sockets; igs Μ1 C3 99 those just posterior to each cleft sometimes set slightly Inglisia inconspicua Maskell, 1892: 19; —Maskell, 1895a: onto dorsum and larger; those at reticulation points also 14 [checklist]; -Cockerell, 1896: 330 [checklist]; — larger; with 3 larger reticulation setae on each side Fernald, 1903: 162 [world catalogue]; —Hutton, 1904: 227 e wee s igma ic c e s eac 3 -3 μm o g a wi 3- [checklist]; —Myers, 1922: 199 [checklist]; -Wise, 1977: 13 sma e se ae e wee c e s e g 1 - μm 105 [checklist]; —Deitz & Tocker, 1980: 29 [checklist]; - S igma ic s i es a e i g o a oi o e s ig y cu e Ben-Dov, 1993: 148 [world catalogue] wi a a ow asa socke ; se s ig y o o o sum; e g 1 -1 μm Eyes o usua y ai y is i c μm Unmounted material: "Test of adult female white, wi e us o sa o ma gi a s i es elongated, narrow, convex, not conical, with [marginal] Venter: pregenital disc-pores with mainly 5 (range 3- fringe [of wax plate filaments] either absent or very small; 6) loculi; distributed across most posterior abdominal texture glassy, very thin and delicate and brittle; segments segments; number on each segment (medially across each polygonal, marked with very delicate radiating striae of air segment/in each lateral group): VII, 0-2/5-23; VI, 9-12/ cells and still finer concentric lines. Length of test 0-5; V, 11-16/0-4; IV, 7-12/0-3; III, 2-7/0; and II, 1/0; averaging about 1/5 in. [5 mm]; height 1/30th in. [0.8 none laterad to metacoxae. Spiracular disc-pores with 3- mm]. Although the test itself is white, the general 6 (mainly 5) loculi in bands of variable width; with 30-53 appearance of the insect on the twig is brown, the colour of in each anterior band and 23-62 in each posterior band; the insect showing through the translucent segments." with 0-3 pores extending medially past peritreme. Ventral "Adult female brown or reddish-brown, filling the test but microducts of 1 type: present rather sparsely throughout shrivelling at gestation." (Maskell, 1892, p. 19). except absent medially on posterior 4 abdominal segme s e a se ae e a a a o e se ae 1-7 μm Mounted material: body elongate oval; anal cleft about l/ o g; wi 3- ai s o a e io a a c e se ae; y o ygia 9th body length; stigmatic clefts shallow but distinct; se ae a se ; wi 1 ai o o g ege i a se ae ese o length 2.0-5.0 mm; breadth 1.35-2.45 mm. segme II o y; o ges a ou 5 μm; um e o se ae Dorsum: dorsal pores in a reticulate pattern, with 7 me ia y o eac a omi a segme II o g - -9 rows of reticulation areas across dorsum, 9 reticulation me ium; I - me ium (occasio a y s ig y a ge ; areas between anal plates and anterior margin, and 29 3-11 s o (occasio a y 1- me ium ; I -I 7-1 areas around margin. Dorsal pores of 3 types: (i) s o ; wi 1 - se ae me ia y o me a o a a wi a microductules: sparse, restricted to lines of reticulation; a ge g ou me ia y o meso o a ; wi a g ou o 5-7 (ii) simple pores of 2 types, both rather variable in size: (a) us a e io /mesa o mesoco ae a -5 ea eac darkish pores: common but restricted to reticulation lines; oco a; e g o se ae associa e wi oco ae 1 -1 and (b) slightly larger, paler pores: common along μm; wi 5-11 su ma gi a se ae o eac si e e wee reticulation lines but also present sparsely within s igma ic a eas ese o ge a ose mo e me ia y; reticulation areas and with 9-11 submarginally on each wi 3- ai s o i e -a e a se ae A e ae we side between stigmatic clefts; and (iii) large, very convex, e e o e - ( o - segme e ; 3 segme wi bollard-like macropores, heavily sclerotised, basal portion seu osegme s (w ic some imes a ea o e is i c slightly sunken, with an inner duct: distinctive in side view segme s ; o a e g 319- 3 μm; e g o a ica (Fig. M16) but appearing round in dorsal view: frequent in segme 3 - μm; e g o a ica se a 3 - 5 μm all reticulation lines except near margin; with 5-7 e g o c y eo a a s ie 17 - 7 μm Wi o macropores in posterior medial macropore line. Anal s i acu a e i emes a e io 39- μm os e io -7 a es 1 -1 9 μm o g com i e wi s 1 3-195 μm; μm egs wi a se a a e i ia a a sus; e g s wi 1- mi u e o es o o sa su ace o eac a e; (me a o acic co a 11 -1 μm oc a e + emu e g o se ae i e ma gi 1 (a ou 1/3 om a e 159- 1 μm i ia 135-1 μm a sus 1 -1 7 μm c aw 1 - μm; i e ma gi ( us a e io o a e 1 - 1 - 5 μm 72 d n & nd r n (2000 : C d (In t : pt r : C d Material examined: LECTOTYPE: NEW ZEALAND: However, Maskell's remaining dry material in the ex W.M. Maskell's dry collection of Inglisia inconspicua, second, round pillbox appears to be a mixture of #149 (rectangular box), NZAC #95-018c: 1/1[f] ad. collections, as at least two other coccid species have been PARALECTOTYPES: (i) `Inglisia inconspicua, adult identified within it and the identity of the immatures male, 1890, W.M.M."; NZAC: 1/l [m] ad.; (ii) "Inglisia mounted from it is consequently doubtful. This material inconspicua, larvae, 1891, W.M.M."; NZAC: 1/21 1sts. may not, therefore, be syntypic (see comments in Deitz & Tocker, 1980, p.17). Maskell (1892) indicated that the Other material: NEW ZEALAND:, ex W.M. Maskell's type series was off Corokia cotoneaster (Escalloniaceae) dry collection of Inglisia inconspicua, (round pillbox) (i from the Reefton District of the South Island. [mounted by J.A. de Boer]: 1/1[f] ad; (ii) [mounted by Adult female A. inconspicuus, along with A. RCH] on leaves of Coprosma sp., #95-018a-b, d—f: 6/1[f] chionochloae, A. matai, and A. pubens, have more than 3 ad (+1 [f] adKalasirisandCoccus 1[f] perforata 3rd spinose setae between the lateral stigmatic areas, but A. ?hesperidum), 2[f] 2nd, 1 [m] 2nd, 1 1st, imma u es of inconspicuus can easily be separated from the latter three uncertain species). Mask. coll. #149, USNM: 1/3 1st + 1 species because: e iococci 1s ΤO M ua e u Aristotelia fruticosa [as (i) the marginal setae are broadly spinose (rather than fruiticosa], 10 Mar 1958, J.I. Townsend: 3/3 [ff] ad. NN: finely spinose to setose); Nelson, Plagianthus divaricatus, 22 Nov 1972, J.S. (ii) there are no hypopygial setae (present in the other ug a e 9 Β 1/1[ ] a e e a o sum s i C species); Wai a a Muehlenbeckia australis, 21 Nov 1915, G. (iii) the pregenital disc-pores are not restricted to the Brittin Coll., Let. [sic!] T.D.A. Cockerell, Jan 1916, submedian folds on the abdomen but are also found USNM: 1/1[f] ad [as Eriochiton spinosus]. MC: Chaney's medially. Corner, [no host], Feb 1915, G. Brittin #76: l/1[f] ad. New Brighton, Corokia sp., 25 May 1915, G. Brittin, #62: 1/1 [f] Biology. Overwinters as the adult female. The male tests ad. New Brighton, [no host], 26 May 1915, G. Brittin, and nymphs are found on the leaves in the spring #76: 1/1[f] ad. Birdlings Flat, on stems of Coprosma (September) and the adult females on twigs of the host propinqua, 25 Aug 1996, G. Hall, #96-156a-i 9/9[ff] ad. plants. Birdlings Flat, property G. Taylor, Coprosma propinqua, Distribution. Currently known from the middle of the stems and leaves, 26 Sept 1997, RCH, #97-133a—j: 10/ North Island to the south of the South Island (Map 4). 6[ff] ad, 2[ff] 3rd, f] 2nd, 3[mm] 2nd, 4 pupae. Banks Peninsula, Hinewai Reserve, Pseudowintera colorata stem, 6 Apr 1999, R. Macfarlane, #99-017a-b: 2/1[f] ad, 9 1sts. SL: Tiwai Point S22/35/78, Olearia nummularifolia, Aph n h t n h e e so & o gso ew 16 Sept 1974, G. Collett, FRNZ: R(b)44: 4/4[ff] ad. s ecies Remarks. The available syntype series of 2 original slides igs Μ 3 C33 1 contains an adult male and numerous crawlers, all in poor Unmounted material: Adult females almost round in condition. It is here considered that none of these shape, very thin and nearly transparent, often only noticed specimens is really suitable for primary type designation on undersurface of leaves because of more yellowish and so the lectotype has been selected from Maskell's dry unborn nymphs clustered in abdomen; female test of thin collection of Inglisia inconspicua. This dry material is glassy plates, marginal fringe plates rather pointed, but stored in two boxes: the first, a small rectangular box pair in each stigmatic cleft curve in towards each other. (labelled #149), contains twigs that each have a pin-hole through the stem, and several triangular pieces of card with Mounted material: body broadly elongate-oval; with "149" written on them, also each with a pin-hole. It is clear distinct, moderately deep, stigmatic clefts; anal cleft about that at one time the specimens in this box were held on 1/6th body length; length 1.4-3.7 mm; breadth 1.2-3.1 labelled pins, indicating that they were part of Maskell's mm. original material. An adult female which was found glued to one of these twigs in the rectangular box and which Dorsum: dorsal pores in a reticulate pattern, with 7 appears to fit Maskell's description of I. inconspicua, has rows of reticulation areas across dorsum, 7 or 8 been slide-mounted (#95-018c) and designated as the reticulation areas between anal plates and anterior margin lectotype. The above description is based on this specimen and 29 areas around margin. Dorsal pores of 3 types: (i) and on subsequent material from other collectors. microductules: frequent within lines of reticulation; (ii) n f l nd 4 73 slightly larger simple pores: frequent along margins of all PARATYPES: as for holotype [f], NZAC #93-284b: 1/1[f] e icu a io i es; a (iii a ge mac o o es ( ig Μ 3 ad, 1[m] 2nd, pupa. -11 μm wi e (a eas wi o ege i a isc- o es slightly convex, with a sclerotised rim and a granulate Other material: NEW ZEALAND: ND: Waipoua, surface: infrequent in median reticulation lines and scarce Weinmannia silvicola [as sylvicola], 6 Nov 1961, J.M. or absent on head; with 2-3 in posterior medial macropore Cox, FRNZ R(a) 73: 1/1[f] ad. AK: Hunua Ra, i e A a a es 13 -171 μm o g com i e wi s Mangatangi Reservoir, Workman Tk, Weinmannia 131-191 μm; eac wi 1-3 mi u e o es o o sa silvicola leaf, 31 Jan 1998, C.J. Hodgson, #98-082: l/1[f] su ace; e g o se ae i e ma gi 5-7 μm; i e ad. BP: Rereauira Swamp, Beech Ridge, Weinmannia ma gi 9-11 μm; a ica 11-17 μm; ou e ma gi 9-17 racemosa, 17 Sep 1992, RCH, #92-310a-b: 2/5[ff] ad. As μm; a ea a e o a a a es A oge i a o wi previous except 29 Jan 1993, RCH, #93-063 & #93-278a- pairs of setae along anterior margin and with a single pair C: 4/5[ff]9 ad, 3[f] 2nd. Te Koau, 200 m, Bushwalk Tk, a e a y; o ges 3 -5 μm Weinmannia racemosa, 31 Oct 1994, RCH, #94-106a-d: Margin: marginal setae finely spinose, with 3 on each 4/2[ff] ad, 4[f] 3rd (1 pharate), 1[f] 2nd, 7 1sts, l prepupa. si e e wee s igma ic c e s e g 9- μm ose us Mamaku, Aquarius Rd, Weinmannia racemosa, 22 Sep posterior to each cleft slightly larger than elsewhere. 19 1 M McΚe ie 73 /1[ ] a o o ua 19 Stigmatic spines: of rather uniform thickness, often rather a 199 Weinmannia racemosa, C.J. Hodgson, #98- e wi a u a e ; e g 5 -99 μm 042: 1[m]ad.Weinmannia/1 GB: Paoneone, racemosa Venter: pregenital disc-pores restricted to a group of leaves, 2 Nov 1994, RCH, #94-107a-b: 2/2[ff] ad, 1 [m] ad, 6-16 on either side of anterior end of anal cleft on segment 1 [m] 2nd, 1 prepupa. TK: Awakau Rd,Weinmannia VII. Spiracular disc-pores: in bands mainly 1 pore wide; racemosa leaves, 12 Dec 1993, RCH, #93-373a-i: 9/ with 15-33 in each anterior band and 24-47 in each 14[ff] ad, 2[ff] 3rd, 1[f] 2nd, 4[m] 2nd, pupa, 14 1st. Awakau posterior band. Ventral microducts as for genus. Ventral Rd, Weinmannia racemosa underside leaves, 13 Nov se ae e a a a o e se ae u a s o 5- 1 μm 1994, RCH, #94-124a-c: 3/3[ff] ad, 2[f] 3rd, 4[m] 2nd, 10 long; with 4-6 pairs of anterior anal cleft setae pointing in 1 st. As previous except Melicytus ramiflorus [tree beside several directions but more or less in a line; hypopygial W. racemosa], RCH & FL Henderson, #94-115c, 1/4[m] setae absent; with 2-3 long setae present medially on 2nd. TO: Ohakune, Weinmannia racemosa, 23 Sept 1958, segment VII and 5-10 distinctively long setae on segment G.B. Rawlings & R. Zondag, FRNZ R48: 1/3[ff] ad. VI, these all generally nearly as long as those on segment Pokaka, Weinmannia racemosa, 27 Jan 1982, C.F. VII; other setae almost all short; number of short setae Butcher, #83-325f: 3/3[ff] ad. FD: Chalky Inlet, medially on each abdominal segment: VII, 5-10; VI, l-4; Edwardson Sound, L. Cadman Island, Weinmannia V, 7-11; IV, 5-11; III, 8-12; and II, 8-12; with 9-20 setae racemosa, 12 Feb 1996, RCH, #96-088: l/2[ff] ad. Dusky across metathorax, 2-11 medially on mesothorax (none Sound, Resolution I, Facile Harbour, Weinmannia just anterior to coxae) and 0-2 near each procoxa, all short, racemosa, 5 Feb 1996, RCH, #97-075a-c: 3/1[m] ad, 2[m] e g o se ae associa e wi eac oco a 5- μm; wi 2nd, pupa; moulted skin. SI: Stewart I, Halfmoon Bay, 2-4 pairs of inter-antennal setae; with 5-7 submarginal Weinmannia racemosa leaves, 18 Feb 1996, RCH, #96- setae on each side between stigmatic areas. Antennae 6- 089a-b: 2/5[ff] ad. segmented, with 0-2 pseudosegments in segment III; total e g -37 μm; e g o a ica segme 3 - μm; e g o a ica se a 3-5 μm Mou a s usua y Remarks. A. kamahi is one of the Aphenochiton species displaced towards 1 procoxa; length of clypeolabral shield which have large, rather flat, dorsal macropores (along 11 -135 μm o g Wi o s i acu a e i emes with A. dierama and A. grammicus). It differs from the a e io 1- 9 μm os e io 5-3 μm egs i io- other two species in the following combination of a sa se a a io usua y is i c u a e y e uce a characters: appearing fused on an individual leg; lengths (metatho- (i) dorsal macropores rather few, restricted to median acic co a 111-153 μm oc a e + emu 151-1 5 reticulation lines; μm i ia 1 -13 μm a sus 91-1 1 μm c aw 11-15 μm (ii) very long pregenital setae present on both segment VI hooked part of claw very short. and VII; (iii) 29 reticulation areas around the margin; Material examined: HOLOTYPE [f]: NEW ZEALAND: (iv) marginal setae just posterior to each stigmatic cleft GB: Kakanui, 300 m, 30 Apr 1993, R.C. Henderson, only slightly longer and more spinose than those Weinmannia racemosa, NZAC #93-284a: 1/1[f] ad. elsewhere. 7 d n & nd r n (2000 : C d (In t : pt r : C d Biology. It seems likely that this species has one 1 -153 μm; eac wi -1 mi u e o es o o sa generation per year overwintering as adult females. The su ace; e g o se ae i e ma gi 5- μm; i e adult female is probably the most thin and round coccid (as ma gi 7-9 μm; a ica 9-13 μm; ou e ma gi 1 -15 opposed to thin and long with A. r in New μm A oge i a o usua y wi ai s o se ae a o g Zealand, appearing as nothing more than a film on the a e io ma gi a 1 ai a e a y; o ges 37- 7 μm undersurface of the leaf and is nearly impossible to Margin: marginal setae finely spinose, with 3-7 on each distinguish if wet with rain. Neonate nymphs have been side between stigmatic clefts; setae of 2 sizes: most setae observed emerging one at a time from beneath the sma a i e 7-1 μm o g u ose us os e io o posterior end of the test; there does not appear to be a s igma ic c e s o ge a mo e s i ose 1 - μm o g; brood chamber as on females of Ct n h t n or K l r reticulation setae clearly enlarged when setae frequent but spp. Apparently restricted to W n nn spp. this less obvious when setae infrequent. Stigmatic spines of a rather uniform thickness, sometimes slightly bent, and Pathogens and parasitoids. Hymenopterous parasite wi a u a e ; e g 1-7 μm recorded: Pteromalidae: Aph b t nana ( ouček Venter: pregenital disc-pores with mainly 5-6 loculi Distribution. Throughout, from the far north of the North (range 3-8), all associated with mediolateral folds on Island to Stewart Island in the far south (Map 5). abdomen; number on each side of segment: VII, 5-8; VI, 2-4; V, 1-3; IV, 1-3; III,1l-2; and II, 0-1; none laterad to Name derivation. As this species appears to be restricted metacoxae. Spiracular disc-pores: with 12-18 in each to species of W n nn , it is named after the host plant's anterior band and 16-22 in each posterior band; bands Maori name h . narrow but broadening slightly near spiracle and margin and with 0-2 pores present a short distance mesad to peritremes. Ventral microducts: frequent, distributed as for genus. Ventral setae: anal lobe setae sometimes Aph n h t n t e e so & o gso ew a se w e ese s o 7-3 μm o g; wi - pairs of anterior anal cleft setae; with a group of 8-10 s ecies hypopygial setae, often in 2 longitudinal lines and of rather variable length; with a pair of long pregenital setae Fig. 101 restricted to segment VII; number of medium/short setae Unmounted material: unknown. medially on each abdominal segment: VII, 0-4 medium + 2-8 short (total 6-10); VI, 6-10 short; V, 9-16; IV, 12- Mounted material: body elongate-oval, with shallow 15; III, 8-11; and II, 10-12; with 3-4 setae associated with stigmatic and anal clefts, latter about 1/9th body length; each meta- and mesocoxa plus 2-5 medially, and 1-3 near length 1.5-1.8 mm; breadth 0.85-0.93 mm. each procoxa, all rather short, length of setae associated wi eac oco a 9-13 μm o g; wi -3 ai s o i e - Dorsum: dorsal pores in a reticulate pattern, with 7 a e a se ae; wi 3-7 su ma gi a se ae o eac si e rows of reticulation areas across dorsum, with probably 9 between stigmatic areas. Antennae 6- or 7-segmented, reticulation areas between anal plates and anterior margin with 0-2 pseudosegments in 3rd segment; total length and 29 areas around margin. Dorsal pores of 4 types: (i) 75-31 μm; e g o a ica segme 3 - μm; e g microductules: frequent, associated with lines of o a ica se a 3-7 μm Mou a s occasio a y reticulation and also in a band along margins of anal cleft, displaced to one side; length of clypeolabral shield 126- where slightly larger; (ii) flattish, simple pores, slightly 135 μm Wi o s i acu a e i emes a e io -3 larger than microductule: frequent in all reticulation lines μm os e io 3 - μm egs wi a se a a e i ia a only; (iii) larger, somewhat convex pores, rather scarce, a sus; e g s (me a o acic co a 1 -117 μm located near reticulation lines but absent from submargin; oc a e + emu 131-1 μm i ia 93-119 μm a sus a (i e a i e y sma mac o o es (3-5 μm iam 75-9 μm c aw 1 - μm su equa i wi o ege i a isc- o es ea i y sc e o ise a co e mos a u a i me ia a Material examined: HOLOTYPE [f]: NEW ZEALAND: su me ia e icu a io i es a se ea ma gi a WO: SF 97, Waimiha, (Te Kuiti) , 1 Oct 1957, R.C. ecomi g ess eque o o a a ea ; wi 5- Howie, d rp p t [= r n p t t x f l ], macropores in posterior medial macropore line. Anal FRNZ, R32: l/4 ad, 2[mm] ad, 2 pupae, holotype a es a e a ow 15 -17 μm o g com i e wi s clearly marked. n f l nd 4 75 PARATYPES: on same slide as holotype: 1/4[ff] ad mac o o e i e A a a es 13 - 3 μm o g com i e (holotype [f] plus 3 [ff] paratypes), 2[mm] ad, 2 pupae. wi s 1 - 1 μm; wi 1- mi u e o es o o sa su ace o eac a e; a a a e se ae i e y s i ose a Other material: WO: as for holotype, except FRNZ R34: s o ; e g o se ae i e ma gi 1 5-1 μm; i e 1/2[mm] ad (poor condition). ma gi 9-1 μm; su a ica 1 - μm; ou e ma gi se ae 1 - μm A oge i a o wi ( a e y 3 ai s o Remarks. A. t belongs to the group of species in se ae a o g a e io ma gi a a o e ai o a e a Aph n h t n which possess hypopygial setae and small, ma gi s; o ges 3 -5 μm o g convex dorsal macropores (i.e., A. h n hl , A. matai, Margin: marginal setae finely spinose, with 3 on each and A. p b n . si e e wee s igma ic c e s; mos se ae 7-9 μm o g A. t shares with A. h n hl the presence of ose us os e io o eac s igma ic c e s ig y a ge pregenital disc-pores mediolaterally on all/most abdomi- 1 - μm o g S igma ic c e s is i c a a a e — nal segments, but differs from it in being much smaller. si e ; s igma ic s i es s o s ou a u some imes A. ιn t differs from A. p b n in having more s ig y cu e wi a we - e e o e oa asa abundant pregenital disc-pores which extend onto the socke ; e g 55-99 μm more anterior abdominal segments (restricted to laterad to Venter: pregenital disc-pores restricted to a group of anogenital fold on A. p b n . 2-13 on either side of anogenital fold. Spiracular disc- pores: with 19-39 in each anterior band and 19-43 in each Biology. Unknown. posterior band; anterior bands with 0-2 pores present just mesad to peritremes. Ventral microducts as for genus. Distribution. A. matai is only known from this one Ventral setae: ventral anal lobe setae rather short, 18-32 collection made from e Kuiti in 1957 (Map 6). μm o g; wi a i e o -7 mo e a e y o g a e io a a c e se ae ese o a ea i g s i a o a oi i g Name derivation. This species has been named after the i wa y owa s a a c e u o e yi g c ose o a a Maori name t for the host plant r n p t c e ma gi ; y o ygia se ae a se ; wi a ai o o g t x f l , from which it was collected. ege i a se ae ese me ia y o segme VII, but with medium length setae on segments VI, V, and sometimes IV; number of setae medially on each abdominal segment: Aph n h t n pr n e e so & o gso ew VII, 2 long + 6-12 short (total 8-15); VI, 1-13 medium s ecies length + 0-13 short (total 4-20); V, 1-7 medium + 1-12 short (total 5-15); IV, 0-2 medium + 3-14 short (total 4- Figs C36, 102 14); III, 4-14 short; and II, 3-17 short; with 10-22 setae Unmounted material: fresh material of adult females seen across metathoracic segment, 2-4 near mesocoxa and 1-3 τom b p r were yellow-green, flat above, (rather spinose) near procoxa; length of setae associated more convex ventrally. wi eac oco a a ou 9 μm o g; wi mai y 3- (occasio a y ai s o i e -a e a se ae; wi 3- Mounted material: body oval, with shallow but distinct su ma gi a se ae o eac si e e wee s igma ic a eas; stigmatic clefts and shallow anal cleft, latter about 1/9th wi a ew sma se ae is i u e a e a om y i a body length; length 1.8-4.9 mm; breadth 1.4-3.5 mm. oa su ma gi a a Wi o eac s i acu a e i eme a e io 7- 3 μm os e io 3 -5 μm Dorsum: dorsal pores in a reticulate pattern, with 7 A e ae -segme e some imes wi a seu osegme rows or reticulation areas across dorsum, 7 or 8 (o e o g a e a 7-segme e ; i segme a ou reticulation areas between anal plates and anterior margin su equa o o e i e segme s oge e ; o a e g 311- and perhaps 29 areas around margin. Dorsal pores of 3 1 μm; e g o a ica segme 3 - μm; e g o types: (i) microductules: most frequent associated with a ica se a 3 -5 μm e g o c y eo a a s ie 1 - lines of reticulation; (ii) simple pores, slightly larger than 175 μm o g egs wi a se a a e i ia a a sus; microductules: frequent along margins of all reticulation e g s (me a o acic co a 1 - 3 μm oc a e + lines; some dark and some paler; and (iii) larger, convex emu 1 - 9 μm i ia 159- 13 μm a sus 1 -135 mac o o es 7- μm iam (su equa i si e o μm c aw 19- 5 μm ege i a isc- o es ; some imes wi a o g i a k i e g a sys em wi i su me ia a me ia Material examined: HOLOTYPE [f]: NEW ZEALAND: e icu a io i es; wi - mac o o es i os e io me ia CO: Rock and Pillar Range, Summit Stonehenge, 2 Dec 7 d n & nd r n (2000 : C d (In t : pt r : C d 1993, B.H. Patrick, Hebe pauciramosa leaves, NZAC Distribution. Four out of the five lots of material were #93-375h: 1/1 ad. collected in the South Island and the fifth near Auckland PARATYPES: as for holotype : NZAC #93-375a—g, i—k: (Map 7). 10/18 [ff] ad, []ad, 15 1sts. Name derivation. The specific name is taken from the Other material: NEW ZEALAND: AK: Huia, Coprosma Latin pronus meaning lying flat or face downwards on the sp. leaves, 13 Jan 1983, J.M. Cox, #138: 5/4[ff] ad, 1 ground. 3rd. WD: Otira, Coprosma sp., Christmas 1915, G. Brittin, #97: 1/1[f] ad. SL: Southland, Blue Mountains, 1020 m, pitfall trap in tussock, 5 Jan 1985, B.I.P. Barratt: Aph n hlt n p b n e e so & o gso ew 6/7[ff] ad. s ecies igs Μ 5 C3 1 3 Remarks. This material is rather variable in size, that from Otira being large (particularly the leg measurements) Unmounted material: nearly indistinguishable from A. and one specimen from the Blue Mountains appearing to subtilis; both species nearly transparent, but adult female be a runt. All of the Blue Mountain specimens are small A. pubens more yellow-green than those of A. subtilis, compared with other specimens but, as they were caught in which are rather blue-green. Test of A. pubens with 9 wax pitfall traps, perhaps they were dispersing pre- plates between anterior margin and anal plates. Body reproductive females. shape usually long and very flat; detected on plant by A. pronus is similar to A. subtilis but the latter has: reflective shine of wax test. (i) fused tibia and tarsus (separate on A. pronus); (ii) anterior anal cleft setae long, in a line facing inwards Mounted material: body elongate-oval; with shallow but and appearing rather stiff, like a comb (not as long or distinct stigmatic clefts; anal cleft quite deep, about 1/6th as formally placed in A. pronus); body length; length 2.0-5.7 mm; breadth 1.1-3.2 mm. (iii) an unusually long apical segment to the antenna (>50 Dorsum: dorsal pores in a reticulate pattern, with 7 μm o A subtilis, μm o A pronus). rows of reticulation areas across dorsum, 9 reticulation A. pronus also resembles A. kamahi in: areas between anal plates and anterior margin, and 29 (i) having few dorsal macropores restricted to median and areas around margin. Dorsal pores of 3 types: (i) submedian reticulation lines; microductules: frequent, associated with lines of (ii) absence of hypopygial setae; reticulation and also frequently abundant in a band along (iii) pregenital disc-pores few and restricted to laterad to margins of anal cleft, particularly just posterior to anal anogenital folds. plates, when slightly larger; (ii) flattish, simple pores of 2 A. kamahi differs in having: sizes: small pores, subequal to that of microductule: (i) much larger dorsal macropores (about twice the width frequent along margins of all reticulation lines; and much of the pregenital disc-pores), which are more or less larger pores: rather scarce but also near reticulation lines; restricted to the median reticulation lines and absent no submarginal band of dorsal pores present; and (iii) from the head (more abundant and widespread on A. e a i e y sma " um-s a e " mac o o es ( ig Μ 5 pronus); (su equa i wi o ege i a isc- o es ea i y (ii) muc sma e egs ( oc a e + emu 151-1 5 μm as sc e o ise a ig y co e mos a u a i me ia com a e wi 1 - 9 μm o A pronus). a su me ia e icu a io i es ecomi g ess eque ea ma gi a a e io y; wi - mac o o es i Biology. The mature adult females collected off Hebe os e io me ia mac o o e i e A a a es 135- 5 pauciramosa were not entirely sessile because they were μm o g com i e wi s 133-1 μm; eac wi 5-11 able to walk away from the preferred host site in the natural mi u e o es o o sa su ace; e g o se ae i e depression in a leaf's upper surface when disturbed. It ma gi 1 7-11 μm; i e ma gi 9-15 μm; a ica 1 - seems possible that the long legs of this species are an μm; ou e ma gi (ac ua y i isc a osi io 1 - 7 adaptation for moving to new host sites, perhaps after μm A oge i a o wi 3-5 ai s o se ae a o g a e io overwintering as adult females on older leaves. All known ma gi a 1 ai a e a y; o ges 3 -7 μm collections of A. pronus have been taken in midsummer, so Margin: marginal setae finely spinose, with 7-13 on it is probably univoltine; those off H. pauciramosa in eac si e e wee s igma ic c e s e g 9- 9 μm a ges December had begun to deposit nymphs. us os e io o s igma ic s i es; e icu a io se ae a so n f l nd 4 sometimes slightly enlarged. Stigmatic spines of rather 028i—k: 3/1[f] ad (pharate), 6[f] 3rd. As previous, except 7 uniform thickness, often rather bent with a blunt apex; July 1998, #98-080a—e: 5/3[ff] ad, 11 crawlers (from e g 5-9 μm brood chamber). As previous except 19 July 1998, #98- Venter: pregenital disc-pores with mainly 6 loculi: 081: 1/1[f] ad. CL: Little Barrier I, Valley Tk ridge, Mida restricted to a group of 5-21 on either side of anterior end salicifolianew & old leaves, 17 Sept 1994, RCH, #94- of anal cleft in segment VII; when frequent, group 071c: 1/1[f] ad. BP: Lottin Point, Otanga, Litsea calicaris, extending a short distance down margins of anal cleft. 27 Jan 1993, RCH, #93-272: 1/1[f] ad, 1[f] 2nd (pharate), 4 Spiracular disc-pores: with 22-47 in each anterior band 1 st. Lottin Point, Otanga, Hedycarya arborea old leaves, and 29-58 in each posterior band; anterior bands with 0- 29 Sept 1993, RCH, #93-328b: 1/2[ff] ad. Te Koau, 130 3 pores just mesad to peritremes. Ventral microducts as m, bushwalk to Twin Puriris, Hedycarya arborea, 2 Nov for genus. Ventral setae: ventral anal lobe setae unusually 1993, RCH, #93-347: 3/1 [f] ad (+ [f] adAphenochiton o g 7 -1 μm; wi - ai s o a e io a a c e subtilis), 1 [m]2nd, pupa, 2 [mm] ad. GB: East Cape, setae; with a group of 5-16 hypopygial setae; with long Lighthouse Tk, Pittosporum tenuifolium leaves, 1 Nov pregenital setae restricted to segment VII; number of setae 1994, RCH, #94-105: 3/1[f] ad, 1 [m]2nd (pharate), 10 1sts. medially on abdominal segments: VII, 2-6 long + 0-6 As previous, except 3 Nov 1995, underside leaf, #95-107: short; VI, 7-14 short; V, 8-17; IV, 8-21; III 8-16; and II, 1/1 [f] ad. WO: Mangapohue Bridge,Beilschmiedia tawa, 11-16; with 8-17 setae medially on metathorax, 9-16 ec 1993 C 9 - 3 1/1[ ] a 1 1s Ο medially on mesothorax and 2-5 near each procoxa, all a gi o o S a io Ma ga u u Beilschmiedia tawa rather short; length of setae associated with each procoxa underside leaves, 9 Nov 1996, RCH, #96-209: 1/1[f] ad. 9-13 μm o g; wi -5 (usua y 3- ai s o i e - As previous, #96-210: 2/1 [m]2nd, 1 [f] 3rd. TK: Mt Egmont, a e a se ae; wi - su ma gi a se ae o eac si e 800 m, Griselinia lucida leaves, 24 Feb 1983, C.F. between stigmatic areas. Antennae 6- to 8-segmented, Butcher, #83-067a: 2/4 [ff] ad. NN: Riwaka, [no host], 2 with 0-2 pseudosegments in 3rd segment when 6- or 7- Nov 1924, Brittin, #90-215: 4/5 [f]f ad. segme e ; o a e g 3 9- 59 μm; e g o a ica segme 3 -5 μm; e g o a ica se a 59- 5 μm Remarks. A. pubens, although rather variable, is a quite Mouthparts often displaced to one side on larger distinctive species. It shares with A. chionochloae and A. s ecime s; e g o c y eo a a s ie 139-1 μm matai: o g Wi o s i acu a e i emes a e io 31-51 μm (i) the presence of hypopygial setae (absent on other os e io 39-5 μm egs wi a se a a e i ia a a sus; species in Aphenochiton); e g s (me a o acic co a 1 5-19 μm oc a e + (ii) (usually) more than 4 marginal setae on each side emu 19 - 5 μm i ia 1 9-1 9 μm a sus 99-1 μm between the stigmatic clefts (3 on other Aphenochiton c aw 1 - μm species, except A. inconspicuus). It differs from A. chionochloae and A. matai in having Material examined: HOLOTYPE [f]: NEW ZEALAND: the pregenital disc-pores restricted to segment VII, AK: Waitakere Range, Sharp Bush, 22 Mar 1998, R.C. whereas they extend onto the more anterior segments on Henderson, Mida salicifolia large-leaved sapling, NZAC the other two species. #98-043d: l/1[f] ad. A. pubens is very similar to A. subtilis in general PARATYPES: as for holotype: #98-043a—c, e-f: 5/4[ff] appearance and the two species have been found together ad, l[m] 2nd. on the same Mida salicifolia leaf! (but the presence of hypopygial setae in A. pubens and the fused tibia+tarsus in Other material: NEW ZEALAND: ND: Kamo,Podocarpus A. subtilis quickly separates these two species). totara, 9 Oct 1968, R.A. Cumber, #1643: 1/2[ff] ad. ND/ One specimen off Mida from Little Barrier Island had AK: Kaiwaka, Podocarpus totara, 18 Sept 1968, R.A. a single tubular duct submarginally between the lateral Cumber, #1620/1643: 1/2[ff] ad. AK: Waitakere Ra, Old stigmatic clefts. Coach Road Tk, Cordyline banksii underside leaves, 18 Oct 1994, RCH, #94-092: 1/1 [f] ad. Riverhead Forest, Biology. Clearly rather polyphagous. It appears to have a Barlow Road reserve, Podocarpus totara leaves, 16 Aug similar life history to that of A. subtilis, overwintering as 1996, RCH, #96-152: 1/1[f] ad. Waitakere Ra, Sharp Bush, the adult female and probably with one generation a year. 6 Feb 1998, RCH, Mida salicifolia both sides lvs, #98- 010a-b: 2/3[f] 3rd, 2[m] 2nd [3[f] 3rd, 4[m] 2nd Distribution. Throughout the North Island and to Nelson Aphenochiton subtilis]. Waitakere Ra, Sharp Bush, Mida in the South Island (Map 8). salicifolia large-leaved sapling, 22 Feb 1998, RCH, #98- 7 d n & nd r n (2000 : C d (In t : pt r : C d Name derivation. The name pubens (L. = pubic hair) disc-pores in bands mainly 1 pore wide; with 15-47 in refers to the presence of the hypopygial setae between the each anterior band and 21-57 in each posterior band; vulva and the anogenital fold. occasionally with 1-2 disc-pores mesad to peritremes. Ventral microducts as for genus. Ventral setae: ventral a a o e se ae se ose 3 - 7 μm o g; wi 5-9 ai s o Aph n h t n bt l e e so & o gso ew anterior anal cleft setae, generally forming a distinct line, s ecies setae quite long and pointing inwards towards anal cleft; hypopygial setae absent; with 2-3 long pregenital setae Figs C35, 104 present medially on segment VII and 4-10 moderately Unmounted material: nearly indistinguishable from A. long setae on segment VI (about 1/2 length of those on pubens; both species nearly transparent, but adult female segment VII); other setae mostly short; number of short A. subtilis rather blue-green, those of A. pubens more setae medially on abdominal segments: VII, 6-13; VI, l- yellow-green. Test of A. subtilis with 8 wax plates 3; V, 5-12; IV, 5-1l; III, 7-11 and II, 11-20; with 12-21 between anterior margin and anal plates; paired marginal setae medially on metathorax (one specimen had a single wax plates in area of each stigmatic cleft tending to be seta anterior to metacoxa), 11-23 medially on mesothorax unusually narrow, lying parallel with margin rather than (none just anterior to coxae) and 0-2 near each procoxa, all curving outwards. Body usually long and very flat, short, length of setae associated with each procoxa very detected on plant by reflective shine of wax test. s o 5-7 μm o g; wi 3-5 ai s o i e -a e a se ae; with 3-9 submarginal setae on each side between stigmatic Mounted material: body broadly elongate-oval; with clefts. Antennae 6-segmented, with 0-2 pseudosegments distinct, moderately deep stigmatic clefts; anal cleft about i o g segme III; o a e g 3 7-5 μm; a ica 1/7th-1/8th body length; length 1.90-4.35 mm; breadth segme a icu a y o g (5 -57 μm ; e g o eac 1.37-3.33 mm. a ica se a 5-77 μm Mou a s usua y is ace o o e si e; e g o c y eo a a s ie 1 -1 μm o g Dorsum: dorsal pores in a reticulate pattern, with 7 Wi o s i acu a e i emes a e io 7-3 μm rows of reticulation areas across dorsum, 8 reticulation os e io 9-3 μm egs i ia a a sus use u areas between anal plates and anterior margin and 27 or 29 separation rarely faintly indicated; lengths (metathoracic): areas around margin. Dorsal pores of 3 types: (i) very co a 1 1-1 μm oc a e + emu 1 - μm i ia small, dark microductules: frequent within lines of + a sus - μm c aw 13-1 μm ooke a e y reticulation; (ii) flat, simple pores, at least 2 x size of short. microductules, some at least 3 x larger: frequent along margins of all reticulation lines; and (iii) fairly small Material examined: HOLOTYPE [f]: NEW ZEALAND: mac o o es ( -5 μm wi e su equa o si e o ege i a AK: Waitakere Range, Sharp Bush, 22 Mar 1998, R.C. isc- o es sc e o ise a co e a e ew mai y i Henderson, Mida salicifolia large-leaved sapling, NZAC me ia e icu a io i es; sca ce o a se o ea ; #98-044a: 1/1 [f] ad. macropores absent in posterior medial macropore line. PARATYPES: collection data as for holotype: (i) NZAC A a a es 1 -173 μm o g com i e wi s 1 -1 #98-044b—h: 7/7[ff] ad (one pharate); and (ii) NZAC #98- μm; eac wi -7 mi u e o es o o sa su ace; e g 045a—b: 2/2[ff] ad. o se ae i e ma gi 1 5-7 μm; i e ma gi 9-1 μm; a ica 1 -1 μm; ou e ma gi 1 -13 μm; a ea a a O e ma e ia EW EA A ΤΗ ee Ki gs Is a e a e A oge i a o wi ai s o se ae a o g West I, Meryta sinclairii leaves, 29 Nov 1983, C.F. a e io ma gi a a si g e ai a e a y; o ges 39-5 Butcher, #83-347b: 1/1[f] ad. As previous, but on leaves of μm Elingamita sp. (or Corynocarpus laevigatus?), #83-347e: Margin: marginal setae finely spinose, with 3 on each 4/3[ff] ad, 1[f] 3rd. AK: Waitakere Ra, Walker Bush Tk, si e e wee s igma ic c e s e g 9- 3 μm ose us Mida sp. leaves, 5 Nov 1976, M.F. Tocker, #76-316a: 1/ os e io o eac c e s ig y a ge a e sew e e 1[f]ad.Midaboth Waitakere salicifolia Ra, Sharp Bush, S igma ic s i es o a e u i o m ick ess o e sides leaves, 1 Dec 1994, RCH, #94-123a—c: 3/5[f] 2nd, 3[mm] somew a e wi a a e u a e ; e g 5 - 5 μm 2nd, 16 1st. As previous except 29 Apr 1995, #95-042a: Venter: pregenital disc-pores generally slightly oval, 1/2[ff] ad. As previous except 13 July 1997, #97-081a—b: with 5-9 loculi (generally 7-8); with 9-17 on each side of 2/2[ff] ad, 1 [m]2nd. As previous except 6 Feb 1998, #98- anterior end of anal cleft on segment VII (one specimen 110a-b: 2/3[f] 3rd, 4[m] 2nd [+3[f] 3rd, 2[m] 2nd A. pubens]. had a single disc—pore laterad on segment VI). Spiracular As previous except 22 Feb 1998, #98-029a—b& 98-030a, n f l nd 4 79 c-d: 5/9[f] 3rd, 9[m] 2nd. As previous except 6 Mar 1998, most favoured host plants are M d l f l and #98-041a: 1/1[m] 2nd. As previous, except 22 Mar 1998, d r rb r . #98-046a—e: 5/5[ff] ad, 2[f] 3rd, 5[mm] 2nd. As previous, except 7 July 1998, #98-079a—b: 2/2[ff] ad. CL: Little Distribution. Rather anomalous, with most records from Barrier I, Valley Tk, ridge, M d l f l new and old the North Island but also from Fiordland and Invercargill leaves, 17 Sept 1994, RCH, #94-071a-h: 8/6[ff] ad, 2/5[f] (Map 9). 3rd, 2[f] 2nd, 2[mm] 2nd. As previous except Hamilton Tk, #94-081 a-i: 9/7[ff] ad, 3[ff] 3rd, 1[f] 2nd, 1 [m]2nd, ?5 1sts. Name derivation. From the Latin bt l , meaning thin, BP: Te Koau, d r rb r , 23 Sept 1992, RCH, referring to the very thin and flat nature of this species, #92-278: 1/1 [f] ad. Te Koau, 130 m, Bushwalk to Twin which resembles a film of water on the underside of a leaf. Puriris, d r rb r , 2 Nov 1993, RCH, #93- 347a—b: 2/2[ff] ad (+ [f] adGB:A. Paoneone,p b n . d r rb r both sides lvs, 2 Nov 1994, RCH, #94-127a-b: 3/3[ff] ad, 12 1sts. TO: Rangitoto Stn, Mangatutu, M d l f l leaves, 9 Nov 1996, RCH, #96-211a--f: 5/3[ff] ad, 3[ff] 3rd, 5[mm]2nd. WN: Rimutaka Ge us C YS A O ES A e e so & Ra, Mt Mathews, 300 m, d r rb r , 25 Dec o gso ew ge us 1932, D. Zotov: 1/1[f] ad. FD: Doubtful Sound, Bauza 1, Type species: In l f Maskell (here designated). [as d p n x] pl x leaves, 28 Jan 1996, RCH, #96-043: 1/1 [f] ad. Breaksea Sound, Breaksea I, Diagnosis. Adult female. Test: conical, elongate-convex, d r rb r underside of leaves, 29 Jan 1996, open beneath, with a distinct fringe of small plates that are RCH, #96-072: 1/1[f] ad, 2 lsts. SL: Invercargill, often sharply triangular; texture glassy. Thomsons [as Thompsons] Bush, tt p r sp. leaves, Body: elongate oval to rather broad; anal cleft 27 Jan 1983, C.F. Butcher, #84-024e: 2/2[ff] ad, 1[f] 2nd, shallow; stigmatic clefts absent or distinct but shallow. 1[m] 2nd. Dorsum: derm membranous. Dorsal setae absent. Dorsal pores distributed in a reticulate pattern, delineating Remarks. A. bt l can be separated from other species reticulation areas in 5-7 longitudinal rows, 5-8 areas in this genus by the fused tibia and tarsus. It most closely between anal plates and anterior margin (middle resembles A. pr n and A. h , the latter also being reticulation area sometimes particularly large), and with particularly thin and flat in life, but differs from them in the 26-28 areas around margin on most species but fewer on following combination of characters: Cr t ll t t rn t and C. rn t ll . Dorsal pores of 3 (i) pregenital disc-pores usually with 7 or 8 loculi and or 4 types: (i) minute, dark microductules, with a long often rather oval in shape; inner ductule: mainly associated with lines of reticulation; (ii) the presence of 5-9 pairs of quite long anterior anal (ii) larger, flat, simple pores of variable size: smallest cleft setae in a line facing inwards, often appearing associated with reticulation lines, slightly larger pores stiff and comb-like; often in a submarginal band; large simple pores sometimes (iii) the presence of 4-10 moderately long setae on present laterad to anal plates; (iii) quite large macropores, abdominal segment VI, more or less 1/2 as long as usually more or less round, slightly to distinctly convex, long setae on segment VII (A. h has 5-10 setae generally with a granulate surface: restricted to on segment VI almost as long as long setae on segment reticulation lines; and (iv) in C. rn t and C. rn t ll VII); only: inverted, tubular pores with a sclerotised inner base (iv) long antennae, with a long apical segment of more (small on C. rn t , large on C. rn t ll : restricted to a 5 μm reticulation lines. Anal plates tapering to rather pointed A. bt l is also similar to A. p b n in general apex, with anterior margins much shorter than posterior appearance and the two species have been found together margins; with or without minute pores on dorsal surface of on the same leaf of a plant f M d l f l , but they can each plate; each plate with 4 spinose setae, 2 along inner be easily separated by the presence of hypopygial setae in margin, an apical seta and another on posterior margin. A. p b n . Anogenital fold with 2-3 pairs of setae along anterior margin and 1 pair on each lateral margin. Anal tube quite Biology. Life cycle probably similar to that of A. p b n , long, when inverted extending well anterior to anal plates; with only one generation a year, overwintering as the adult anal ring with 3 pairs setae (4 pairs on C. rn t ll . Anal female. Although it seems to be fairly polyphagous, the plate sclerotisation and supporting bars absent. 80 d n & nd r n (2000 : C d (In t : pt r : C d Margin: marginal setae broadly spinose with narrow crystal, and t t (L. f.) meaning shell. This name was basal sockets; abundant but absent from margins of anal suggested by the late C. Tymone Duval and we here cleft (except on C. n f where a few extend up anal dedicate it to his memory. cleft); reticulation setae may be significantly larger and then often displaced onto dorsum. Stigmatic spines stout, Remarks. This genus contains six species: Cr t ll t t tapering, with a narrow basal socket, 1 per stigmatic cleft; f (Maskell) n. comb., C. f (Maskell) n. comb., C. quite short on some species, very long on others; set l pt p r (Maskell) n. comb., C. n f Henderson & distinctly onto dorsum (significantly so on C. f . Hodgson n. sp., C. rn t (Maskell) n. comb., and C. Eyespot obscure and set slightly onto dorsum, or absent. rn t ll Henderson & Hodgson n. sp. These species are Venter: pregenital disc-pores with mainly 10 loculi; characterised by the presence of: distributed across most or all abdominal segments and (i) numerous pregenital disc-pores (with mainly 10 loculi) sometimes medially on metathorax and laterad to each extending across all abdominal segments: metacoxa. Spiracular disc-pores with mainly 5 loculi, in (ii) 2 types of ventral tubular duct, with the larger duct in bands 1-5 pores wide between each spiracle and margin a submarginal band and the slightly smaller duct (and extending onto dorsum on C. f with 0-2 pores medially on posterior abdominal segments and laterad extending medially past peritreme. Ventral microducts of to anterior end of anal cleft; I type, with a rather broad inner ductule; frequent in a (iii) marginal setae abundant and spinose; broad submarginal band, otherwise rather sparse (iv) reticulation setae often clearly differentiated; throughout but absent from posterior 4 abdominal (v) generally with pairs of long setae on 2 pregenital segments. With 0-1 preantennal pores. Ventral simple segments (3 on C. rn t and C. rn t ll pores present in a submarginal band on C. rn t and C. (vi) generally with 2 or 3 pairs of anterior anogenital fold rn t ll absent on other species. Ventral tubular ducts setae; of 2 types: (i with a long outer ductule, long inner ductule (vii) often with a group of simple pores laterad to anal and a well-developed terminal gland: frequent throughout plates; except medially on more posterior abdominal segments (viii) test tending to be highly convex, stout and glassy. where replaced with (ii) a slightly smaller duct with a Species in the genus Cr t ll t t resemble species broader cup-shaped invagination, with or without a in the genera Ct n h t n and K l r in having abund- glandular end to inner ductule: most frequent on either ant pregenital disc-pores medially across the abdominal side of anal cleft, becoming less frequent on next few segments and numerous spinose marginal setae around the anterior abdominal segments. Ventral setae: ventral anal margin. Cr t ll t t species also resemble K l r lobe setae rather spinose, tapering to a blunt or finely species in having: pointed apex; with 1-5 pairs of anterior anal cleft setae; (i) 2 types of ventral tubular duct (but the distribution on with long pregenital setae present medially on abdominal the two genera differs); segments VII, VI (and V on C. rn t and C. rn t ll (ii) large simple pores on the dorsum, generally associated with few setae medially on each abdominal segment; with the anal plates. usually with 1-3 submarginal setae between stigmatic K l r differs in having ventral tubular ducts: areas; with 3-4 pairs of interantennal setae; other setae (i) in narrow submarginal bands, with the ducts tending to a e ew a sma 5-9 μm o g A e ae we lie radially; developed and 6- to 8-segmented; when 6- or 7- segmented, 3rd segment much longest and generally with (ii) absent medially on the thorax. l-2 pseudosegments. Spiracular peritremes moderate to Ct n h t n species differ from Cr t ll t t species in: quite large. Legs normal to rather small; tibia and tarsus (i) having a thin test; sometimes without an articulation; with 2 long setae on (ii) having very large spiracles; each trochanter; tarsal digitules both slightly longer than (iii) having spinose marginal setae rather unevenly spaced claw digitules, generally subequal in length and with one around margin; narrower; claw digitules both broad and only slightly (iv) having antennae generally without pseudosegments; longer than claw; claw with or without a minute denticle. (v) lacking ventral tubular ducts medially on the thorax; Vulva on segment VII, immediately posterior to long (vi) t he displacement of the mouthparts to one side. pregenital setae. The species currently placed in Cr t ll t t are all endemic to New Zealand. They are possibly univoltine Generic name derivation: the name refers to the apart from C. rn t and C. rn t ll , which may have appearance of the test: r t ll (L.) meaning ice or rock two generations a year. n f l nd 4 8 ig. 0 . Crystallotesta fagi (Maske , . com ., a u ema e Α si e iew o es . 82 d n & nd r n (2000 : C d (In t : pt r : C d ig. 06. Cr t ll t t f (Maske , . com ., a u ema e. n f l nd 4 8 ig. 0 . Cr t ll t t l pt p r (Maske , . com ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . Α ig a o a ium. 84 d n & nd r n (2000 : C d (In t : pt r : C d ig. 08. Cr t ll t t n f e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . Α ig a o a ium. n f l nd 4 8 C ig. 0 . Cr t ll t t rn t (Maske , . com ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . Α ig a o a ium. 86 d n & nd r n (2000 : C d (In t : pt r : C d ig. 0. Cr t ll t t rn t ll e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . Α ig a o a ium. n f l nd 4 7 Key o a u ema e Cr t ll t t Cr t ll t t f (Maske ew com i a io Spiracular disc-pore bands extending past marginal Figs C43, C44, 105 spinose setae onto dorsum; stigmatic spines either Inglisia fagi Maskell, 1890b: xv (nomen nudum). absent or set some distance onto dorsum at end of Inglisia fagi Maskell, 1891: 13; –Maskell, l895a: 13 spiracular disc-pore band fusca [checklist]; –Cockerell, 1896: 330 [checklist]; –Fernald, —Spiracular disc-pores not extending past marginal 1903: 162 [world catalogue]; –Hutton, 1904: 226 spinose setae onto dorsum; stigmatic spines present [checklist]; –Myers, 1922: 199 [checklist]; –Miller, 1925: on, or very close to, margin 2 33, 65 [host, illus.]; –Vayssière, 1927: 3 [host comment]; –Wise, 1977: 105 [checklist]; –Deitz & Tocker, 1980: 28 Stigmatic spines very long, at least 10 x longer than [checklist]; –Hosking & Kershaw, 1985: 201 [ecology]; – marginal spinose setae 3 Ben-Dov, 1993: 149 [world catalogue]. —Stigmatic spines shorter than 5 x length of marginal Unmounted material: "Test of adult female usually spinose setae 4 conical, less frequently elongato-convex, open beneath, with a distinct fringe of small segments which are often 3 Dorsal macropores highly convex; with an invaginated sharply triangular; main segments large, patched with pore (much smaller than dorsal macropore) also brownish-green or often altogether light-green, sometimes present in reticulation lines; ventral tubular ducts few white, distinctly striated with air-cells, which are generally or absent medially on pro- and mesothorax; with 2 largest near the margins of the segments. Length of test pairs of setae along anterior margin of anogenital fold often reaching 1/5th in. [5 mm], height from 1/10th-1/7th ornata in [2.5-4 mm]. Texture glassy." (Maskell, 1891, p. 13). —Dorsal macropores (or largest dorsal pore) tubular and Specimens on thin twigs, although very conical, narrow invaginated, at least 4-5 x longer than broad, with a laterally. sclerotised distal end, present in reticulation lines; ventral tubular ducts frequent medially on pro- and Mounted material: when off narrow twigs, body mesothorax; with 3 pairs of setae along anterior elongate-oval but rather more broadly oval when off margin of anogenital fold ornatella thicker twigs, and pointed anteriorly; anal cleft about 1/ 12th body length. Length 2.2-3.4 mm; breadth l.3-1.7 4 Reticulation point setae on margin noticeably long, ratio mm. of length of ordinary marginal setae: reticulation setae: stigmatic setae = 1: 2: 4-5; large dorsal simple pores Dorsum: dorsal pores delineating reticulation areas in absent laterad to each anal plate; pairs of long 5 longitudinal rows, with a very large reticulation area pregenital setae only present on abdominal segment medially on dorsum, and with rather large reticulation VII neofagi areas mediolaterally; with 5 reticulation areas between anal plates and anterior margin and 26 smaller areas —Reticulation point setae on margin only slightly longer around margin. Dorsal pores of 3 types: (i) minute, dark than ordinary marginal setae, ratio (as above) = 1 : 1.3 microductules: restricted to lines of reticulation, rather : <4; large dorsal simple pores present in a group on sparse; (ii) larger, simple pores of variable size: smallest either side of anal plates; pairs of long pregenital setae pores throughout each reticulation area and also present on abdominal segments VI and VII 5 associated with reticulation lines; slightly larger pores in a submarginal line, with 24-37 on each side between 5 With >20 large simple pores on either side of anal plates; stigmatic clefts; largest pores in an elongate group on pregenital disc-pores extremely abundant medially on either side of anal plates, with 21-32 in each group; and all abdominal segments and with more than 20 across (iii mo e a e y a ge mac o o es (5 μm iam s ig y metathorax; pregenital disc-pores absent from co e wi icke e ma gi s a a g a u a e su ace segment VII; only on Nothofagus fagi es ic e o i es o e icu a io mos a u a a ou —With about 10 large simple pores on either side of anal a ge me ia e icu a io a ea u wi a ew mo e plates; pregenital disc-pores about 1/3 as numerous a e a y; wi -31 i os e io me ia mac o o e i e and with less than 6 across metathorax; pregenital A a a es 1 5-19 μm o g com i e wi s 1 -195 disc-pores present on segment VII; only on Kunzea μm; wi - mi u e o es o o sa su ace o eac a e; and Leptospermum leptospermi e g o se ae i e ma gi 1 (a ou 1/3 om a e 88 d n & nd r n (2000 : C d (In t : pt r : C d 7- μm; i e ma gi ( ea a e 1 - 7 μm; a ica NZAC: 1/l pharate [f] ad here designated. Maskell (1891) 7-3 μm; a ou e ma gi (o o sa su ace ea a e states off Fagus [=Nothofagus] var. sp., Reefton district 7-3 μm A oge i a o wi ai s o se ae o (BR). a e io ma gi a 1 ai a e a y; o ges 59- μm PARALECTOTYPES: NEW ZEALAND: (i) as for Margin: marginal setae strongly spinose: with 37-63 lectotype: NZAC: 3/1 [m] ad, 2 1st, 1 [f] 2nd (latter stained on each side between stigmatic clefts, those on either side and remounted by RCH); (ii) mounted from Maskell's dry of stigmatic spines extending slightly onto dorsum, collection (blue pill box), by J.A. de Boer, NZAC: 2/6[ff] forming a small group in each cleft; non-reticulation setae ad; & by C.J. Hodgson: 10/11[ff] ad, 4 lsts. eac 1 -1 μm o g; e icu a io se ae is ace s ig y o o o sum a a ge eac - μm o g wi 7 o Other material: NEW ZEALAND: labelled Inglisia fagi either side of abdomen, 2 on each side between stigmatic Maskell, #118, USNM: 3/2[ff] ad, 4 1st, (in quite good clefts and 8 around head; spines absent from along condition). TO: Ohakune, Nothofagus [as Fagus] fusca, margins of anal cleft. Stigmatic spines set distinctly onto 10 Jun 1924, G. Brittin, #133: [one each NZAC & USNM] o sum; e g 1- 5 μm 2/3[ff] ad (poor). NN: Riwaka, Fagus sp., 8 Feb 1925, G. Venter: pregenital disc-pores distributed across all Brittin, #133: 1/1 [f] ad (poor). As previous, except 25 Jul abdominal segments; number on each segment (medially 1926: 1/1[f] ad (poor). Motueka, Nothofagus fusca, 3 Jan across segment/in each lateral group): VII, 0 (very rarely 1938, G. Brittin, #133: 1/3[ff] ad (poor). Eve's Valley, 1)/32-43; VI, 86-88/63-75; V, 85-120/38-49; IV, 90- Nothofagus menziesii, 29 Feb 1972, J.A. de Boer, #795: 1/ 100/29-38; III, 61-94/20-28; and II, 27-43/7-14; with 2[ff] ad (poor). Mt Arthur, Flora Hut, Nothofagus fusca, 26-33 across metathorax and 0-8 disc-pores laterad to 4 Feb 1983, C.F. Butcher, #83-314b: 1/1[m] 2nd. BR: Slab each metacoxa; none on meso- or prothorax or near scape. Hut Creek, nr Reefton Saddle, Nothofagus fusca, 11 Feb Spiracular disc-pores: with 25-32 in each anterior band 1970, A. Holloway, # 673: 1/1[m] ad. Capleston, nr and 30-37 in each posterior band (latter band usually with Reefton, Nothofagus menziesii, 10 Nov 1972, J.A. de some larger disc-pores closely associated); with 1-2 pores Boer, #926a: 1/2[ff] ad (good; dorsum and venter split). (rarely none) extending medially past peritreme. Ventral Hochstetter Forest, nr Reefton, Nothofagus menziesii, 9 microducts and tubular ducts as for genus. Ventral setae: Nov 1972, J.A. de Boer, #945: 11 [m] ad, 1 [m] 2nd. Reefton, e a a a o e se ae 9-5 μm o g a e s i ose th f nz ί twig, 4 Feb 1987, J.M. Cox, #644: tapering to a blunt or finely pointed apex; with 1-4 pairs of 1/2[ff] ad (poor). anterior anal cleft setae; with a pair of long pregenital setae me ia y o segme s II a I o ges 77- μm o g; Remarks. Adult females of C. fagi are very similar to number of setae medially on each segment: VII, 2 long + females of C. neofagi but differ in having: 8-9 short setae; VI, 2 long + 3-5 short setae; V—II, 1-4 (i) no pregenital disc-pores medially on segment VII short setae; with 2-3 setae just anterior to meta- and (present on C. neofagi); mesocoxa; 2 just posterior to procoxa, 1 longer than other, (ii) long pregenital setae on two posterior abdominal a ou 5-3 μm o g; su ma gi a se ae ose o o a segments (in segment VII only on C. neofagi); and abdomen near reticulation marginal spines particu- (iii) from 2 x to 5 x more pregenital disc-pores on each a y a ge wi a ge asa socke s e g 13-17 μm; abdominal segment, both medially and mediolaterally; with only 1 large submarginal seta laterally between (iv) a group of about 20-30 large dorsal simple pores stigmatic clefts; smaller setae also occasionally present laterad to each anal plate (only 2-6 small simple pores e wee a ge su ma gi a se ae eac 9-11 μm o g; in C. neofagi); with 7 interantennal setae. Antennae 6-segmented: total (v) i egs a ou 1/3 sma e (a ou 1 μm a e a e g 33 μm 3 segme muc o ges (1 - 1 μm a ou μm o C. neofagi); long; ratio to total length 0.52:1); length of apical seta 36- (vi) distinctive macropores; 1 μm e g o c y eo a a s ie Ι 7 -1 μm Wi (vii) only 5 longitudinal lines of reticulation areas (seven o eac s i acu a e i eme a e io 5 -7 μm os e io on C. neofagi). 7 - 1 μm egs se a a io o i ia a a sus i is i c ; C. fagi differs from other females of Crystallotesta in e g s (me a o acic co a 9 -1 3 μm oc a e + having a particularly large central reticulation area and emu 115-1 7 μm i ia + a sus 1 -17 μm c aw 1 - rather short, conical stigmatic spines. μm; c aw wi ou a e ic e Biology. When mature and larvipositing, the body of the Ma e ia e ami e EC O Y E Ϋ EW EA A female bends as though hinged between the thorax and labelled "Inglisia fagi, adult female, 1889, W.M.M." abdomen, so that the abdomen slowly turns up vertically n f l nd 4 8 beneath the test, in a concertina fashion as it does so. Old Dorsum: dorsal pores delineating reticulation areas in specimens are extremely difficult to slide mount due to this 5 longitudinal rows across body; with 7 or 8 reticulation position. Possibly (as on K l r d pr the nymphs areas between anal plates and anterior margin and 26 areas come to fill the resulting space beneath the test and around margin. Dorsal pores of 3 types: (i) small, dark disperse from under its slightly loosened posterior. microductules: present throughout, but most frequent C. f (as In l f was implicated in the death of within reticulation lines; (ii) simple pores all slightly a large number of red beech trees ( th f f in an larger than microductules, of 2 sizes: smaller pores, outbreak of the scale insect in the Maruia Valley, South frequent along lines of all reticulations, and slightly larger Island, between 1976 and 1978. However, Hosking & pores, restricted to a submarginal band and laterad to anal Kershaw (1985) concluded that the scale insect population plates; and (iii) heavily sclerotised, convex macropores: expansion was incidental and due to changes induced in present medially and mediolaterally on thoracic and the trees having been stressed by several years of drought. abdominal segments and occasionally on head. Anal They noted that (Hosking & Kershaw, 1985, p.206) "the a es wi a a e u a e ; 153-1 7 μm o g males and younger female instars occur on the underside com i e wi s 1 - 3 μm; wi 1-3 mi u e o es o of leaves, but the adult female occurs on the twigs as white o sa su ace o eac a e; se ae s i ose; e g o se ae conical protuberances". Unfortunately, specimens i e ma gi 1 1 -1 μm; i e ma gi 1 -1 μm; collected at that time have not been located for the present a ica 1-3 μm a ou e ma gi 1 -17 μm study. A oge i a o wi ai s o se ae a o g a e io ma gi a a si g e ai a e a y; o ges 5 μm o g Pathogens and parasitoids. Hosking & Kershaw (1985) Margin: marginal setae spinose with a blunt or also noted that "the collapse of the epidemic appeared to slightly swollen apex and with a rather thin basal socket, be caused by an entomogenous fungus, probably e g 9- μm; wi 3 -59 o eac si e e wee a e a p r ll d pl x". s igma ic c e s; e icu a io se ae o i e e ia e S igma ic s i es ese o a se ; w e ese Distribution. With a single site record from central North is ace some is a ce o o o sum a ou e e o Island (Ohakune), otherwise in the northern third of the s i acu a isc- o e a ; eac s i e s o a s ou South Island (Map 10). sometimes sharply pointed and then similar to marginal se ae o e wise u a a e e ; e g 1-3 μm Eyes o o scu e o a se Cr t ll t t f (Maske ew com i a io Venter: pregenital disc-pores with mainly 10-11 loculi (range 8-14); number on each abdominal segment (medially/laterally on one side): VII, 11-13/70-73; VI, Fig. 106 60-64/26-32; V, 36-54/15-27; IV, 54/15-23; III, 44-49/ Ct n h t n f Maskell, 1884: 131; -Maskell, 1887: 8-17; and II, 16-41/6-17; metathorax with 21-37 70 [description]; -Maskell, 1895a: 13 [checklist]; - medially and 15-29 laterad to each coxa. Spiracular disc- Cockerell, 1896: 330 [checklist]; -Fernald, 1903: 160 pores with mainly 5 loculi (range 3-10), in a band l-3 [world catalogue]; -Hutton, 1904: 226 [checklist]; - pores wide between each spiracle and margin and Myers, 1922: 199 [checklist]; -Wise, 1977, 104 extending some distance onto dorsum through line of [checklist]; -Deitz & Tocker, 1980: 29 [checklist]; -Ben- marginal setae; with 29-44 disc-pores in each anterior Dov, 1993: 102 [world catalogue]; -Henderson, 1995: band and 27-46 in each posterior band (each band with 6- 106 [lectotype designated]. 12 disc-pores on dorsum); anterior bands with 0-1 pores a short distance mesad to each peritreme. Preantennal pores Unmounted material: available dry material consisted of apparently absent. Ventral microducts and tubular ducts large, nearly round, moderately convex adult females with as for genus. Ventral setae: ventral anal lobe setae 21-22 thick, white wax test plates. As pointed out by Henderson μm o g; wi 1 ai o a e io a a c e se ae; wi 1 ai (1995), the description given by Maskell (1884: 131) o o g ege i a se ae me ia y o segme s I a II; seems to be more appropriate for l h t n l rp um e o se ae me ia y o eac a omi a segme (Maskell) than for C. f . II - ; I -3; -II - (? (a e y s o a a om ege i a ai s ; wi 1- se ae ea eac me aco a - Mounted material: body oval to almost round and rather ea eac mesoco a a - s o se ae ea eac convex; anal cleft about 1/9th body length. Length 3.0- procoxa; longest seta on thorax associated with 5.0 mm; breadth 2.1-3.6 mm. mesoco ae 3 -35 μm o g; wi 1-3 ai s o 9 d n & nd r n (2000 : C d (In t : pt r : C d interantennal setae laterally plus 0-1 seta medially; with dorsum and by either the absence of stigmatic spines or, 1-3 submarginal setae between each stigmatic area; with when present, their displacement some distance onto the small setae distributed rather randomly submarginally. dorsum. Other important characters are the presence of: Antennae 6- to 8-segmented; 3rd (or 3-5) segment(s) (i) ventral tubular ducts throughout the venter; about as long as other 5 segments together; total length (ii) a second type of ventral tubular duct around the 95-35 μm; e g o a ica se a 19- μm e g o anogenital area; c y eo a a s ie 1 3- 7 μm Wi o s i acu a (iii) only 5 longitudinal rows of reticulation areas dorsally; e i emes a e io 3-7 μm os e io 7 - 3 μm egs (iv) ventral microducts throughout the venter; well developed; separation of tibia and tarsus usually (v) pregenital disc-pores medially on metathorax and also is i c ; e g s (me a o acic co a 5 - 1 μm laterad to each metacoxa; oc a e + emu 135-1 μm i ia 97-119 μm a sus (vi) the presence of a very small denticle on the claw; -97 μm c aw 1 μm; c aw wi a is i c u e y sma (vii) the convex structure of the dorsal macropore; denticle. (viii) the shape of the marginal spines, which are often slightly dilated at their apex. Material studied: LECTOTYPE [f]: NEW ZEALAND: labelled: "Ctenochiton fuscus, adult female, from Biology. Nothing known Brachyglottis sp., Jan. 1883, W.M.M." CMNZ: 1/1[f] ad; designated by Henderson, 1995. Distribution. From Cook Strait (Trio Islands) to Lake PARALECTOTYPES: NEW ZEALAND: (i) from Panax Tekapo in the South Island (Map 11). (=Pseudopanax) sp., Jan. 1883, W.M.M., CMNZ: 2/1[m] 2nd (although labelled female 2nd), 1 [f] ad test. (ii) as previous, except NZAC: 1/1 "antenna of female". Cr t ll t t l pt p r (Maske ew com i a io Other material: NEW ZEALAND: from W.M. Maskell's dry material #37, mounted 2 Mar 1972 by J.A. de Boer: 1/ Figs C46, C47, 107 1 pupa. SD: Trio Is, Hymenanthera sp., Aug 1965, D.J. Campbell, #61: 5/10[ff] ad. NN: Motueka, Melicytus Inglisia leptospermi Maskell, 1882: 220; —Maskell 1885: ramiflorus, 9 Jan 1937, 8 Jan 1938 & 12 Mar 1938, G. 27 [male]; -Maskell, 1887: 75 [description]; —Maskell, Brittin, #326: 3/69 [f] ad. Motueka,Myoporum[as laetum 1895a: 14 [checklist]; -Cockerell, 1896: 330 [checklist]; — ngaio], 4 Jul 1917, G. Brittin, No. 113, NZAC: 1/1 [m] 2nd Fernald, 1903: 162 [world catalogue]; —Hutton, 1904: 227 pharate prepupa. Golden Bay, Tarakohe, Melicytus sp. , 14 [checklist]; —Myers, 1922: 199 [checklist]; -Green, 1929: Aug 1925, (G. Brittin dry collection), #90-213b: 1/1[f] ad. 377 [record];-Hoy, 1954:601 [distribution];-Hoy, 1961: MC: Lyttelton, Myoporum laetum, no date, G. Brittin, 52, 57 [distribution];-Wise, 1977: 105 [checklist];—Deitz #110, BMNH: 7/78, 2/2[ff] ad + NZAC (mounted from & Tocker, 1980: 29 [checklist]; —Ben-Dov, 1993: 148 dry material by C.J. Hodgson): 6/3[ff] ad, 1 [m]ad, 8 1sts (as [world catalogue]; —Henderson, 1995: 107 [lectotype `Ctenochiton testudo', Brittin manuscript name). Lyttelton, designated]. Myoporum laetum [as ngaio], 1914, from Brittin dry collection, #110, #90-220a-d: 4/4[ff] ad. As previous, Unmounted material: "Test of adult female white, glassy except 13 Jan 1917, G. Brittin: 2/2[ff] ad. Springfield, or waxy, elongated, convex above, flat and open beneath, Coprosma sp., 26 Dec 1914, G. Brittin, #71: 1/1[f] ad [as formed of several agglutinated segments, each segment Inglisia ornata]. Okuti Valley Scenic Reserve, Melicytus more or less convex or conical, median segments usually ramiflorus stems, 19 Jan 1983, L.M. Emms, #83-052d: 4/ five in number; at the edge an irregular fringe, but the 8[ff] ad. MK: Lake Tekapo, Hymenanthera sp., 4 Feb fringe is often absent. Average length of test, 1/10th in 1968, J.A. de Boer, #352: 2/3[ff] ad (l split [2.5 mm]. The marginal segments sometimes assume the dorsoventrally). form of small cones, as if a number of secondary tests were attached to the principle one. All the segments are marked Remarks. Maskell's material was collected at with striae radiating from the apex of each: the striae, Canterbury, the Brachyglottis species was B. repanda and which are composed of air-cells, widen from the apex to the n x sp. was P. arboreum (=Pseudopanax arboreus) the base." (Maskell, 1887, p. 75). Young female chestnut (Maskell, 1895a). brown; adult female shrivels to anterior end of test whilst The adult females of C. fusca are easily recognisable ovipositing, causing near empty test to appear very white by the extension of the spiracular disc-pore bands onto the (this is stage most often collected). n f l nd 4 Mounted material: body elongate oval, about equally 7 -9 μm; um e o s o se ae me ia y o eac rounded at both ends and parallel-sided; anal cleft about 1/ segme II— I -5; -1; I - ; a III—II ; wi 11th body length; stigmatic clefts not indented. Length -3 se ae us a e io o me aco a; - ea eac 1.9-2.4 mm; breadth 0.89-1.22 mm. mesoco a a 3 us os e io o eac oco a a e muc o ge a ose ea o e co ae e g 3 -3 Dorsum: dorsal pores delineating reticulations in 7 μm; wi 1-3 su ma gi a se ae o eac si e e wee longitudinal rows across dorsum, with 5+ reticulation s igma ic a eas; wi 3- ai s o i e a e a se ae areas between anal plates and anterior margin and about 28 a e a y + - me ia y A e ae -segme e wi o areas around margin. Dorsal pores of 3 types: (i) minute, seu osegme s; o a e g 5 -3 9 μm; e g o dark microductules: present throughout, but most frequent a ica se a -5 μm e g o c y eo a a s ie 15 - associated with lines of macropores; (ii) simple pores of 171 μm Wi o s i acu a e i emes a e io 3 - 1 various sizes: flat simple pores, slightly larger than μm os e io 37- 5 μm egs wi i ia a a sus microductule: present throughout but most frequent is i c y se a a e; e g s (me a o acic co a 133-1 associated with lines of macropores, slightly larger near μm oc a e + emu 1 9-17 μm i ia 1 -1 μm margin; larger simple pores: forming a distinct a sus 91-11 μm c aw 1 μm; c aw wi o wi ou a submarginal band; still larger simple pores are present in a mi u e e ic e line on either side of anal plates, with 6-11 in each group; and (iii) macropores, approximately similar in size to Material examined: LECTOTYPE [f]: NEW ZEALAND: simple pores near anal plates: frequent within inner labelled "Inglisia leptospermi from manuka tree, two adult reticulation lines; with about 8 in posterior medial females, Oct. 1880, WWM"; CMNZ: 1/2[f] ad, lectotype mac o o e i e A a a es 1 9-157 μm o g com i e arrowed; designated by Henderson, 1995. wi s 113-1 1 μm; wi -3 mi u e o es o o sa PARALECTOTYPE: 1 [f] ad on slide with lectotype. su ace o eac a e; e g o se ae i e ma gi 1 1 - [Paralectotype: collection data as for lectotype except 1 μm ( i e y s i ose ; i e ma gi 1 - μm (mo e female - 2nd stage; CMNZ: 1/1 [f] 3rd, a misidentification, s ou y s i ose a a mos o a e ; a ica 1-3 μm a now considered to be Crystallotesta ornatella described as ou e ma gi 1 -1 μm ( o a mos o a e a e se new below. This specimen = Slide 2 under Inglisia s ig y o o o sa su ace o eac a e A oge i a o leptospermi, p. 107, Henderson (1995)]. wi -3 ai s o se ae o a e io ma gi a 1 se a o eac a e a ma gi ; o ges - 5 μm Other material: NEW ZEALAND: Maskell coll., no. 47, Margin: marginal setae conical and spinose, 10-15 USNM: 4/2[ff] ad, (one very poor), 1 1st, 4 ?eggs. New μm o g; wi 9-3 a e a y e wee s igma ic c e s; Zealand, Kunzea ericoides [as Leptospermum], 30 Apr e icu a io se ae s ig y a ge 15- μm o g; wi 19 Mye s M 1/1[ ] a ΤΗ ee Ki gs Is G ea sim e o es e o o e wee ma gi a se ae wi 5-1 I a i Kunzea ericoides, underside leaves, 14 Apr o eac si e e wee s igma ic a eas S igma ic s i es 1999, T.K. Crosby, #99-026a—d: 4/2[m] 2nd, 2[f] 3rd. CL: e i g o e oa u i e y ea a e s ig y cu e ; Little Barrier I, Te Maraeroa West, Kunzea ericoides, 17 o se s ig y o o o sum; e g 5 -7 μm Eyes o Sept 1994, RCH, #94-077a—e: 5/1[f] ad, 29 3rd (one o scu e o ma gi pharate), 2[f] 2nd, 2[m] 2nd, 1 1st. Tairua SF, Leptospermum Venter: pregenital disc-pores with mainly 10 (range scoparium, a 19 1 A E Ma sack ΝΖ o 1/ 6-13) loculi; number on each segment (medially across 1 [ ] a G ao eo e Kunzealeaves & ericoides, twig, segment/laterally on one side): VII, 11-16/17-34; VI, 28- 15 Mar 1994, & 2 Nov 1994, RCH, #94-050b: 1/1[f] 2nd, 38/17-20; V, 26-32/6-14; IV, 23-28/7-12; III, 18-26/6- 2[m] 2nd, & #94-101: 1/1 [f] ad (good). TO: Kaingaroa SF 8; and II, 12-16/2-5; metathorax generally with at least 1 Cpt 1164, Leptospermum scoparium, 10 Aug 1959, N.O. disc-pore laterad to each coxa and often with 1-2 medially. Secom e ΝΖ 99 1 /7[m] S U i eI, Spiracular disc-pores in bands l-2 pores wide between Leptospermum sp., 17 Feb 1971, J.A. de Boer, #700: 2/ each spiracle and margin; with 12-18 in each band, with 3[ff] ad (one split dorsoventrally). NN: Riwaka, none extending medially past peritreme. With preantennal Leptospermum sp., Jan 1918, no coll., USNM #72: 1/2[ff] pores single. Submarginal band of simple pores absent. ad. Nelson, Rockville, Arorere R, Kunzea ericoides [as Ventral microducts and tubular ducts as for genus. Ventral Leptospermum], 4 Aug 1948, T.G. Sewell, #59 J.M. Hoy se ae e a a a o e se ae qui e o g 1- 5 μm a Collection: 5/1[f] 3rd, 4[f] 2nd, 4[m] 2nd. Onekaka,Kunzea ge e a y wi a u o e e s ig y swo e a e ; wi ericoides [as Leptospermum], 4 Aug 1948, T.G. Sewell, - ai s o a e io a a c e se ae; wi a ai o o g #62 J.M. Hoy Collection: 2/2[m] 2nd. Pokoporo, Motueka ege i a se ae me ia y o segme s I a II o ges Valley, Kunzea ericoides [as Leptospermum], 4 Dec 1959, 92 d n & nd r n (2000 : C d (In t : pt r : C d G Mc u ey ΝΖ (a 3 / [ ] a e so were misdiagnosed and the key characters refer to a third K nz r d [as pt p r ], 19 Dec 1968, J.A. species, Cr t ll t t rn t ll , described as new below. de Boer, #471: 2/4[ff] ad (one split dorsoventrally). BR: Buller R. Gowanbridge [as nr Gower Bridge], K nz Biology. Juvenile instars of both sexes are found on the r d [as pt p r ], 3 Mar 1969, J.A. de Boer, leaves of the host plant, whereas the adult females settle on #494: 1/2[ff] ad. Maimai, K nz r d [as pt the twigs. The very young adult female is a cryptically p r ], 1 Aug 1948, T.G. Sewell, #67 J.M. Hoy coloured chestnut brown with spots and stripes and closely Collection: 5/1[f] 2nd, 4[m] 2nd, 6 1sts. Reefton, 3 miles resembles the pattern of the leaf bud bracts of K nz north, K nz r d [as pt p r ], 2 Aug 1948, r d (Colour plate 1). From the collection records, T.G. Sewell, #64 J.M. Hoy Collection: 4/3[ff] ad (old), 1 [m] there is apparently one generation per year, with the 1st- 2nd, 8 1sts. Reefton, 4 miles south of Ahaura, K nz and 2nd-instar nymphs overwintering and the adult r d [as pt p r ], 2 Aug 1948, T.G. Sewell, females reproducing over the summer and into autumn, #68 J.M. Hoy Collection: 5/1[f] ad (pharate), 1[f] 3rd, 3[f] depending on geographic location. 2nd (one slide too poor). Inangahua Landing, pt p r p r , 2 Aug 1948, T.G. Sewell, J.M. Pathogens and parasitoids. Hymenopterous parasitoids Hoy Collection: 3/1[f] 3rd, 1 [f] 2nd,[m]2nd.4 Waipuna recorded are: Encyrtidae: Ad l n rt d v r b l (Reefton District), K nz r d [as pt p r ], Noyes; Pteromalidae: Aph b t n n ( ouček Oc 19 o ag ΝΖ 5 /3[ ] u ae aw ai S. ., K nz r d [ pt p r ], Distribution. Throughout areas with a drier climate in the 18 Jan 1960, J.G.R. McBurney, FRNZ R(a)36: 3/3[ff] ad. North Island and the northern half of the South Island MC: Springfield, pt p r sp., 25 Dec 1914, (Map 12) Brittin, #72: 1/1[f] ad. Remarks. Also recorded from: MC: Governors Bay Cr t ll t t n f e e so & o gso ew (Green, 1929: 377). s ecies Cr t ll t t l pt p r has been recorded only Fig C45, 108 from pt p r p r —manuka— and K nz r d [also as pt p r r d ]— n . C. Unmounted material: unknown. l pt p r can be identified by having the following combination of characters: Mounted material: body broadly oval, rather more (i) a line of large dorsal simple pores on either side of anal pointed anteriorly. Length 2.3-2.6 mm; breadth 1.5-1.7 plates; mm. (ii) dorsal reticulation areas in 7 longitudinal rows; Dorsum: dorsal pores delineating reticulation areas in (iii) 8-segmented antennae; 7 longitudinal rows across dorsum; with 5 reticulation (iv) pairs of long pregenital setae on abdominal segments areas between anal plates and anterior margin, middle 3 VI and VII; rows with rather fewer areas than on other species in this (v) the convex shape and unusual distribution of the dorsal genus, and with 27-28 areas around margin. Dorsal pores macropores; of 3 types: (i) rather large, dark microductules (whose (vi) the narrow, moderately short, curved stigmatic spines. orifice often looks as though it is bilocular): common on In possessing a line of large simple pores laterad to the either side of macropores in reticulation lines; (ii) similar anal plates, it resembles C. f but these two species are sized, flat to slightly convex, simple pores: variable in size, otherwise easily separable by the characters listed above. associated with lines of reticulation but those nearest An attempt was made (Henderson, 1995) to sort out margin distinctly larger: with 12-17 in a submarginal line the Maskell coccid species recorded from pt p r on each side between stigmatic clefts and with 2-6 in a line p r and K nz r d (as pt p r laterad to each anal plate; and (iii) quite large macropores r d . Further study has shown that it was only partly (3-5 μm iam o a y a e a ei e mo e o ess successful: the key characters and the discussion given in round or roundly lemon-shaped, with lateral walls Henderson (1995) for In l rn t Maskell are correct thickest: with 9-12 in posterior medial macropore line. but with regard to In l l pt p r Maskell, the type A a a es 153-171 μm o g com i e wi s 1 -11 specimens are those illustrated by Maskell in 1887; the μm; mi u e o es o o sa su ace a a e y a se ; characters of the female lectotype given by Henderson e g o se ae i e ma gi 1 1 μm ( ossi y a se o n f l nd 4 93 some specimens but when present about middle of inner acic co a 155-1 μm oc a e + emu 19 - 1 ma gi ; i e ma gi ( ea a e 5-35 μm; a ica μm i ia 1 3-1 μm a sus 9 -1 3 μm c aw - 5- μm a ou e ma gi 3 - 5 μm (se o o sa μm; c aw wi ou a e ic e surface near apex). Anogenital fold with 2 pairs of setae Material examined: HOLOTYPE [f]: NEW ZEALAND: on anterior margin and 1 pair on lateral margins; longest BR: Granville Forest, 8 Aug 1957, E.E. Ensor, 5 - 3 μm th f f , FRNZ R28-30: 1/1 [f] ad. Holotype Margin: marginal setae spinose: with 2-4 spines slide marked as such; [misidentified as In l f extending a short distance up anal cleft; with 32-39 on Maskell by C.F. Butcher 1984] (remounted by C.J. each side between stigmatic clefts, those on either side of Hodgson). stigmatic spines displaced slightly onto dorsum, forming a PARATYPES: as for holotype: NZAC: 11/2[ff] ad, 7[ff] sma g ou i eac c e ; mos se ae 1 - μm o g u pharate ad., 1 [m]2nd , 1[f] 3rd . e icu a io se ae sig i ica y a ge -3 μm a displaced slightly onto dorsum; with 7 larger, reticulation Other material: NEW ZEALAND: GB: L. Waikaremoana, setae on either side of abdomen, 2 on each side between 19 Jun 1991, C.F. Morales, th f f , #97-074a— stigmatic clefts and 8 around head. Stigmatic spines set c: 3/1[f] 3rd, 6[m]2nd, 5 pupae. TO: SF 90, Taupo, s ig y o o o sum; eac 7 -99 μm o g th f f , 8 Dec 1975, M. Kay, FRNZ R(b)73- Venter: pregenital disc-pores with mainly 10 (7-11) 78: 6/10[ff] ad (very poor). BR: Maimai [as Mai-Mai], loculi; distributed across abdominal segments; number on th f tr n t , Aug 1977, T. Johnson, #77-251 a: 4/ each segment (medially across segment/laterally on 1 1[f] ad, 3 pharate [ff] ad, 1[f] 2nd, 2[f] 3rd. side): VII, 5-9/14-21; VI, 40-61/7-14; V, 36-52/3-7; Remarks. The adult females of Cr t ll t t n f are IV, 22-32/4-6; III, 11-17/4-6; and II , 8-10/1-5; with 5- very similar to those of C. f but differ in having: 9 medially across metathorax and with 1-3 laterad to each (i) long pregenital setae on only abdominal segment VII metacoxa; 0-1 mesad to each mesocoxa; 0-2 mesad to (segments VI and VII on C. f each procoxa and 0-1 mesad to each scape. Spiracular disc-pores with 3-7 (mainly 5) loculi in bands 1-5 pores (ii) at least 5 pregenital disc-pores medially on segment wide between each spiracle and margin; with 36-43 in VII (none or very rarely 1 on C. f each anterior band and 41-47 in each posterior band; with (iii) one third as many pregenital disc-pores medially and 1-2 pores (rarely none) extending medially past peritreme. mediolaterally on all abdominal segments; With 1-2 preantennal pores just anterior to each scape. (iv) few (2-6) dorsal simple pores laterad to each anal plate Ventral simple pores absent. Ventral microducts and (about 30 on C. f tubular ducts as for genus. Ventral setae: ventral anal lobe (v) legs about 1/3rd longer than on C. f (a ou μm se ae -7 μm o g a e s i ose wi a u o e e a e a a ou 1 μm o C. f slightly capitate apex; with 0-1 pair of anterior anal cleft (vi) rather few (<15) macropores in the transverse median setae; with a pair of long pregenital setae medially on line anterior to the anal plates (C. f has more than segme II o y 3-9 μm o g; um e o s o e se ae 20); medially on each segment: VII, 7-9; VI—V, 2-4; IV-II, 0- (vii) many fewer pregenital disc-pores medially on the 4; with 1-3 setae just anterior to each meta- and mesocoxa thorax (5-9 as compared with 23-26 on C. f and and 1-3 just posterior to each procoxa; length of setae these extending onto meso- and prothorax; associa e wi eac oco a 13-1 μm; su ma gi a (viii) the test with seven lines of reticulation plates (only 5 setae: those on thorax and abdomen near reticulation setae on C. f . particularly large, with large basal sockets, length 27-33 μm; wi 1 a ge se a o eac si e e wee s igma ic c e s Biology. C. n f has been collected in March, June, u a so wi occasio a sma e se ae 1 -15 μm o g August, and December. Several life stages were collected between larger submarginal setae; interantennal setae: in June and August and therefore C. n f may have just with 1 pair of long setae + 1-3 pairs short setae. Antennae one generation a year. Found only on th f spp. 6-segmented, with 2 pseudosegments in long third Distribution. The distribution is apparently disjunct, with segme ; o a e g 3 -39 μm; i segme muc two records from central North Island and two records o ges (1 - 1 μm o g; a io o o a e g 5 1 ; from north-west South Island (Map 13). e g o a ica se a 39- μm e g o c y eo a a s ie 1 5-17 μm o g Wi o s i acu a e i emes Name derivation. A combination of n , new, with fagi, a e io 55-7 μm os e io 1-7 μm egs se a a io for the similarity to that species. of tibia and tarsus generally distinct; lengths (metatho- 4 d n & nd r n (2000 : C d (In t : pt r : C ld Cr t ll t t rn t (Maske ew com i a io reticulation lines and most abundant medially on thorax and abdomen; with 18-21 in posterior medial macropore i e A a a es 1- 1 μm o g com i e wi s igs Μ19 C C 9 1 9 1 -173 μm; wi -3 mi u e o es o o sa su ace o Inglisia ornata Maskell, 1885: 27; —Maskell, 1887: 76 eac a e; e g o se ae i e ma gi 1 5 μm; i e [description]; —Maskell, 1895a: 14 [checklist]; — ma gi μm; a ica 5 μm a ou e ma gi 3 - Cockerell, 1896: 330 [checklist]; —Fernald, 1903: 162 μm (se o o sa su ace o eac a e ea a e [world catalogue]; —Hutton, 1904: 227 [checklist]; - Anogenital fold with 1-3 pairs of setae on anterior margin Myers, 1922: 199 [checklist]; —Miller, 1925: 33, 63 a wi 1 ai o a e a ma gi s; o ges -9 μm [host]; —Hoy, 1954: 601 [distribution];—Hoy, 1961:52,57 Margin: marginal setae spinose: with 50-67 laterally [distribution]; -Wise, 1977: 105 [checklist]; -Deitz & between stigmatic clefts, those just posterior to each cleft Tocker, 1980: 30 [checklist]; —Ben-Dov, 1993: 149 some imes i a g ou -3 wi e; e g 9- μm; a ge [world catalogue]; -Henderson, 1995: 107 [lectotype setae at intersegmental boundaries and not coinciding with designated]. reticulation junctions at margin. Simple pores present in a submarginal ring next to or between marginal setae, with 13-20 pores on each side between stigmatic areas. Unmounted material: "Test of adult female reddish Stigmatic spines very long, slightly tapering and curved; brown, the base more or less oval, the rest elevated in a e g 3 -3 μm Eyes o o scu e cone and ending in a prominence standing up like a more Venter: pregenital disc-pores with mainly 10 (6-11) or less sharp horn; sometimes there are two of these horns. loculi; number on each segment (medially across segment/ The test is formed of a number of polygonal segments, laterally on one side): VII, 40-62/26-30; VI, 17-27/49- each slightly elevated, and all marked with the striae 63; V, 9-14/44-64; IV, 7-13/29-44; 111, 4-11/26-41; and peculiar to the genus. There is a fringe of sharply II, 5-12/12-19; one specimen also has a single disc-pore triangular segments, also striated. Average length of test laterad to a metacoxa. Spiracular disc-pores with mainly 5 about 1/6 inch [4 mm], but specimens attain a length of (range 3-7) loculi, in bands 1-5 pores wide; with 29-41 in 1/4 inch [6 mm]; height about 1/10 inch [2.5 mm]" each anterior band and 42-57 in each posterior band; with (Maskell, 1885, p. 27). The reddish-brown colour is the 0-2 pores (usually at least 1) extending medially past animal within showing through its nearly transparent test. peritreme. With 0-1 preantennal pores. Also with a line of simple pores around submargin, approximately in line Mounted material: body oval, about equally rounded at with submarginal setae; with 12-29 on each side between both ends. Length 2.4-4.2 mm; breadth 1.7-2.5 mm. stigmatic clefts (when abundant, band more than 1 pore wide). Ventral microducts and tubular ducts as for genus. Dorsum: dorsal pores delineating reticulation areas in e a se ae e a a a o e se ae 3 -3 μm o g 5 longitudinal rows; reticulation areas in middle 3 rows all rather spinose; with 4-5 (generally 4) pairs of anterior anal large; with 5 or 6 areas between anal plates and anterior cleft setae; with long pregenital setae medially on margin and about 19 areas around margin; each ege i a segme s II I a o ges 75- μm; reticulation line broad, composed of 3 parallel lines of number of setae medially on each segment: VII—VI, 4-8 pores: a median line with macropores interspersed with long setae, no short setae; V, 2-3 long setae + 0-1 tubular pores and microductules (microductules clustered moderately long setae; IV—II, 2-4 short setae; with 1 seta around each tubular pore and each macropore in small just anterior to each meta- and mesocoxa; 0-2 setae just groups) and 2 lateral lines with sub-groups of 3 or more posterior to each procoxa; length of setae associated with pores, consisting of 1 or more simple pores with a eac oco a 1 -1 μm; wi 1 su ma gi a se a o eac microductule on either side. Dorsal pores of 4 types: (i) side between stigmatic areas; with 3-5 pairs of small, dark microductules: present within reticulation interantennal setae; other setae rather few and small. lines; (ii) slightly larger, flat, simple pores: present in both A e ae -segme e ; o a e g 3 - μm; sca e reticulation areas and in reticulation lines; of 2 sizes, those wi o y se ae; e g o a ica se a 7- 9 μm e g ea es ma gi a ges ; (iii u u a o es 7-1 μm ee o c y eo a a s ie 1 5- 7 μm Wi o s i acu a uc u e a ou 1/ -1/3 e g o a e a u u a uc e i emes a e io 55- 1 μm os e io -9 μm egs mi us i e i ame a wi a a k i e o e ese with a distinct tibio-tarsal articulation; lengths (metatho- e wee mac o o es wi i me ia e icu a io i e; a acic co a 1 - 5 μm oc a e + emu 17 - (i a ge ig y co e mac o o es o a -s a e μm i ia 135-1 5 μm a sus 9 -1 7 μm c aw 5- 9 wi wa s ickes a ase ( ig M19 ese wi i i e μm; c aw wi ou a e ic e n f l nd 4 95 Material examined: LECTOTYPE [f]: NEW ZEALAND: these specimens refer to Cr t ll t t rn t ll , which is [WN: Wellington], "In l rn t , adult female, from only known from species of pt p r , K nz El rp , Mar.1884, W.M.M." NZAC / Maskell coll.: (previously pt p r , and p n, and is very 1/1 [f] ad.; designated by Henderson, 1995. similar to Cr t ll t t rn t in outward appearance. PARALECTOTYPES: NEW ZEALAND: (i) As for This is also probably true for the material collected off lectotype , but labelled "Female — 2nd stage / stained and pt p r spp. by Hoy (1961), some of which is remounted in Canada balsam, 16 Jan 1995, R.C. included in "Material examined" under C. rn t ll Henderson" [= 2nd-instar male] NZAC: 1/1 [m] 2nd. (ii) below. "Young insect" [= 1st-instar], NZAC 1/1 1st. (iii) "Antenna, foot and spines of adult female, Oct 1884" Biology. C. rn t probably has one generation a year as NZAC: 1/bits. (iv) "Male, Aug 1884" NZAC 1/1[m] ad. adults have most often been collected in October (Spring) while nymphs have been collected throughout the summer Other material: NEW ZEALAND: Maskell collection no. and winter at the Riverhead Forest site. Male nymphs are 45, USNM: 1/1[f] ad, 1/1 [m]ad. AK: Riverhead Forest, found on leaves, while the female nymphs move to the Barlow Road reserve, d rp t t r leaves & stems, twigs where the 3rd instar and adult females are found. 4 May 1995, RCH, #95-040a+d: 2/2e 2nd. As previous, except 13 Oct 1995, #95-108a–f: 6/5[ff] ad, 2[m] 2nd. As Pathogens and parasitoids. Hymenopterous parasitoid previous, except 17 Jul 1997, #97-085a–e, 5/2[f] 3rd,[m]3 recorded: Encyrtidae: Ad l n rt d v r b l Noyes. 2nd. As previous, except 14 Aug 1997, reared to 10 Sep 1997, #97-121a–g: 6[mm] ad, 7 moults, 3[m]2nd, 3 DiStribution. C. rn t is only known from the North prepupae, 1 pupa. As previous, except 6 Feb 1998, C.J. Island (Map 14). Hodgson, #98-070a–f: 6/3[f] 2nd, 14 1sts. As previous, except 25 Jul 1999, RCH, #99-093a–j: 10/1[f] ad, 3 [f] 3rd, 6[m] 2nd, 2 prepupae, 1 pupa. BP: Te Koau, 360 m, Cr t ll t t rn t ll e e so & o gso ew d r rb r , 26 Oct 1992, J.S. Dugdale, #92-340: s ecies 1/1 [f] ad. Waiaroho,28 d r Jan rb r , 1993, RCH, #93-268: 1/1[m] 2nd. As previous, except 3 Nov 1993, #93-339: 1/1[f] ad (split dorso-ventrally). Rereauira Figs C50, C51, 110 Swamp, Beech Ridge, th f tr n t , 26 Apr 1993, Unmounted material: test of adult female similar to a J.S. Dugdale, #93-032: 1/1 [f] 2nd. Lottin Point, Otanga, small C. rn t , with a convex median horn and a series of t x l n twigs, 5 Nov 1993, RCH, #93-051a–b: 1/1[f] pointed plates at its base, which easily break apart (unlike ad, 5 1sts. GB: Taikawakawa, nr. Maungakaka, the fused test f l h t n p ll n , which can appear l h d t r , 18 Mar 1993, RCH, #93-084: 1/ similar and is about same size); young females reddish- 1 [f] ad (poor). WN: Otaki Forks,4 d rp Oct t t r , brown; mature adult females shrink towards anterior end 1980, C.F. Butcher, #83-292: 1/1[f] ad. of test and become sclerotised. Remarks. Adult females of C. rn t can be distinguished by the presence of the following characters: Mounted material: body oval, slightly more pointed at anterior end; stigmatic clefts shallow or absent. Length (i) very long stigmatic spines; l.2-2.4 mm; breadth 0.85-1.2 mm. (ii) large sclerotised, bollard-shaped macropores; (iii) only 5 lines of reticulation plates; Dorsum: dorsal pores delineating reticulation areas in (iv) 8-segmented antennae; 5 longitudinal rows across dorsum; with 5 or 6 areas (v) 6 anal ring setae. between anal plates and anterior margin and an unknown Cr t ll t t rn t ll (described below) is similar number of areas around margin; each reticulation line but differs in: broad, composed of 3 parallel lines of pores: median line (i) being nearly half the size; of tubular macropores interspersed with microductules (ii) lacking the bollard-shaped dorsal macropores; and an occasional simple pore (usually with several (iii) in having 8 anal ring setae; microductules clustered around each tubular macropore); (iv) 1 pair of long pregenital setae on segments VII and VI lateral lines with microductules and occasional simple (rather than many on segments VII–V in C. rn t . pores. Dorsal pores of 3 types: (i) small, dark Maskell (l885a, 1887) states that C. rn t was also microductules: present throughout, but most frequent collected on pt p r sp. but it seems probable that associated with lines of macropores; (ii) slightly larger, 6 d n & nd r n (2000 : C d (In t : pt r : C d flat simple pores: most abundant near margin; and (iii) 5 μm os e io 3 -5 μm egs wi i ia a a sus long, inverted tubular macropores (or largest dorsal distinctly separate; lengths (metathoracic): coxa 73-122 pores), about 1/2-2/3rds length of a ventral tubular duct μm oc a e + emu 1 -1 μm i ia -119 μm minus inner filament, with a heavily sclerotised inner end, a sus - 5 μm c aw 1 - μm; c aw wi ou a e ic e which tends to be slightly broader than duct, appearing round in dorsal view: present in all lines of reticulation; Material examined: HOLOTYPE [f]: "NEW ZEALAND: with 8-13 in posterior medial macropore line. Anal plates: AK: Hunua Ra, Oct 1982, C.F. Butcher, #83-320d, 1 9-1 μm o g com i e wi s 115-1 μm; eac NZAC: 1/1[f] ad. a e a e i g o a a icu a y a ow a e ; wi -3 PARATYPES: collection data as holotype: NZAC: 2/1 [f] mi u e o es o o sa su ace o eac a e; e g o 3rd, 1 [m] 2nd. se ae i e ma gi 1 1 - 5 μm; i e ma gi 1 -1 μm; a ica 3 - 1 μm a ou e ma gi -3 μm (se o Other material: NEW ZEALAND: "In l l pt p r " o sa su ace o eac a e ea a e A oge i a o [misidentification] ,Leptospermum p r [as manuka wi 3 ai s o se ae (o e s ecime wi o a e io tree], Oct 1880, W.M. Maskell: 1/1[f] 3rd [previously ma gi a 1 ai a e a y; o ges 3 -5 μm A a i g included as a paralectotype f r Cr t ll t t l pt p r ]. wi se ae ΤΗ: Three Kings Is, Great I, Bald Hill, K nz r d , Margin: marginal setae spinose and conical; with 29- stem, 14 Apr 1999, T.K. Crosby, #99-028: 1/1[f] ad. ND: 3 43 laterally between stigmatic clefts, those just posterior to miles north of Dargaville, pt p r p r , 9 each cleft sometimes set slightly onto dorsum; setae 9-13 Aug 1954, J.M. Hoy, #160: 5/5[ff] ad, 1 pupa, 1 [m] ad. CL: μm o g; wi a su ma gi a a o sim e o es o Little Barrier I, Te Maraeroa west, K nz r d , 17 o sum e o o e wee ma gi a s i es wi 5-1 o Sept 1994, RCH, #94-083a-b: 2/1[f] ad, 1 pharate [f]. GB: eac si e e wee s igma ic a eas S igma ic s i es e y Kokakonui Stm headwater, Kaharoa Station, K nz o g s ig y a e i g a cu e wi a we - e e o e r d , 20 Dec 1993, J.S. Dugdale, #93-376: 1/1[f] ad asa socke ; e g 11 -191 μm Eyes o o scu e (parasitised, poor). Paoneone, K nz r d twigs, 4 Venter: pregenital disc-pores with mainly 10 (6-11) Nov 1993, RCH, #93-340: 1/1[f] ad. As previous, 15 Mar loculi; number on each segment (medially across segment/ 1994, #94-049a-c: 3/1[f] ad, 1 [m]2nd, 1 pupa, l [m] ad. As in each lateral group): VII, 2-5/12-20; VI, 11-24/11-18; previous, except K nz r d , both sides leaves, 4 V, 9-21/5-9; IV, 7-15/0-5; III, 4-13/1-5; and II, 3-8/1- Nov 1995, #95-102a-b: 2/1[m] 2nd, 3 1sts. Paoneone, 3; one specimen also had a single disc-pore laterad to a p n f l t , 2 May 1993, RCH: 2/1[f] ad, 1[m] metacoxa. Spiracular disc-pores with mainly 5 (range 3- 2nd. WA: Te r , M tr d r sp. (rata), 29 May 1983, 7) loculi, in bands 1-4 pores wide; with 8-18 in each band, C.F. Butcher, #83-160a: 2/1 & 2 part [ff] ad, 2[m] 2nd. NN: but tending to have more in anterior than in posterior Motueka, pt p r p r , 13 Apr 1925, G. bands; pores not extending medially past peritremes. Brittin Collection #71: 1/1[f] ad. Onekaka, K nz Preantennal pores single. Also with a submarginal line of r d [as pt p r ], 4 Aug 1948, T.G. Sewell, simple pores, approximately in line with submarginal J.M. Hoy Collection #62: 6/2[ff] ad (one pharate), 7[f] 3rd, setae; with 7-10 on each side between stigmatic clefts. 3[m] 2nd, [+?3/3 prepupaeor?? rn t ll l pt p r ]. Ventral microducts and tubular ducts as for genus. Ventral Collingwood, K nz r d [as pt p r ], 4 se ae e a a a o e se ae 1 - 3 μm o g; wi 1- Aug 1948, T.G. Sewell, J.M. Hoy Collection #60: 4/3 [f] (ge e a y ai s o a e io a a c e se ae; wi a ai o 3rd, l[m]2nd,pupa. 1 Garden prepupa, Valley, nr Wairoa o g ege i a se ae me ia y o ege i a segme s II Gorge, K nz r d [as pt p r ], 19 Dec VI, a o ges - 5 μm o g; um e o se ae 1968, J.A. de Boer, #470: 1/2[ff] ad. Eves Bush, me ia y o eac segme II- I o g se ae; o g p n f l t [as C th d f l t ], 10 se ae a - s o se ae; I -II s o se ae; wi 1 se a May 1973 A e oe 1 19Β 1/ 2[m]2nd. MC: New us a e io o eac me a- a mesoco a; a 1- se ae Brighton, pt p r sp., 24 May 1915, G. Brittin, us os e io o eac oco a; e g o se ae associa e #71: 1/1[f] ad. wi eac oco a -9 μm; wi 1 su ma gi a se a o eac si e e wee s igma ic a eas; wi - ai s o i e - Remarks. Adult females of C. rn t ll and C. rn t are a e a se ae A e ae -segme e ; 3 segme o y very similar and share: s ig y s o e a o e 5 segme s oge e wi (i) the same distribution of reticulation areas on the seu osegme s; o a e g 1 - 3 μm; e g o dorsum; a ica se a a ou 3 μm e g o c y eo a a s ie (ii) the same basic dorsal pore distribution in the 13 -153 μm Wi o s i acu a e i emes a e io 37- reticulation lines; n f l nd 4 97 (iii) the tubular pores on the dorsum (although of very Ge us C E OC I O Maske different size in the 2 species); (iv) the submarginal band of simple pores on the venter; Ct n h t n Maskell, 1879: 208; —Atkinson, 1886: 277 (v) the very long stigmatic spines. [taxonomy]; -Maskell, 1887: 62, 65 [description]; — C. rn t ll can be separated from C. rn t by: Maskell, 1895a: 12 [checklist]; —Cockerell, 1894b: 1053 (i) its small size; [distribution]; —Cockerell, 1896: 330 [checklist]; — Cockerell, 1899a: 394 [checklist]; —Cockerell, 1899b: 16 (ii) the long tubular macropores (or largest dorsal pores) [mention]; -Cockerell, 1899c: 332 [key]; -Farquhar, (macropores convex and bollard-shaped on C. 1900: 247 [zoogeography]; —Fernald, 1903: 159 [world rn t catalogue]; -Hutton, 1904: 126 [checklist]; -MacGillivray, (iii) 8 anal ring setae (6 setae on C. rn t 1921: 171, 178 [catalogue]; —Froggatt, 1921, 22 (iv) only single pairs of long pregenital setae in segments [distribution]; —Morrison & Morrison, 1922: 71 V-VII (all setae long on segments VI and VII on C. [redescription of type]; —Steinweden, 1929: 234 rn t . [classification]; —Bodenheimer, 1935: 243 [zoogeography]; As indicated in the remarks under C. rn t b v , the —Balachowsky, 1936: 124 [French sp.]; —Archangelskaya, material ff pt p r sp. mentioned by both Maskell 1937: 35, 37 [mention]; -Lindinger, 1937: 182 (1885) and Hoy (1961) almost certainly refers to this [checklist]; —Brittin, 1940a: 412 [mention]; —Takahashi, species. The material from Dargaville (ND) collected by 1942: 27, 29 [Asian sp.]; —Lindinger, 1943: 218 Hoy in 1954 and that collected from Onekaka and [mention]; -Balachowsky, 1948: 255 [key]; -Borchsenius, Collingwood (NN) by Sewell in 1948 are also probably of 1957: 48, 271 [mention]; -Wise, 1977: 104 [checklist]; - this species. Ben-Dov, 1993: 99 [world catalogue]. Biology. Adult female C. rn t ll are generally found on Cnetochiton of a ac owsky 193 36 [miss e i g] the twigs of K nz , pt p r , and p n species. Adult females have been collected in the spring Type species: Ct n h t n v r d Maskell (designated by (August—December) and autumn (April—May) and adult Fernald, 1903: 159) males in August and March and so C. rn t ll may be bivoltine; apparently overwinters as the 1st- and 2nd- Diagnosis. Adult female. Test: glassy, flat, thin, and instar nymphs. The empty glassy tests of the males persist i e ya i e; s owi g e icu a e a e ( igs Μ Μ1 on the twigs through the summer and autumn and can often internal wax structure of plate on C. v r d . be found when no other life stages are apparent on the host. Shape: mature adults mostly large, up to 10 mm or more in length, pyriform, flat to slightly convex, with a Distribution. Throughout areas with a drier climate from shallow anal cleft. Colour green. Northland to Mid-Canterbury (Map 15). DorSum: derm membranous. Dorsal setae absent. o sa o es i a e icu a e a e ( igs Μ M1 o C. p r v r d , delineating reticulation areas in 7 longitudi- Name derivation. L rn , rn t , to decorate; rn t ll , small ornate scale similar to rn t . nal rows across body, 9 reticulation areas between anal plates and anterior margin and with 26 or 28 areas around margin. Dorsal pores of 2 to 4 types: (i) minute, dark microductules, each with a long inner filament with a small balloon-like proximal end; (ii) simple pores of various sizes: mostly within lines of reticulation but also occasionally present medially on plates; (iii) larger convex, simple pores, apparently similar in structure to macropores: forming a distinct submarginal band (except on C. v r d and (iv) macropores, quite large and rather flat (`button-shaped'): most abundant in median 3 reticulation lines, becoming less frequent laterally and absent near margins (absent on C. v r d . Preopercular pores, dorsal tubercles, and dorsal tubular ducts absent. Anal plates widest on anterior third, each plate with 4 setae: 2 stout setae along inner margin, one finer, longer seta ± apically and another spinose seta posteriorly on 98 d n & nd r n (2000 : C d (In t : pt r : C d upper surface near apex; without a sclerotised collar ema ks. This genus contains four species: Ct n h t n around anterior margin of anal plates on dorsum. h l n Henderson & Hodgson . s ., C. p r v r d Anogenital fold with 2 large sclerotised plates arising Henderson & Hodgson . s ., C. t r Henderson & internally and extending anteriorly from anterior margin; Hodgson . s ., and C. v r d Maskell. The main with 2-4 pairs of long setae along anterior margin and with diagnostic characters are: a single seta on each lateral margin. Anal ring with 3 pairs (i) their large size and generally pyriform shape; of setae. (ii) the distribution of the pregenital disc-pores across Ma gi : marginal setae abundant, broadly conical, most abdominal segments, and the oval, parallel-sided strongly spinose (except finely spinose on C. v r d , shape of the inner loculus; rather blunt, shape of basal sockets rather variable within (iii) the large size of the spiracles, subequal to the size of genus; in a single line not extending up margins of anal the coxae; cleft; reticulation setae usually larger; setae on anal lobe (iv) the displacement of the mouthparts to nearer one not differentiated from other marginal setae. Stigmatic procoxa; clefts shallow (particularly on C. t r but distinct; (v) the distribution of the ventral tubular ducts, when without stigmatic sclerotisations; each cleft with a single present; spinose, rather short, stout stigmatic spine, never longer (vi) the rather irregular distribution of the marginal setae. than 3 x length of marginal setae. Eyespot present. Species in the genus Ct n h t n are similar to those Venter: pregenital disc-pores with mainly 10 (range in Cr t ll t t and K l r in having abundant pre- 6-10) outer loculi and a single oval inner loculus, usually genital disc-pores medially across the abdominal with distinctive parallel sides; distributed across most or segments, each with mainly 8-10 loculi. Species in both all abdominal segments medially and in groups on Cr t ll t t and K l r differ from those in mediolateral folds of most segments. Spiracular disc- Ct n h t n in having: pores with mainly 5 loculi (range 3-7), in fairly narrow to (i) 2 types of ventral tubular duct; very broad bands between spiracles and margin, each band (ii) small spiracles; often broadening along margins of stigmatic cleft; with (iii) abundant spinose marginal setae approximately few or none extending medially past peritremes. One or 2 evenly spaced around margin; preantennal pores present near each scape. Ventral (iv)large simple pores on the dorsum, generally associated microducts: distal apex of inner ductule with 3-4 finger- with the anal plates. like lobes: sparsely distributed in broad marginal band and abundant medially on head, thorax, and more anterior Cr t ll t t species also differ in having ventral abdominal segments. Ventral tubular ducts of 1 type (2 tubular ducts which are: present medially on the thorax; types on C. h l n , terminal glands on distal ends on (i) inner ductules rather spiky: distribution highly variable (ii) in broad submarginal bands, with the ducts not tending between species and generally diagnostic (sometimes to lie radially. K l r species have ventral tubular absent on C. v r d . Ventral setae: hypopygial setae ducts in complete, relatively narrow submarginal absent; with a pair of long pregenital setae on abdominal bands, with the ducts tending to lie radially. segments VII and VI only; generally with groups of small Ben-Dov (1993) included thirteen non-New Zealand setae present on either side of posterior spiracular disc- species in the genus Ct n h t n Maskell. The senior pore band (absent on C. v r d other setae distributed as author has seen many of these and none appear to be for family. Antennae well developed, 6-segmented, 3rd congeneric with C. v r d . Indeed, most do not even segment much longest, often subequal to other 5 segments, belong to the same subfamily! As species in the genus often showing some constriction between segments II and Ct n h t n appear to be endemic to New Zealand, those III; setal distribution as for family. Mouthparts generally from other parts of the world will have to be transferred distinctly displaced to one side. Spiracles large, width of either to new or to other existing coccid genera. peritremes usually longer than length of coxae. Legs well developed but rather small for size of mature female, each with a separate tibia and tarsus but no tibio-tarsal articulatory sclerosis; tarsal campaniform pores absent; distribution of leg setae as for family; claws small and short, without a denticle; claw digitules similar and broad; tarsal digitules dissimilar, one shorter and narrower than other. Vulva opening on segment VII. n f l nd 4 . . Ct n h t n h l n e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . 00 d n & nd r n (2000 : C d (In t : pt r : C d . 2. Ct n h t n p r v r d e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . n f Ν Ζ l nd 4 0 ig. . Ct n h t n t r e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . 02 d n & nd r n (2000 : C d (In t : pt r : C d ig. 4. Ct n h t n v r d Maske , a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . Α s i ac e. n f l nd 4 0 Key o a u ema e Ct n h t n composed of thin glassy wax plates in a reticulate pattern Ventral tubular ducts extremely scarce, 1-2 sometimes of rows, with 7 rows of wax plates across abdomen and 9 present mediolaterally on posterior abdominal plates between anal plates and anterior margin; teneral segments; dorsal reticulation lines without macropores, female with a narrow marginal fringe of pointed wax plates replaced by simple pores subequal in size to associated similar to that of C. paraviridis, which tend to disappear as microductule pores; marginal row of medium to large insect grows. simple pores absent; spiracular disc-pore bands fairly Mounted material: body shape as above, round-oval to narrow throughout. viridis pyriform, slightly narrower at head end; length 2.62-6.30 —Ventral tubular ducts present in a submarginal band mm, breadth 1.94-5.40 mm. (although sometimes absent on C. paraviridis); dorsal reticulation lines with macropores, these much larger Dorsum: dorsal pores distributed in a reticulate than associated microductule pores; with a complete pattern as for genus, but with 26 reticulation areas around row of medium to large simple pores close to marginal margin; dorsal pores of 3 types: (i) microductules with spines; posterior spiracular disc-pore bands generally dark oval pore, about half size of smaller disc-pores: broadening near margin 2 present as 2 border rows along reticulation lines and in a submarginal row; (ii) simple pores, variable in size: small 2 Ventral tubular ducts forming a sparse submarginal band pores about twice size of microductule pore, occasionally mainly close to margin; ventral tubular ducts absent on with a few slightly larger simple pores: within reticulation either side of mouthparts toru lines between macropores, and with a few scattered within plate areas; medium-sized simple pores also present —Ventral tubular ducts either forming a clear, dense within plate areas and in a row on submargin, but not in a submarginal band or with only a few ducts scattered marginal row along anal cleft (those present part of may from margin; ventral tubular ducts usually reticulation lines); and (iii) large, button-shaped present as a group between mouthparts and procoxae macropores with a sclerotised rim and granular centre (rarely absent on paraviridis) 3 ( ig Μ o e wi a sma c ea o i mi e; su equa i si e o ege i a isc- o es; wi 1 -1 i Ventral tubular ducts usually forming a clear, broad os e io me ia mac o o e i e is i e o e wi a ga submarginal band (occasionally in a scattered band); me ia y w e e mac o o es a se A a a es e g dorsal macropores almost as large as pregenital disc- 177- 5 μm com i e wi s 15 - 3 μm; eac a e pores on venter; with 4 pairs of setae along anterior wi - mi u e o es o u e su ace; se ae o i e a margin of anogenital fold chelyon ou e ma gi s s o a s a y s i ose 1 - μm o g —Ventral tubular ducts scattered or absent submargin- a ica se ae u o ea y 3 o ge 33- μm o g i e y ally; dorsal macropores of medium size, smaller than s i ose- age a e A oge i a o wi 3- ai s o se ae pregenital disc-pores on venter; with 2 pairs of setae a o g a e io ma gi o ges a ou - μm along anterior margin of anogenital fold .. paraviridis Margin: marginal setae moderately spinose to near- conical with blunt tips, in staggered groups, with 10-26 on each side between stigmatic clefts; size rather variable, 15-3 μm o g a ge s i es mo e o icea e a Ct n h t n h l n e e so & o gso ew e icu a io oi s o a ome S igma ic s i es a e s ecies s o eac 53-73 μm o g Venter: pregenital disc-pores present medially across igs Μ1 Μ C5 111 posterior 5-6 abdominal segments and in small groups on Unmounted material: large, up to 10 mm long, green, mediolateral lobes: number of disc-pores on each segment with a reticulate pattern in double lines of slightly paler (medially/mediolaterally on one side): anal cleft+VII, 25- green and blue green, and with an irregular-shaped patch 40/24-45; VI, 54-72/11-21; V, 62-106/7-15; IV, 48-95/ in centre of each wax plate area bounded by same coloured 0-12; III, 20-56/0-8; and II, 0-6/0. Spiracular disc-pores double lines; body shape usually acuminate-oval and with 40-74 in anterior bands and 48-122 in posterior sometimes asymmetrical when adpressed to main leaf bands; each band 2-6 disc-pores wide near anterior vein; usually convexly rounded on dorsum and ventrally stigmatic spines and 4-10 disc-pores wide submarginally nearly flat. Size increase to maturity can beat least 12-fold near posterior stigmatic spines. Ventral microducts as for before oviposition begins, when median underside of genus. Tubular ducts of 2 types, a larger type in a broad abdomen gradually retracts to form a brood chamber. Test band on inner submargin, numbering 147-890 (mean 350) 104 d n & nd r n (2000 : C d (In t : pt r : C d ducts, and on head by mouth, and with 0-5 smaller ducts ad, 5[f] 3rd, 5[f] 2nd, 7[m] 2nd, 3 pupae,[mm]ad. Waiaroho, mediolaterally on abdomen and on either side of anal cleft; Str bl h t r ph ll , 26 Jan 1993, 29 Sept 1993 & 3 absent medially on thorax. Ventral setae: ventral anal lobe Nov 1993, RCH, #93-076a-f, #93-323a-b & #93-342a-f: se ae 3 - 1 μm o g; wi a ai o o g ege i a se ae 12/9[ff] ad, 4[f] 3rd (2 pharate), 3[f] 2nd, 9 crawlers,8[m] o a omi a segme s I a II o ges 5- μm; 2nd, 2 pupae, 3[mm] ad. Te Araroa beach north end, wi - ai s o a e io a a c e se ae; um e s o se ae C r n rp l v t , 31 Oct 1994, RCH, #94-132a- me ia y o a omi a segme s II 3-7; I -7; 1- h: 8/3[ff] ad (1 pharate), 7[f] 3rd, 10[m] 2nd, 4 pupae. GB: 7; I -7; III -5; a II 1- ; wi - se ae ea eac Karakatuwhero V Rd, Waipiata, M l p pl x under- me aco a -5 ea eac mesoco a a - ea eac side leaves, 28 Sept 1993 & 4 Nov 1993, RCH, #93-317b- oco a; wi 11-3 se ae o ei e si e a wi i eac e & #93-348b-d: 7/1[f] ad, 5[f] 3rd (2 pharate), 2[f] 2nd, os e io s i acu a isc- o e a ; wi -1 i e a e a 13[m] 2nd, 1 crawler, 1 prepupa, 2 pupae, 1 [m] ad. Paoneone, se ae; wi 3-5 su ma gi a se ae o ei e si e e wee Cl t nn n h , underside leaves, 2 Nov 1994, s igma ic c e s A e ae -segme e o a e g 3 7- RCH, #94-103a-e: 5/6 [ff] ad, 2[f] 3rd. Hexton, Grays Bush, 7 μm; a ica se a 3 - 5 μm o g C y eo a a s ie Str bl h t r ph ll young leaves, 25 Nov 1993, J.S. 173- 3 μm o g S i ac es wi o e i emes a e io Dugdale, #94-001: 1/1 [f] ad. WN: Wellington, t x l n 73-1 3 μm wi e os e io 9 -1 1 μm wi e egs e g leaves, 16 Jan 1981, J. Campbell, #81-40a: 7/7 [ff] ad. (me a o acic co a 119-1 μm; oc a e + emu 1 -1 μm; i ia 1 -1 μm; a sus 53-73 μm; c aw Remarks. C. h l n is very similar to C. p r v r d in 1 - μm o g having: (i) a marginal row of dorsal medium-sized simple pores; Material examined: HOLOTYPE [f]: NEW ZEALAND: (ii) nearly conical marginal spines; BP: Waiaroho, 3 Nov 1993, R.C. Henderson, Str bl (iii) ventral tubular ducts near the mouth; h t r ph ll new young leaves, #93-342a, NZAC: 1/1[f] (iv) in the shape of the dorsal macropores. ad. However, on C. h l n the macropores are larger, the PARATYPES: collection data as for holotype, NZAC #93- ventral tubular ducts are always present in a submarginal 342b-f: 5/3[ff] ad, 3[mm] ad, 2 pupae. band, often in Iarge numbers (they are rarely like this on C. Other material: New Zealand (Hawkes Bay), labelled p r v r d , and the posterior spiracular pore bands on C. "Ct n h t n d pr , [misidentification] from h l n widen substantially near the margin. C. t r also l nth , adult female, May 1883, W.M.M."; CMNZ: has spiracular bands which widen significantly near the 1/1[f] ad. [previously included as a paralectotype for margin but has no tubular ducts near the mouth and the K l r d pr ]. ΤΗ: Three Kings Islands, Southwest ventral tubular ducts are nearer the margin. C. v r d can I, C r n rp l v t leaves, 26 Nov 1983, J.C. Watt, be separated from C. h l n in: #83-340a: 6/6[ff] ad. As previous, but A. Wright, #83- (i) having slender marginal spines; 346e, 3/3[ff] ad. Three Kings Islands, West I, El n t (ii) having narrow spiracular pore bands; sp., 28 Nov 1983 & 29 Nov 1983, J.C. Watt, #83-346c & (iii) lacking dorsal macropores; 83-347f: 2/2[ff] ad. AK: "Ct n h t n v r d Mask., (iv) lacking a marginal row of simple pores; Kotinsky Coll. #282, on Ct h p r v r d [unknown ( lacking ventral tubular ducts near the mouth and plant name], Auckland, NZ, Koebele (1909) Nov. 23 1899" submarginally. USNM: 1/1[f] ad. "Ct n h t n v r d , Kot. Coll. #24, The three USNM slides listed above-originally Auckland, NZ, Koebele (1914), Nov. 27 1899", USNM: identified as C. v r d r here considered to be C. 1/1[f] ad. "Ct n h t n v r d Mask., Auckland, New h l n. Zealand, Geo. Compere Coll. #713", USNM: 1/2[ff] ad (incomplete). Owairaka, C r n rp l v t [as Biology. Similar to C. p r v r d . l v t ] leaf, Dec 1948, K.P. Lamb: 1/1[f] ad (poor). Pathogens and parasitoids. Hymenopterous parasitoid Auckland, St Heliers, C r n rp l v t , 23 Jan recorded: Pteromalidae: Aph b t ll Howard. 1977, L.L. Deitz, #77-39a: 8/8[ff] ad. Auckland Univer- sity grounds, C pr sp. leaf, 10 Oct 1980, #80-291c: Distribution. Known from Three Kings Islands and the North Island (Map 16). 3/3 [ff] ad. BP: Murupara, Motumoku Bush C1230, t l r , 4 Nov 1959, N.O. Secombe, FRNZ R(a)4-10, Name derivation. The specific name h l n is taken 12-13: 13/11[ff] ad, 3[m] 2nd,[mm]ad.1 Lottin Point, from h l n (Gr. n.): tortoise-shell: patterned and shaped Otanga, t x l n , 27 Jan 1993, 29 Sept 1993 & 3 Nov like a turtle, referring to the pattern of colourful plates on 1993, RCH, #93-276a-d, #93-330a,c, #93-350a-f: 12/6[ff] the dorsum. n f l nd 4 0 Ct n h t n p r v r d e e so & o gso ew Venter: pregenital disc-pores as for genus: number s ecies per segment (medially/mediolaterally on one side) laterally: anal cleft+VIl 44-62; VI, 48-53/8-10; V, 27- Figs Μ Μ1 C53-5 C5 -57 C 9-71 11 60/4-13; IV, 29-53/2-5; III, 7-26/1-2; and II, 0-2/0. Spiracular disc-pores with mostly 5 (range 3-5) loculi; Unmounted material: large, up to 10 mm long, slightly with 11-38 in each anterior band and 16-53 in each convex on dorsum but more convex on venter, lying in a posterior band; anterior band only slightly broader near depression on underside of leaves, most often adpressed to margin but posterior band broadening more obviously. main leaf vein and consequently asymmetrical, or near a Ventral microducts as for genus. Ventral tubular ducts subsidiary vein, when not so asymmetrical; generally sometimes scattered in a submarginal band (but with none green, sometimes with blue-green spots and sometimes close to margin), numbering 0-180 (proportion of with a dorsal reticulate pattern in a single different shade of specimens with 0-5 ducts: 80%; 6-50 ducts: 10%o; 51-180 green; test glassy, smooth, made up of a series of wax ducts: 10%), also in a group of 0-55 near mouthparts, plates, with 7 longitudinal rows of plates across body, 9 usually with more ducts on compressed side when plates between anal plates and anterior end and with a specimen asymmetrical. Ventral setae: ventral anal lobe marginal fringe of rather pointed wax plates. se ae 3 -57 μm o g; wi a ai o o g ege i a se ae o a omi a segme s II a I o ges 1- 5 μm Mounted material: body becoming more oval at maturity; o g; wi 1- ai s o a e io a a c e se ae; eque cy specimens taken from near a main leaf vein asymmetrical, o se ae me ia y o a omi a segme s II - ; I - with mouthparts twisted towards one side; specimens from ; - ; I 3-5; III 1-3; a II - ; wi 1-5 se ae other leaf sites usually more symmetrical; anal cleft about ea eac me aco a - ea eac mesoco a a -3 ea 1/5th body length; length 2.90-7.85 mm, breadth 2.35- eac oco a; wi g ou s o 7- 3 se ae a o g ma gi s o 6.50 mm. os e io s i acu a isc- o e a s; wi -7 i e a e a se ae; wi - su ma gi a se ae e wee s igma ic c e s Dorsum: derm membranous except at maturity when o ei e si e A e ae -segme e wi ou reticulation lines become sclerotised. Dorsal pores seu osegme s; o a e g 3 -37 μm; e g o is i u e i a e icu a e a e ( igs Μ Μ1 as o a ica se a 3 -73 μm C y eo a a s ie 15 -1 μm ge us u wi e icu a io a eas a ou ma gi ; o sa o g S i ac es wi o e i emes a e io 5 -1 μm; o es o 3 y es (i mic o uc u es i a ow o ei e si e os e io 9-115 μm egs e g (me a o acic co a o e icu a io i es a a o g su ma gi ; (ii sim e 1 -119 μm; oc a e + emu 13 -173 μm; i ia - o es a sma es sca ce; s ig y a ge o es ese 173 μm; a sus 1-9 μm; c aw 15- μm wi i e icu a io a eas a e icu a io i es; a ges i a ow e wee a esi e ma gi a s i es; a (iii Material examined: HOLOTYPE [f]: NEW ZEALAND: mac o o es mo e a e y a ge u sma e a ege i a AK: Waitakere Range, e ow Track, S h ffl r d t t isc- o es u o -s a e wi a ea i y sc e o ise ou e u e si e ea es 1 Oc 1997 Α Ma i AC 97- i g a a i e a k g a u a su ace ese i me ia 1 1/1[ ] a a su me ia e icu a io i es wi 1 - 1 i com e e PARATYPES: collection data as for holotype, NZAC #97- os e io me ia mac o o e i e A a a es wi es a l44a, c-l: 11/13[ff] ad (1 pharate), 2[f] 3rd instars. a e io i a e i g o a e oge e o a ; e g 13 - 15 μm com i e wi s 13 - 3 μm; eac a e wi 1- Other material: "Ct n h t n v r d Mask., New Zealand, 3 mi u e o es o u e su ace; i e ma gi se ae s a ex coll. W.M. Maskell, Newstead Coll/1945-121, Cotype a s i ose e g 1 - μm; a ica se ae s e e lot", USNM: 1/1[f] ad (split dorsoventrally). "Ct n h t n s i ose a usua y a ou wice as o g as i e ai s v r d Mask., New Zealand, Mask. Coll. #33", USNM: 1/ e g 3 - 7 μm; ou e ma gi se ae s ou s a - oi e 1 [f] ad.AK:RangitotoGr l n I, Kidney Fern Glen, s i es e g 1-3 μm A oge i a o wi 3 ai s o l d , 12 Sept 1976, G.Hall, 76-274a: 4/11[ff] ad, 39 se ae o ges -57 μm 2nd. Auckland, Titirangi, d r rb r leaves, 11 Margin: marginal spines with blunt tips; with 7-24 on Jan 1983, J. Cox: 1/1[f] ad. Waitakere Ra, Fairy Falls Tk, each side between stigmatic spines; size rather variable, d r rb r leaves, 4 Dec 1979, M.F. Tocker, #79- 15-3 μm o g sma e s i es ai y a ow a ge 347c: 4/4[ff] ad. Waitakere Ra, Rangemore Tk, east end, e icu a io s i es oa a a mos co ica mo e S h ffl r d t t underside leaves, 26 Dec 1994, R.E. o ious a o g ma gi s o a ome a o ce a o- Beever, #95-005a-c, -006a-b, -007: 6/6[ff] ad. Waitakere o a S igma ic s i es 3 - μm o g Ra, Sharp Bush, d r rb r underside leaves, 13 06 d n & nd r n (2000 : C d (In t : pt r : C d Jul 1997, RCH, #97-082a-b: 2/1[f] 2nd, 3[m]2nd. As pre- N.Z., from Hedycarya arborea, Jan 1921, coll. J.G. Myers, vious, except 6 Oct 1997, #97-140: 1/1[m] 2nd. Paparata ΒΜ 19 1 " ΒΜΝΗ; 1/ [ ] a We i g o Rd, neighbouring property to P.S. Dale, Hedycarya Pseudowintera axillaris, 26 Feb 1980, S. Norton, #80- arborea, 26 Feb 1995, RCH, #95-025a-b: 2/2[ff] ad. As 71 / [ ] a Aka a awa Sa e Ν 1 1 7 Schefflera previous, but on property of P.S. Dale, #95-026a-b: 2/ digitata, 2 Feb 1976, M.A. Stoodley: 1/19 ad. SD: Opouri 3[ff] ad. As previous except 1 Feb 1997, #97-032a--j, & Valley, Marlborough, Pseudowintera axillaris, 10 Jan #97-033a-j: 40/20[ff] ad (split dorsoventrally). BP: Te 1969, J.S. Dugdale, No. 477: 2/29 9 ad (one split dors- Koau, 360 m, Hedycarya arborea, 26 Oct 1992, J.S. oventrally). NN: Motueka, Hedycarya arborea, Dec 1937, Dugdale, #92-339a-b: 2/4[ff] ad. Te Koau, Hedycarya G.Brittin, No.8, USNM: 1/1[f] ad, & 29 Dec 1937, NZAC: arborea, 31 Jan 1993, RCH, #93-077a-c & 4 Nov 1993, 2/2[ff] ad. Eves Bush, Pseudowintera axillaris, 9 Dec #93-355a-c: 5/6[ff] ad, 1[f] 3rd, 1[f] 2nd, 10 crawlers. Te 1970, J.A. de Boer, No. 671: 3/3[ff] ad (split dorsoven- Koau, Ripogonum scandens leaves, 4 Nov 1993, RCH, trally). Upper Maitai R, Dad's Creek, 450m, Griselinia #93-354a-e & Bushwalk tk, cnr to Lookout tk, 31 Oct littoralis (broadleaf), 20 Oct 1963, E.S. Gourlay, #83-298b 1994, #94-l33a--h: 13/10[ff] ad (1 pharate), 2[f] 3rd, 1[f] & #90-221a--f: 7/7[ff] ad. Maitai R, Dodonaea viscosa, 2nd, 7[m] 2nd, 2 1st, 2 pupae, 2[m] ad. Lottin Point, Otanga, 17 Jan 1968, J.A. de Boer, No. 341: 1/2[ff] ad (split dors - Hedycarya arborea new young leaves, 29 Sept 1993, RCH, oventrally). Sherry Valley, `from leaves', 4 Feb 1942, (no #93-327a-c: 3/2 [ff] (pharate), 4[f] 3rd,[m]2nd,2 & #93- coll.), No. 687, #83-304c & #90-229a-e: 10/10[ff] ad. 329: 1/2[m] 2nd, 2 prepupae, 2 pupae. As previous, except Golden Downs, Pseudowintera axillaris [as Drimys], 9 undersurfaces new leaves, 3 Nov 1993, #93-343a-f: 6/5[ff] Mar 1961, J.G.R. McBurney, FRNZ No. R11, 1/2[ff] ad, ad, 1[f] 3rd. Waiaroho, Hedycarya arborea new shoots & & Νo 19 1s Ma s e a ey Hedycarya arborea, leaves, 3 Nov 1993, RCH, #93-341d-e: 2/2[ff] ad. Orete 16 Apr 1969, E.W. Valentine, No. 508: 2/3[ff] ad (split Forest, Te Puia Hut, Myrsine salicina, 25 Jan 1993, RCH, dorsoventrally). Motupiko, Griselinia littoralis, 19 Dec #93-215a-b: 2/2[ff] ad (split dorsoventrally). Rereauira 1967, E.W. Valentine, No, 304: 2/2[ff] ad (split dorsoven- Swamp, Beech Ridge, Myrsine australis, 29 Jan 1993, trally). BR: Maruia Springs, Peraxilla colensoi, 8 Jan RCH, #93-079a-e & 29 Sept 1993, #93-324a-c: 6/6[ff] 1976, B.P.J. Molloy, (submitted by Somerfield): 2/2[ff] ad, 13[f] 3rd, 1 [m]2nd.Griselinia Whakarewarewa (F.R.I.), ad. Alfred R, L Daniells Tk, Griselinia littoralis under- littoralis ea es o 19 o ag ΝΖ (a 55- side leaves, 12 Jan 1994, RCH, #94-016a-b: 2/2[ff] ad. 57; (a - 9 3/13[ ] a ; /7[mm] a G Kaka ui Fletchers Creek, West Inangahua, Reefton, Pseudowintera 5 m Pseudowintera axillaris, 1 Feb 1993, RCH, #93- colorata, 7 Mar 1972, J.A.K. Farrell, No. 816: 2/2[ff] ad 275a-b: 2/1[f] ad (split dorsoventrally), 10 1st. Kakanui, (split dorsoventrally). Springs Junction, Palmer Road, 300m, Myrsine australis, 1 Feb 1993, RCH, #93-280a-e Peraxilla colensoi new shoots, 4 Nov 1993, J.S. Dugdale, & 16 Mar 1993, #93-080: 6/6[ff] ad, 1[f] 2nd, 1[m] 2nd, 8 #93-330Ba-e: 5/2[ff] ad (one pharate), 4[f] 3rd, l[m]2nd, 2 1st. Paoneone, Hedycarya arborea leaves, 2 Nov 1994, pupae, 7[mm] ad. Charleston - Addison Flat, Myrsine RCH, #94-126: 1/3[ff] ad, 1[f] 3rd. Pohutu, Awatere R salicina, 7 Nov 1972, J.A. de Boer, No. 936: 1/3[ff] ad (2 bridge, Hedycarya arborea underside leaves, 4 Nov 1993, split dorsoventrally). NC: Wandle R, Alepis [as Peraxilla] RCH, #93-357, 1 Nov 1994, #95-022a-j, 3 Nov 1995, flavida, 9 May 1973, B.P.J. Molloy, No. 1023: 1/2[ff] ad #98-060a-b: 13/17[ff] ad, 2[f] 3rd,TO:[m]2nd,3 1 pupa. (one split dorsoventrally). MC: Banks Peninsula, Akaroa, Hauhungaroa Ra, 650m, Waihora, Griselinia lucida leaves, Griselinia littoralis, 16 Jan 1972, R.C. Close, No. 788: 2/ 11 Jan 1995, RCH, #95-015a-e: 5/5[ff] ad. Rangitoto Sta- 2[ff] ad (split dorsoventrally). Akaroa, Pseudopanax tion, Mangatutu, Pseudowintera axillaris new shoots, 9 arboreus [as Nothopanax], 16 Jan 1972, R.C. Close, No. Nov 1996, RCH, #96-236a-b: 2/9[f] 3rd. As previous, 787: 4/4[ff] ad (split dorsoventrally). Banks Peninsula, except Hedycarya arborea new shoots, #98-059: 1/1[f] ad. Otanerito Reserve, Pennantia corymbosa leaf, 14 Jan 1994, HB: Black Birch Ra, Kaweka Rd, Griselinia littoralis un- RCH, #94-015: 1/1[f] ad. DN: "Ctenochiton viridis Mask., derside leaves, 14 Mar 1998, RCH, #98-054a-b: 2/1[f] ad, on Griselinia littoralis, Oamaru, N.Z., May 24, 1914, 3[m] 2nd, 1 1st.TK:Hoheriasp., e We a S Ν 3/15/75 Brittin no.8", USNM; 1/1[f] ad. "Ctenochiton viridis, 1 e 197 M A S oo ey ΝΖ ( 1- 3 3/7[ ] Oamaru, on Griselinia littoralis, G. Brittin, BM 1945 121, a RI: S ua i es Wes amaki a ey Ν/13 /5 / 9 Newstead Coll., stained Oct 25th 1916, RN, No. 20", Schefflera digitata leaves, (no date or collector), FRNZ BMNH; 1/1[f] ad. Ardgowan, Griselinia littoralis, 12 Jan R(b)43: 1/4[ff] ad (+ 9 of another unknown ?coccid). S. 1912, G. Brittin, No. 8: l/1[f] ad. FD: Breaksea Sound, Ruahine SF 3/04, West Tamaki Valley, Schefflera digitata Gilbert I 6, Hedycarya arborea, 2 Feb 1996, RCH, #96- o iage 1 ec 197 M A S oo ey ΝΖ ( -5 087: 1/1[f] ad. 7/1 [ ] a WN: "Ctenochiton viridis Mask., Wellington, n f l nd 4 0 Remarks. This species is highly variable and may be Post-oviposition females die and fall off the host, leaving displaying unstable phenotypic characters or may be a permanent depressions in the lamina of the leaf. complex of species. Amongst several collections on d r rb r an occasional specimen had large Pathogens and parasitoids. Hymenopterous parasitoids numbers of ventral submarginal tubular ducts and could recorded are: Encyrtidae: Ad l n rt d inconstans thus be keyed out to the closely related new species C. Noyes; Ad l n rt d sp. Pteromalidae: Aph b t h l n, whereas other specimens with the same ll Howard; Aph b t n n ( ouček I a i io collection data were invariably C. p r v r d . In contrast, C p r v r d is often attacked by several entomogenous adult females of C. h l n invariably exhibit a more fungal species, especially under damp conditions. The stable phenotype and in addition, there are no known two most common are A r t bb r H.S. Fawc., the populations of C. h l n n . rb r . The live "brown blobs" fungus, which usually infests early instars, appearance of adult females, particularly the lack of completely obliterating the shape of the insect, while multi-hued reticulate bands on the dorsum, can help rt ll l n (Zimm.) Viégas is a white fungus distinguish C. p r v r d from C. h l n. Individual C. generally killing adult females which retain their shape. p r v r d specimens without tubular ducts near the mouthparts may key out to C. t r (described as new Distribution. Recorded from Auckland in the north to below) but this latter species has very wide posterior Fiordland in the south (Map 17). spiracular disc-pore bands near the margin and has only been collected from r n t r , on which C. p r v r d Name derivation. From para (Gr.) meaning near and has never been found. v r d , the species it resembles described by W.M. C. p r v r d may be separated from C. h l n and Maskell. Although the taxonomic characters of these two C. t r in lacking the very wide posterior spiracular disc- species differ widely, they share a remarkably similar pore bands near the margin, and from C. v r d by: outward appearance and are both so-called "sixpenny (i) the broader marginal spines; scales". (ii) the presence of button-like macropores; (iii) a dorsal submarginal row of simple pores. The three USNM slides and two BMNH slides listed Ct n h t n t r e e so & o gso ew s ecies above that were originally identified as Ct n h t n Fig. 113 v r d are here considered to be C. p r v r d n. sp. The collection of C. p r v r d ff d p n x rb r Unmounted material: large, up to 8 mm long, elongate from Akaroa is unusual, as this is the only record of a and green; restricted to lower leaf surface of host plant Ct n h t n species other than C. v r d (which is almost r n t r usually sited with stylets inserted into main restricted to this genus) being recorded from it. leaf vein; a depression is caused in leaf beneath body with a consequent bump in upper leaf surface; test of glassy wax Biology. C. p r v r d is probably univoltine, with an plates covering dorsum. extended generation during the summer although it may occasionally have 2 generations. It overwinters as the 2nd- Mounted material: body elongate-oval, slightly narrower instar nymph. At the spring flush, the juvenile females at head end; length 2.6-7.3 mm, breadth 1.4-4.0 mm; anal migrate to the outside of the young shoots of their host cleft shallow, with margins meeting along their length; plant and quickly moult to the 3rd-instar. By the time the stigmatic areas not indented. leaves have fully unfurled, females have completed their final moult to become adult. The 2nd-instar males remain Dorsum: dorsal pores distributed in a reticulate on the old leaves and develop through prepupa and pupa to pattern as for genus, but with 28 reticulation areas around the adult male, emerging to mate with the adult females. margin; dorsal pores of 3 types: (i) microductules: present The preferred settling site for the adult females is by the mainly as 2 border rows alongside reticulation lines, also main leaf vein of new shoots, where they cause distinct with slightly larger pores in a submarginal row laterad to depressions in the lamina. Once settled, they are sessile, marginal row of simple pores; (ii) simple pores of various continuing to grow in size and eventually producing about sizes: small pores very few, present within reticulation 500 eggs. The eggs hatch into crawlers in the brood lines and reticulation areas; medium-sized pores present chamber formed beneath the abdomen and emerge from within middle row in reticulation lines between under the posterior end, where the wax test is loosened macropores; larger simple pores in a complete row near from the leaf substrate, and then settle on nearby leaves. marginal spines, but not along margins of anal cleft; and 08 d n & nd r n (2000 : C d (In t : pt r : C d (iii) macropores dense and button-shaped, with a dark Henderson, r n t r underside young lvs, NZAC, #95- sclerotised ring and granular centre under light 014d: 1/1[f] ad. microscope, smaller than ventral pregenital disc-pores: PARATYPES: same collection data, NZAC #95-014a—c: restricted to centre of reticulation lines, around median 3/3[ff] ad. and submedian reticulation areas from head to anal area, most abundant on abdomen; with 13-19 in complete Other material: BP: Rereauira Swamp, Beech Ridge, posterior medial macropore line. Anal plates: length 154- r n t r , 19 Sept 1992, RCH, #92-292; 26 Jan 1993, 1 μm com i e wi 13 -177 μm; eac a e wi 1- #93-107a-c & -#93-267a—d, and 19 Apr 1993, #93-266a— 3 minute pores on upper surface; inner margin setae sharp b: 10/5[ff] ad, 3[f] 3rd, 1[f] 2nd, 1 [f][m]2nd, 12 neonates/1st. a s i ose e g 1-33 μm; a ica se ae o ge a s e e s i ose e g 3 - 3 μm; ou e ma gi se a Remarks. C. t r is the least pyriform of the four species s ou y s i ose e g 1 -33 μm A oge i a o wi 3 currently placed in the genus Ct n h t n. In lacking ai s o se ae a o g a e io ma gi o ges 33- 7 μm tubular ducts medially near the mouthparts, it resembles C. Margin: marginal setae spinose to conical, with blunt v r d , but differs from C. v r d in having: tips; with 15-22 on each side between lateral stigmatic (i) a distinct submarginal band of ventral tubular ducts; c e s; si e a e a ia e 19-3 μm o g; usua y (ii) distinct dorsal macropores (absent on C. v r d distributed in groups of about 3 spines on abdomen at (iii) broad spiracular disc-pore bands, which become e icu a io oi s S igma ic s i es eac 3 -5 μm o g much wider near the margin (narrow on C. v r d Venter: pregenital disc-pores in rows across posterior (iv) posterior spiracular disc-pore bands with numerous 5 abdominal segments: numbers per abdominal segment small-medium size setae along margins (absent on C. (medially/mediolaterally on one side): anal cleft+VII, v r d . together 50-68; VI, 59-72/8-14; V, 53-83/5-10; IV, 50- C. t r also resembles C. h l n and C. p r v r d in 59/3-12; III, 2-9/5-6; and II, none. Spiracular disc-pores: having dorsal macropores and numerous small-medium with 27-48 in each anterior band and 41-80 in each setae along margins of posterior spiracular disc-pore posterior band; posterior bands broadening to 3-12 pores bands, but C. t r differs from them in: wide near stigmatic spine. Ventral microducts as for genus (i) more elongate body shape; but absent from a broad band submedially and also (ii) the absence of ventral tubular ducts near the mouth; medially on posterior abdominal segments. Ventral (iii) the distribution of the ventral tubular ducts in a band tubular ducts usually in a row about 2 ducts wide close to close to the margin. margin; absent from elsewhere on venter and sometimes absent altogether (total 0-119 ducts). Ventral setae: Biology. Probably very similar to the other three species e a a a o e se a 7-53 μm o g; wi a ai o o g of Ct n h t n. pregenital setae on both abdominal segments VI and VII, o ges -9 μm; usua y wi 1 ai o a e io a a c e Distribution. Only collected from two localities, both in setae; number of setae medially on abdominal segments: central North Island (Map 18). VII, 3-6; VI, 3-6; V, 3-6; IV, 3-7; III, 1-6; and II, 1-5; with 0-4 setae near each metacoxa, 0-3 near each Name derivation. From t r , the maori name for the host mesocoxa and 3-6 near each procoxa; with 17-27 setae plant ( r n t r . bordering and within posterior spiracular disc-pore bands, eac 7- μm o g; wi -1 i e a e a se ae; wi 3- 4 submarginal setae between stigmatic clefts on either side. Antennae 6-segmented, 3rd segment by far longest, Ct n hlt n v r d Maske wi o seu osegme s; o a e g 33 -3 μm; a ica se a - 1 μm o g C y eo a a s ie 1 -173 μm igs Μ Μ3 C55 11 o g Wi o s i acu a e i emes a e io 57- μm Ct n h t n v r d Maskell 1879: 211; -Maskell, 1885: os e io 73-1 μm egs e g (me a o acic co a 24 [male description]; -Maskell, 1887: 74 [description]; - 1 5-13 μm; oc a e + emu 135-1 μm; i ia 1 - Maskell, 1890a: 278 [biology]; —Koebele, 1893: 12 1 5 μm; a sus -77 μm; c aw 15- μm; c aw igi u es [parasitoid]; -Maskell, 1895a: 13 [checklist]; -Cockerell, usually small, rather narrow and short. Ι 9 33 [c eck is ]; - e a 19 3 159 1 1 [wo ca a ogue]; — u o 19 [c eck is ]; —Mo iso & Material examined: HOLOTYPE [f]: NEW ZEALAND: Mo iso 19 71 [ e esc i io ]; -Mye s 19 199 Ο au u ga oa a ge Wai a a 1 a 1995 C [c eck is ]; —Mi e 19 5 33 [ os ]; -G ee 19 9 n f l nd 4 0 377 [record]; —Steinweden, 1929: 234 [classification]; — Venter: pregenital disc-pores present medially across Green, 1930: 289 [mentions]; -Balachowsky, 1936: 124 posterior 4-6 abdominal segments, seldom extending onto [comparison French sp.]; —Lindinger, 193'7: 183 mediolateral lobes: number of disc-pores per segment [checklist]; —Wise, 1977: 104 [checklist]; —Deitz & (medially/mediolaterally on one side): anal cleft+VII, 0/ Tocker, 1980: 32 [checklist]; —Ben-Dov, 1993: 104 4-12; VI, 25-72/0-2; V, 26-107/0; IV, 32-120/0-2; III, [world catalogue]; —Hodgson, 1994a: 208 [redescription]. 21-54/0, and II, 0-41/0; and with 0-16 medially on metathorax and 0 laterad to metacoxae. Spiracular disc- Unmounted material: very large, up to 10 mm long, pores with mainly 5 (range 3-5) loculi, with 9-44 in each green, sometimes with slightly paler green markings in a anterior band and 22-48 in each posterior band; bands narrow, only 1-5 pores wide near stigmatic clefts. Ventral reticulate pattern, but never with blue-green spots as microducts as for genus but abundant in broad bands sometimes present on C. paraviridis; shape usually across median thorax and abdominal segments II—III, with elongate acuminate-oval and often asymmetrical when adpressed to main leaf vein; usually flat on dorsum and fewer on IV—V and none medially on VI-VII. Tubular ducts: with l-2 ducts present laterad to anal area on about convex ventrally, lying in a depression on leaf. Size increase to maturity can be at least 12-fold before half specimens examined. Ventral setae: ventral anal lobe oviposition begins, when median underside of abdomen se ae eac 3- μm o g; wi a ai o o g ege i a setae on both abdominal segments VI and VII, longest 77- gradually retracts to form a brood chamber. Test of thin g assy wa a es ( ig Μ3 i e a wa s uc u e i a 9 μm; wi -3 ai s a e io a a o e se ae; um e s o reticulate pattern of rows, with 7 plates across abdomen setae medially on abdominal segments: VII, 3-8; VI, 3-9; and 9 plates between anal plates and anterior margin, and V, 3-10; IV, 2-10; III, 1-6; and II, 0-8; with 0-3 setae with a narrow, non-pointed, marginal fringe of wax plates near each metacoxa, 0-2 near each mesocoxa and 0-4 near which tends to disappear as insect grows. each procoxa; with few or no small setae associated with posterior spiracular disc-pore bands; with 4-13 interantennal setae; with 3-8 rather small submarginal Mounted material: body shape as above, oval, slightly setae between stigmatic clefts on either side. Antennae 6- narrower at head end; length l.60-12.45 mm, breadth segme e o a e g 33 - 5 μm; a ica se a 3 -73 1 .13-4.70 mm. μm o g C y eo a a s ie 15 -19 μm o g Wi o s i acu a e i emes a e io -1 7 μm os e io 57- Dorsum: dorsal pores distributed in a reticulate 15 μm egs e g (me a o acic co a 1 7-15 μm; pattern as for genus, but with 28 reticulation areas around oc a e + emu 15 -1 μm; i ia -13 μm; a sus ma gi ; o sa o es o y es ( ig Μ (i 5 - 5 μm; c aw we e e o e 13-19 μm o g; c aw microductules with dark oval pore with a slit-like opening digitules narrowly expanded, subequal. across shorter side, sometimes appearing to be bilocular: in 2 border rows along reticulation lines only; and (ii) Material examined: LECTOTYPE [f]: NEW ZEALAND: small simple pores about same size as microductule pore here designated, Ctenochiton viridis, from `Panax', adult (occasionally a few slightly larger simple pores present): female, June 1878, W.M.M. / stained and remounted in restricted to central band of pores in each reticulation line; Canada balsam, Feb 1998, RCH, CMNZ: 1/1[f] ad. with 13-20 simple pores in posterior medial macropore PARALECTOTYPES: (i) "from Coprosma, skin of adult line; without a submarginal band of pores. Anal plates: female, June 1878, W.M.M." / stained and remounted in e g 13 - 19 μm com i e wi s 13 -1 μm; eac Canada balsam, 2 Feb 1995, RCH, CMNZ: 1/1[f] ad. (ii) plate with 0-4 minute pores on its upper surface; inner and "from Coprosma - five females - 2nd stage, June 1978, outer margin setae all fine and sharply spinose, lengths W.M.M" / stained and remounted in Canada balsam, 16 1 -1 μm; a ica se ae o ges e g -3 μm Feb 1995, RCH, CMNZ: 1/1[f] 2nd, 2[m] 2nd, 2 1st (i.e. not Anogenital fold generally with 2 pairs of setae but all females!). (iii) "from Rubus, old females, June 1977, sometimes with 3 setae along each side of anterior margin W M Μ " AC 1/ [ ] a mou e y w o e ( o a 1- se ae a e a y o ges a ou 7- μm o g parasitised and infested with fungus). (iv) from `Panax' Margin: marginal setae slender-spinose to moderately as above, CMNZ: 2/3[ff] ad mounted dry, whole (alI heavily s i ose wi oi e i s; si e a ia e 7- 3 μm o g infested with fungus). larger spines at reticulation points but also with 2-3 on ei e si e o s igma ic s i es 3-3 μm o g; wi 1 - Other material: Maskell subsequent slide, Ctenochiton on each side between stigmatic clefts. Stigmatic spines viridis, a u ma e 1 9 W M M CM 1/ [m] a ΤΗ s o eac 19-5 μm Three Kings Is, Great I, Pseudopanax lessonii, 1-3 Jan 0 d n & nd r n (2000 : C d (In t : pt r : C d 1963, E.S. Gourlay, #83-304a: 3/3[ff] ad. Great I, Meryta RCH, #95-016a-d: 4/4[ff] ad. NN: Rai Valley, C 2, sinclairii, 27 Nov 1983, C.F. Butcher, #83-349e, 2/2[ff] 300 ft, Marlboro' Prov., S68:90:38, Pseudopanax arboreus ad. West I, [no host], 29 Nov 1983, C.F. Butcher, #83- [as Neopanax arboreum] 23 Jan 1964, W.A. Holloway, 346h: 1/1 [ff] ad (and 2 parasitoids). ND: Te Arai Sanctu- FRNZ R(b)9-11: 3/10[ff] ad. Maitai R, Pseudopanax ary, Houhora, Pseudopanax lessonii hybrid, 27 Jan 1995, arboreus [as Neopanax arboreum], 19 Dec 1967, J.A. de R.E. Beever, #95-021 a-c: 3/3[ff] ad. AK: Waitakere Boer, No, 308: 2/3[ff] ad. Eves Valley, Pseudopanax Range, Sharp Bush, Pseudopanax arboreus leaf, 1 Dec arboreus [as Neopanax arboreum], 29 Feb 1972, J.A. de 1994, RCH, #94-122: 1/1[f] ad. As previous, but 30 Dec Boer, No. 804: 2/3[ff] ad (2 split dorsoventrally). As pre- 1994, #95-002 a-b, #95-003, #95-004:4/4[ff] ad. As pre- vious, but Pseudopanax crassifolium, No. 805: 1/1 [f] ad vious, but 29 Apr 1995, #95-043: 1/3[f] 2nd (2 pharate), 6 (split dorsoventrally). Golden Bay, nr Bainham, Mt 1st. As previous, but 11 Sept 1995, G. Hall & L.H. Clunie, Stevens Tk, Pseudopanax crassifolius juvenile leaf, 12 Mar #95-100a-d: 4/6[f] 3rd (3 pharate), 6[f] 2nd, 2[m] 2nd. As 199 Ν A Ma i 9 - 5 1/1[ ] a M e o us previous, but 23 Oct 1995, RCH, #95-110a-f: 6/4[ff] ad i ge Pseudopanax arboreus [as `Five Finger'], May (2 pharate), 12[f] 3rd. As previous, but 13 Jul 1997, #97- 1952, W. Cottier: 4/4[ff] ad. NC: Lees Valley Rd, near 079a-b: 2/7[f] 2nd, 2[m] 2nd. As previous, but 6 Oct 1997, Oxford, Pseudopanax ?linearis [as ?lineare], 4 Mar 1963, #97-139a-c: 3/1 [f] 2nd, 1 [m]2nd, 1 pupa, 1 [m] ad. CL: Little R.M.J. McKenzie: 2/4[ff] ad. Barrier I, Summit Tk, 243 m, Pseudopanax arboreus, 10 Mar 1974, J.C. Watt, #83-300fZ-243: 3/3[ff] ad. Little In addition, the identity of the following material from Barrier I, Hamilton Tk, Pseudopanax arboreus underside Maskell's syntype series remains unconfirmed: (i)"from leaves, 17 Sept 1994, RCH, #94-080a-d: 4/3[f] 3rd, 16[f] Rubus, old females, June 1877, W.M.M., 1/1[f] ad 2nd, 6 1st. BP: Te Koau, 130 m, Pseudopanax arboreus mounted dry, whole (NZAC)", because no other material new young shoots, 2 Nov 1993, RCH, #93-356a-c: 3/6[ff] is available from this host; (ii) the Maskell subsequent ad, 2[f] 2nd. Te Koau, 175 m, side track from Bush Walk slide labelled "Ctenochiton viridis, [no host], female early track, Pseudopanax arboreus new leaves, (one slide, from 3rd stage, Jan 1890, W.M.M. / stained and remounted in old leaves), 31 Oct 1994, RCH, #94-131 a-h: 8/9[f] ad (1 Canada balsam, 16 Feb 1995, RCH" (NZAC) is a pharate), 3[f] 3rd, 9[m] 2nd. Te Koau,Pseudopanax arboreus misidentification as it is probably Aphenochiton kamahi underside new leaves, 2 Nov 1995, RCH, #95-111 a-d: 4/ Henderson & Hodgson n. sp. 3[f]f ad. 3[f] 3rd. Rereauira Swamp, Pseudopanax arboreus, 29 Jan 1993, RCH, #93-285 a-e: 5/5[ff] ad. GB: Remarks. C. viridis and C. paraviridis are referred to as East Cape, Pseudopanax crassifolius, 3 Feb 1993, RCH, the "sixpenny scales" because of their size and generally #93-202 a-d: 4/4[ff] ad (3 split dorsoventrally). As previ- flat shape. Although the outward appearance of these two ous, but 17 Mar 1993, #93-081 a-d: 4/3[ff] ad (1 split scales is very similar, mounted females of C. viridis can be dorsoventrally), 11 1st. As previous, but 30 Oct 1994, old separated immediately by the lack of: leaves, #94-130a-d: 4/6[f] 3rd (l pharate), 9[f] 2nd, 4[m] 2nd, (i) dorsal macropores; 3 prepupae, 1 pupa. As previous, but Pseudopanax ferox (ii) a submarginal band of simple pores; old leaves & new shoots, #94-129a-e: 5/2[ff] ad (1 (iii) ventral tubular ducts on the submargin and by the pharate), 12[f] 3rd, 6[f] 2nd, 5[m] 2nd, 1 prepupa, 1 pupa. mouth; Paoneone, Pseudopanax arboreus underside leaves, 2 Nov (iv) the presence of fine marginal spines. 1994, RCH, #94-128a-c: 3/5[ff] ad, 1[f] 2nd. Pohutu, Apart from one unconfirmed record (Maskell's whole Pseudopanax arboreus, 30 Jan 1993, RCH, #93-078a-d: mount female "from Rubus"), Maskell's paralectotype 4/3[ff] ad, 13 1st. As previous, but off new shoots & leaves, from Coprosma sp. and the collection off Meryta sinclairii 4 Nov 1993,#93-349a-c: 3/6[ff] ad, 2[f] 3rd. WO: Pangaki from Three Kings Is, C. viridis has been found exclusively Stm, Paemako Reserve, Pseudopanax arboreus, 11 Dec on species of Pseudopanax, whereas there is only one 1993, RCH, #93-374,a-d: 4/4[ff] ad. TO: Arataki SF 98, other instance of any of the other three species of Pseudopanax arboreus [as Neopanax arboreum] leaves, Ctenochiton having been collected off this host plant. 7 a 197 M McKe ie ΝΖ ( 5 -5 3/7[ ] Green (1929) recorded this species off `Hedycarpa' a au u ga oa a ge i o i o oa Pseudopanax arborea (clearly a misspelling of Hedycarya), from crassifolius, 13 Jan 1995, RCH, #95-017a-c: 3/4[ff] ad, Korokoro, Wellington, Jan. 1921. As C. viridis has not 5[f] 3rd. Hauhungaroa Range, Waihaha R,Pseudopanax been collected off this host, this record almost certainly arboreus underside leaves, 14 Jan 1995, RCH, #95-013: refers to C. paraviridis. In addition, Miller (1925) 1/1 [f] ad. Pureora Forest Park, Waipapa Lodge Tk, recorded C. viridis off pukatea (Laurelia novae- Pseudopanax arboreus underside leaves, 15 Jan 1995, zelandiae) and porokaiwhiri (Hedycarya arborea) and n f l nd 4 these records are also presumed to refer to C. p r v r d , Pathogens and parasitoids. Hymenopterous parasitoids while the record off karaka (C r n rp l v t is recorded: Encyrtidae: Ad l n rt d n n t n Noyes; considered to refer to C. h l n, also described as new A. v r b l Noyes. Entomogenous fungal parasites as above. Of the records included under C. v r d by W.M. for C. p r v r d except A r t bb r . Myers (1928) Maskell (1895), those off n x (= d p n x and noted that he never found C. v r d attacked by A. bb r C pr can be confirmed from his original slides but when on d p n x rb r th p n x] whereas those off the other hosts, n l d r , Ath r p r the scale he thought to be C. v r d (now C. p r v r d (= r l , and b are unconfirmed. The present n d r rb r was always heavily attacked by this records for C. v r d are almost restricted to d p n x fungus. spp. and so it seems likely that these last three host genera refer to C. p r v r d . Distribution. From Three Kings Islands, the North Island Morrison and Morrison (1922) wrote a description except south of Lake Taupo, to North Canterbury (Map based on slides mounted from dry material from the W.M. 19). Maskell Collection, of which they stated: "There is no positive evidence that the unmounted specimens were from the original type lot of material". Based on their description and the slides we have studied from the USNM, including the Cotype slide of Morrison & Morrison (which is here identified as Ct n h t n Ge us E E I OC I O e e so & o gso ew p r v r d Henderson & Hodgson n. sp.) we conclude ge us that it was unlikely that they studied C. v r d , but that the slides were most probably a mixture of specimens of C. Type species: Ct n h t n p p r Maskell (here p r v r d (which their description fits) and C. h l n, designated). unrecognised as separate species at that time. In addition, Diagnosis. Adult female. Test: glassy wax plates on Hodgson (1994a), whilst redescribing what he thought young adults, becoming duller due to dark grey sclerotised was C. v r d using USNM material, also commented on derm of mature adult beneath. the great range of variability in the available material and Shape: mature female convex. Body of adult female considered that the material represented more than one round, with a shallow anal cleft and no stigmatic clefts. species. The illustration of C. v r d in Hodgson (1994a) Dorsum: each reticulation area with a small is nearest to C. p r v r d . sclerotised patch, formed initially in centre but growing to fill whole reticulation area on mature insects; anal sclerotisation present anterior to anal plates, horseshoe- Biology. Normally univoltine, but two generations may shaped at first but expanding to join nearby dermal occur in a favourable summer or in a favourable site. C. sclerotisations on old individuals. Dorsal setae absent. v r d overwinters as 1st- or 2nd-instar nymphs. At the Dorsal pores distributed in a reticulate pattern of lines spring flush, female nymphs move to the new leaves on delineating areas underlying wax plates of test; with young shoots (where they settle in preference by a main reticulation areas in 7 longitudinal rows, 9 areas between vein on the undersurface of leaves) and undergo rapid anal plates and anterior margin and with 27 areas around growth and development. The male nymphs remain on the margin. Dorsal pores of 4 types: (i) small, dark old leaves and develop over the same period into adult microductules: present as most abundant pore in all males and emerge from their tests to mate with young adult reticulation lines; (ii) minute simple pores: a few scattered females. The adult females can increase in size at least 12- beside lines of microductules; also 2-3 pores between fold between their last moult and becoming fully grown. reticulation points on submargin; (iii) larger simple pores: Once oviposition begins, the median underside of the present occasionally near reticulation lines and at abdomen gradually retracts to form a brood chamber reticulation points on margin and along anal cleft; (iv) where the eggs are laid and the nymphs shelter after large, sclerotised, flask-shaped macropores, with a hatching. Often about 500 eggs and neonates can be found strongly sclerotised base imbedded in derm and with a in a female's brood chamber. First-instar nymphs emerge ridged top (see SEM 11) — in side view under light from under the posterior end of the test and settle on microscope appearing like a fine brush; interspersed with nearby leaves. Old, post-oviposition females finally drop microductules, mainly within lines around median 3 rows off the host plant, with their telltale depressions remaining of reticulation areas. Preopercular pores, dorsal tubercles for the life of the leaves. and dorsal tubular ducts absent. Anal plates oval-shaped, 112 d n & nd r n (2000 : C d (In t : pt r : C d with minute pores present on upper surface; with 4 pairs of Generic name derivation. The name refers to the long flagellate setae in a row on upper face of posterior half sclerotisations on each of the reticulation areas: from of each plate, usually pointing outwards. Anogenital fold epelis, epelidos (Gr.= freckle) and chiton (Gr.= tunic or with 3 or 4 pairs of setae arising from fleshy basal garment worn close to the skin). extensions along anterior margin and 1 pair laterally. Anal tube short; anal ring with 6 setae. Eyespots present on margin. Margin: marginal setae small and finely spinose, Ep l d h t n p p r (Maske ew com i a io sparse, with 1 seta at each reticulation point on margin, Figs M18, C58, 115 plus an additional seta by each eyespot and 3 setae between stigmatic areas on each side. Stigmatic clefts absent; each Ctenochiton piperis Maskell 1882: 218; —Maskell 1885: stigmatic area with or without a small stigmatic spine; 25 [male description]; —Maskell 1887: '73 [description]; - when present, obviously differentiated but only slightly Maskell, 1895a: 13 [checklist]; -Cockerell, 1896: 330 larger and broader than marginal setae. [checklist]; —Fernald, 1903: 161 [world catalogue]; - Venter: pregenital disc-pores with mainly 5 loculi Hutton, 1904: 226 [checklist]; —Myers, 1922: 199 (range 3-5), present in a group on either side of anterior [checklist];-Green, 1929: 376 [record];-Wise, 1977: 104 end of anal cleft. Spiracular disc-pores with 3-7 (mainly [checklist]; -Deitz & Tocker, 1980: 31 [checklist]; —Ben- 5) loculi in bands between each spiracle and margin. Dov, 1993: 103 [world catalogue]. Multilocular disc-pores with mainly 5 loculi, present in a small group on either side of clypeolabral shield. Ventral Unmounted material: young adults rather flat and round, microducts of 2 types: (i) a small microduct abundant becoming more convex when mature. Test of young throughout; and (ii) a much larger microduct in a female glassy, each wax plate convex, composed of layers submarginal row several pores wide. Ventral tubular of wax, uppermost layers at centre of each plate smallest ducts absent. Ventral setae: anal lobe setae short; anterior and narrowest (secreted when insect was small); most anal cleft setae few; hypopygial setae absent; with 1 pair of basal layers of each plate broadest, filling plate area (and long pregenital setae on segment VII; setae frequent secreted most recently). Teneral female at first light green, medially on abdomen, scarce on thorax and head; other then developing a red-brown horseshoe-shaped pattern on setae as for family. Antennae 6-segmented; third segment each submedian reticulation area; this expands to form a long; setal distribution as for family. Mouthparts usually brownish circular area medially with a light green margin, displaced to 1 side. Spiracles typical of family. Legs well dorsum finally becoming overall dark grey with a dull developed, with a separate tibia and tarsus but no appearance when all sclerotised patches have coalesced articulation; setal distribution as for family; tarsal digitules and thickened. subequal or with 1 distinctly shorter than other; claw Mounted material: as for genus. Length 1.5-2.4 mm, digitules both broad; claw short, without a denticle. Vulva breadth 1.6-2.2 mm. in segment VII. Dorsum: derm of teneral female membranous but becoming sclerotised as described above for genus. Remarks. This genus contains one endemic species, Dorsal pores as for genus; with 0-1 macropores (Fig. Epelidochiton piperis (Maskell). Whilst clearly related to Μ1 i os e io me ia mac o o e i e A a a es the other `Ctenochiton-like' species described here, it is 11 -1 3 μm o g com i e wi s 9 -1 μm; quite distinct. It can be immediately recognised by: sometimes with surface near inner margin of plates (i) the sclerotisation of the reticulation areas on the striated; with 4-8 minute pores on upper surface of each dorsum; plate; setae flagellate, as for genus, in a row on upper (ii) the stigmatic spines which are mostly absent or, if surface, sometimes near outer margin and sometimes present, only slightly larger than the marginal setae; apparently positioned more medially; anteriormost seta (iii) the presence of quinquelocular disc-pores on either ge e a y oi i g ou wa s a e y o g (5 -53 μm side of the clypeolabral shield; o ge a mo e os e io 3 ( a e a a ou 3- 7 μm (iv) the 2 types of ventral microduct, the larger with a long Anogenital fold as for genus; length of longest setae 30-46 inner ductule; μm (v) the distribution of the long, ffagellate setae on the anal Margin: marginal setae as for genus, each marginal plates; se a 7-1 μm o g S igma ic s i es w e ese 1 - (vi) the flask-shaped dorsal macropores. μm o g n f l nd 4 Venter: pregenital disc-pores restricted to a group of ad. Whangaroa, C r n rp l v t , 12 Oct 1968, 9-13 on either side of anterior end of anal cleft. Spiracular R.A. Cumber, No. 1607, E.W. Valentine's Collection: 1/ disc-pores: with 10-35 in each anterior band and 20-45 in 4[ff] ad. Poor Knights Is, Tawhiti Rahi, x l each posterior band. Multilocular disc-pores also in a p t b l , 2 Dec 1980, C.F. Butcher, #81-50b: 2/4[ff] ad. group of usually 1-4 (range 0-9) pores on either side of As previous, except M r p p r x l , #81-58f: 4/7[ff] clypeolabral shield. Ventral microducts of 2 types: (i) ad. AK: Noises Is, Otata I, track E of summit, M r p p r small microducts with a short, tubular inner ductule: those x l leaves, 1 Nov 1977, L.L. Deitz, #77-329g & #80- present medially on thorax and abdomen sometimes larger 289c: 6/11 [ff] ad (with wax plates). As previous, except and more sclerotised than those on submedian areas, and near cottage, #78-152b: 1/2[ff] ad. Noises Is, Otata I, (ii) a microduct about 2 x larger than small microducts, C pr r rp leaves, 1 Nov 1977, L.L. Deitz, with a sunken sclerotised, dark pore and a long inner #77-343b & #80-289d: 4/9[ff] ad. Noises Is, ductule narrowly expanded at the proximal end and Motuharopapa I, SW face, x l p t b l leaves, is a y i ame ous ( o a ways isce a e ese i Ι 2 Nov 1977, L.L. Deitz, #78-11lb: l/19 ad. Noises Is, to 2 submarginal rows. Ventral setae: ventral anal lobe Motuharopapa I, summit, trig point 55 m, C pr se ae s o 5- μm o g; wi -3 a e io a a c e r rp leaves, 4 Nov 1977, L.L. Deitz, #77-329f: 2/ setae; with a pair of long setae on segment VII only, 4[ff] ad. Auckland, Titirangi, t x l n , 13 Oct 1973, o ges 5 -13 μm; um e o se ae me ia y o eac P.S. Dale, No. 1085, J.A. de Boer Collection: 1/5[ff] ad (2 abdominal segment: VII, 3-9; VI, 3-7; V, 4-9; IV, 3-12; split dorsoventrally). As previous, except `shrub', 12 Jan III, 9-15; and II, 4-14; with 4-8 setae medially and 4-9 1983, J.M. Cox, No. 126 and 11 Jan 1983, No. 122: 2/ near each coxa on metathorax; with 5-12 medially and 3- 2[ff] ad. Huia, t x l n leaves, 20 Jul 1975, J. 9 near each coxa on mesothorax; and with 0-1 medially Johanneson: 4/9[ff] ad. Huia , Ar t t l rr t , 26 and 2-4 near each coxa on prothorax; with 2-4 pairs of Sept 1980, G. Hall, #80-282w: 3/6[ff] ad. As previous, interantennal setae. Antennae 6-segmented, occasionally except C r n rp l v t , #80-282x & #90-241: with slight signs of pseudo-segmentation, total length 2/3[ff] ad. As previous, except C pr r b t , C.F. 3 - 9 μm; e g o a ica se a 3 -5 μm Butcher & G. Hall, #80-282s: 2/4[ff] ad. As previous, C y eo a a s ie 115-135 μm o g Wi o except M r p p r x l , C.F. Butcher, #80-282e: 2/ s i acu a e i emes a e io 5-3 μm os e io 3 -3 1[f] ad, 1[f] 3rd. Huia Dam, d r leaf, 28 Mar 1981, μm egs e g s (me a o acic co a 1 -177 μm B.G. Bennett, #81-100f-g: 3/3[ff] ad, 1 [f] 3rd. Huia Dam, trochanter + emu 1 7- μm i ia 115-155 μm a sus d r rb r leaves, 9 Apr 1981, C.F. Butcher, #81 - 77-9 μm c aw 15- μm 105a: 1/1[f] ad. Auckland, Wattle Bay, C r n rp l v t leaf, 5 Jul 1982, P.A. Maddison, #82-187h: 1/ Material examined: LECTOTYPE [f] (here designated): 1[f] ad. Laingholm, M r p p r x l leaf, 14 Jan 1979, NEW ZEALAND: Ct n h t n p p r , from p r W. Park & P.A. Maddison, #79-022a: 2/8[ff] ad. As pre- [=M r p p r] x l , female in test, Jan 1881, W.M.M. vious, x l p t b l leaf, except W. Park, #79- "Entomology Div., DSIR, W.M. Maskell Collection"; 022d: 1/1[f] ad. Bethells, Te Henga Reserve,C r n rp NZAC: 1/1[f] ad. The slide contains a young female in l v t leaf, 19 Oct 1980, P.A. Maddison, #80-296b: good condition, with small amount of wax remaining. 3/3[ff] ad. Waitakere Ra, Walker Bush track, x l PARALECTOTYPES: (i) as for lectotype, except "two fe- p t b l leaves, 5 Nov 1976, M.F. Tocker, #76-316c: 2/ males in tests", a hollow mount slide, the specimens un- 4[ff] ad. Waitakere Ra, x l p t b l , Dec 1979, cleared, in poor condition, CMNZ: 1/2[ff] ad. (ii) as for P.A. Maddison, #79-347b: 1/2[ff] ad. South Piha nr beach, lectotype, "young insect" (a crawler), CMNZ: 1/1 1st. (iii) M r p p r x l leaves, 29 Oct 1979, C.F. Butcher, as for lectotype, "female - 2nd stage", an uncleared speci- #79-305e: 4/11 [ff] ad, 1[m] 2nd, 1 prepupa (pharate). men with most of the pattern of wax plates visible CMNZ: Auckland, Mt Albert Rd, C r n rp l v t [ 1/1 2nd [f]. karaka] leaves, 11 Jun 1992, RCH, #92-188b: 1[f] 3rd (pharate), 2[f] 2nd. Auckland, Glen Eden, t x l n , Other material: NEW ZEALAND: Maskell subsequent Oct 1993, RCH, #93-302a-d: 4/1[f] ad, 1 [f] 3rd,[m]ad,3 2 slide: "male, from p r x l , Oct 1882, W.M.M.", pupae, 2 prepupae (pharate), 1 [m] 2nd. As previous, except NZAC: 1/1 [m] ad - in reasonable condition, uncleared. 1 1 Sept 1994, #94-069a-i: 9/3[ff] ad (1 pharate), 6[f] 3rd From M r p p r x l , J.G. Myers, New Zealand, (1 pharate), 13[f] 2nd, 8[m] 2nd, 1 prepupa, 1 pupa. As pre- 1 ΒΜΝΗ /3[ ] a Ko uko u Auck a vious, except 12 Dec 1995, #95-151 a-b: 2/11 1 sts. WO: C pr r b t , G Mye s o 17 ΒΜΝ 1/ [ ] Raglan, Bridal Veil Falls, r l n v z l nd , 19 ad. Rawene, t x l n , 31 Dec 1982, J.M. Cox: 1/1[f] Sept 1981, C.F. Butcher, #81-282a: 1/1[f] ad. As previ- 11 d n & nd r n (2000 : C d (In t : pt r : C d ous, except S h ffl r d t t , #81-282j: 1/1 teneral [f] io ogy E. p p r is multivoltine, at least in the warmer ad. Raglan, x l p t b l , 19 Sept 1981, C.F. northern areas near Auckland. It is also oligophagous and Butcher, #81-282e: 2/2[ff] ad, 1 [m] 2nd. As previous, ex- able to withstand urban environments (in Auckland at cept tt p r sp., #84-013e: 2/2[ff] ad. Raglan, least). Unlike most other adult female New Zealand M tr d r sp. (rata), 19 Oct 1981, C.F. Butcher, #81- coccids, adult females of Ε. p p r are not entirely sessile 282g: 1/1 teneral [f] ad. CL: Kopu-Hikuai Rd,Al tr n and can walk off the underside of the leaves (their x l , 1 Aug 1968, R.A. Cumber, No. 1583 E.W Val- preferred feeding site) when disturbed. It seems possible entine Collection: 1/1[f] ad.BP:P.Te Puke, 6 Sept 1977, that this is an adaptation for moving to new feeding sites, Ma co m o Ε 3 3 1/ [ ] a M e A o a ui Mi e particularly after overwintering on older leaves. oa si e t x l n , 29 Nov 1991, RCH, #91-346a—b: a oge s a a asi oi s Hymenopterous parasitoids 2/6[ff] ad. Papatea, M r p p r x l , 25 Jan 1993, recorded: Pteromalidae: Aph b t n n ouček e RCH, #93-110: 1/1[f] ad (split dorsoventrally). Hicks Bay, e omoge ous u gi A r t bb r H.S. Fawc. and Wharekahika R, x l p t b l , 20 Sept 1992, Verticillium l n (Zimm.) Viégas occasionally attack E. RCH, #92-277b-c: 3/4[ff] ad (l pharate), 4[f] 3rd, 3[m] 2nd. p p r . As previous, except Macropiper x l , #92-335: 1/3[ff] ad. Te Koau, M r p p r x l , 23 Sept 1992, RCH, is i u io The distribution of E. p p r in the North #92-336: 1/2[ff] ad. As previous, except x l Island lies within the lowland forest from Northland to p t b l , #92-341: 1/1 [f] ad. Lottin Point, Otanga, t x Waikato in the west and to northern Hawkes Bay on the l n ; 29 Sept 1993, RCH, #93-330b: 1/1[m] 2nd. Β east coast, and then from Marlborough Sounds to the White Pine Bush reserve, tt p r n d , 20 Jun Nelson area in the South Island. E. p p r has not been 1991, C.F. Morales, #91-212a—b: 2/1[f] ad (pharate), 2[f] collected between southern Waikato and Wellington 3rd, 1[m] 2nd. G Gisborne, Ch p r l n n (North Island), nor south of Nelson (South Island) (Map [as n ] hedge (close to native trees), 19 Feb 1981, 20). C.F. Butcher, #81-121d: 2/3[ff] ad. SD: d'Urville I x l p t b l , 16 Feb 1971, J.A. de Boer, No. 694: 1/1 [f] ad (split dorsoventrally).NN:Marsden Valley, Ge us I G ISIA Maske d r rb r , 16 Apr 1969, E.W. Valentine, No. 509 J.A de Boer Collection: 2/7[ff] ad (2 split dorsoven- In l Maskell, 1879: 213; —Signoret, 1882a: clviii trally). Motueka, M l t sp., 1 Jul 1917, G. Brittin, No. [synonymy]; -Signoret, 1882b: clxxxiii [synonymy]; — Maskell, 1887: 75 [description]; —Cockerell, 1894b: 1054 131: 2/2[ff] ad. [distribution]; —Cockerell, 1895: 100 [sp. description]; — ema ks This species is easily separable from all other Cockerell, 1896: 330 [checklist]; —Cockerell, 1899a: 394 New Zealand species by the following combination of [checklist]; -Cockerell, 1899c: 332 [key]; —Fernald, 1903: characters: 162 [world catalogue]; —Hutton, 1904: 226 [checklist]; — (i) the round shape, convex at maturity; Green, 1909: 282 [Ceylon sp.]; -Froggatt, 1915: 513 (ii) presence of sclerotised patches on the dorsal derm; [Australian spp.]; -Brain, 1920: 36 [South Africa spp]; - (iii) a sclerotised collar on dorsum anterior to anal plates; Froggatt, 1921: 25 [taxonomy]; —MacGillivray, 1921: (iv) presence of 5-locular disc-pores laterad to the mouth- 171, 178 [catalogue]; —Morrison & Morrison, 1922: 75 parts; [redescription type]; —Mandihassan, 1923: 97 (v) long, outward-pointing discal seta on each anal plate; [relationships]; —Steinweden, 1929: 201, 202, 206, 207, (vi) flask-shaped dorsal macropores; 237 [classification]; —Ramakrishna Ayyar, 1929: 35, 41 (vii) complete absence of ventral tubular ducts, but with [Indian sp.]; —Goux, 1933: 123 [mention]; —Lindinger, numerous microducts throughout the venter. 1937: 187, 191 [checklist]; -Hempel, 1937: 10 [Brazilian Although E. p p r is a widespread and common sp.]; -Lepage, 1938: 357 [Brazilian sp.]; —Hodgson, species, it has been recorded only once from an introduced 1967b: 1 [African spp.]; —Hodgson, 1969: 17 [African host plant, namely from a hedge of Ch p r spp.]; -Wise, 1977: 105 [checklist]; -Tao, 1978: 82 l n n (Lawson's cypress); however, it was noted at [Taiwan checklist]; —Tao et l., 1983:91 [Taiwanese sp.]; the time that the hedge was "close to native trees". This —Hamon & Williams, 1984: 56 [Florida sp.]; —Tang, 1991: isolated record is considered to be a chance occurrence as 321 [Chinese sp.]; —Ben-Dov, 1993: 146 [world there are no other records of an endemic New Zealand soft catalogue]; —Hodgson, 1994a: 289 [redescription type]. scale becoming established on a non-native plant. y e s ecies In l p t ll Maskell (designated by Fernald, 1903: 162) n f l nd 4 . . Ep l d h t n p p r (Maske , . com ., a u ema e. 6 d n & nd r n (2000 : C d (In t : pt r : C d Fig. 116. In l p t ll Maske , a u ema e. Α ig a o a ium. n f l nd 4 117 iag osis a u ema e es glassy, conical and 3rd segment longest; setal distribution: 2 setae on scape; a limpet-like, with 2 concentric lines around it marking long flagellate seta and a short seta on pedicel; 2 short change in growth between instars; margin approximately setae on 3rd segment; 1 fleshy seta on both 4th and 5th 10-sided, each side rather fluted at each indentation and segments; total of 8 setae on apical segment, l very short with median apex roundly pointed. seta, 1 rather short flagellate seta and 6 fleshy seta S a e body rather 10-sided (decagonal), lacking including longest, latter usually clubbed terminally. stigmatic clefts and with a very shallow anal cleft. Spiracles quite small, each with a distinct, sclerotised o sum derm membranous, without reticulation spiracular plate. Legs typical of family but rather small lines. Dorsal setae absent. Dorsal pores of 3 or 4 types and with tibia and tarsus fused; setal distribution: coxa present: (i) minute microductules, with along, filamentous with 4 setae, trochanter with 1 long flagellate seta, femur inner ductule; (ii) small, heavily sclerotised, convex with I seta, tarsus usually with 1 seta (occasionally 2 setae simple pores; (iii) much larger simple pores or on metatarsus); tarsal digitules variable; claw digitules macropores; and (iv) round, simple pores, similar to (iii) alike with broad apices; claws with or without a small but smaller: present sparsely throughout. Two rather large denticle. Anterior margin of segment VII near centre of membranous areas present approximately dorsad to abdomen, far anterior to anal plates, with a distinct vulva metathoracic legs, each with numerous unevenly round immediately posterior to its anterior margin. pores with a granulate surface. Preopercular pores, dorsal tubular ducts, and submarginal tubercles absent. Anal ema ks Although many species have been placed in plates each with anterior margin almost at right angles to this genus in the past, it is here considered that it is body axis and with outer angle almost a right angle; each currently a monotypic genus, as was also concluded by plate with 5 long spinose spines on inner margin and on Morrison & Morrison (1922), and endemic to New apex, plus 1 shorter spine set slightly onto dorsum; plates Zealand. For distinctive characters, see under the species lacking minute pores on dorsal surface. Anogenital fold description below. without setae. Anal ring with 6 setae; anal tube much longer than length of anal plates. Ma gi marginal setae spinose, of 2 types: (i) a club- shaped spine, blunt apically and narrowing abruptly at In l p t ll Maske base, with a large basal socket; and (ii) a sharply spinose seta, with a quite large basal socket; marginal setae on anal igs Μ 9 C59 11 lobe not differentiated from other marginal setae. In l p t ll Maskell, 1879: 213; —Maskell, 1882: 219 Stigmatic clefts absent; stigmatic spines not differentiated [male description]; -Maskell, 1887: 78 [description]; - from marginal spines. Eyespots small, very indistinct on Maskell, 1895a: 14 [checklist]; -Cockerell, 1896: 330 anterior outer corners of decagon. [checklist]; —Fernald, 1903: 162 [world catalogue]; — e e derm membranous. Pregenital disc-pores Hutton, 1904: 227 [checklist]; —Myers, 1922: 199 with 5 or 6 loculi: present in a line on either side of anal [checklist]; —Morrison & Morrison, 1922: 75 area and extending down margins of anal cleft. Spiracular [redescription]; -Miller, 1925: 33, 64 [host]; —Green, disc-pores with mainly 5 loculi; present in bands 1929: 377 [record]; —Steinweden, 1929: 212 extending to margin from anterior spiracles only; posterior [classification]; —Lindinger, 1937: 187 [checklist];-Wise, bands only represented by a group of 2 or 3 pores just 1977: 105 [checklist]; —Deitz & Tocker, 1980: 28, 29, 30, anterior to peritreme. Preantennal pore possibly absent. 31, 32 [checklist]; -Ben-Dov, 1993: 149 [world Ventral microducts present, scattered sparsely throughout catalogue]; —Hodgson, 1994a: 289 [redescription]. much of venter. Ventral tubular ducts of l type, with a rather long outer ductule and a long inner ductule with a U mou e ma e ia test glassy, shaped like a limpet large terminal gland; present in a narrow submarginal ( t ll , as for genus; internal wax structure a band and with a few elsewhere. Ventral setae spinose, combination of horizontal layers near apex, appearing distributed as follows: ventral anal lobe setae absent; more as honeycomb areas in the outer side walls, and solid anterior anal cleft setae present laterad to pregenital disc- wa i e i e si e wa s ( ig Μ 9 pores; hypopygial setae absent; long pregenital setae absent, each abdominal segment with short bands of stout Mou e ma e ia shape of body rather 10-sided setae present medially across each segment; with very few (decagonal), lacking stigmatic clefts and with a very setae on thorax and head; with interantennal setae; shallow anal cleft, about 1/30th body length. Length 0.9- submarginal setae sparse. Antennae 6- or 7-segmented, 2.4 mm, width 0.9-2.8 mm. 11 d n & nd r n (2000 : C d (In t : pt r : C d Dorsum: dorsal pores of 4 types: (i) minute medially across each segment as follows: VII, 4-8; VI, 5- microductules: in an broken submarginal band up to 4 or 10; V, 5-9; IV, 5-9; III, 3-6; and II, 3-5; with 2-3 setae more pores wide but tending to be absent at each corner of just anterior to each metacoxa, 1 anterior to each decagon; largest microductules in a triangular area just mesocoxa, and 0-1 near each procoxa; longest about 16 anterior to anal plates; (ii) small, heavily sclerotised, μm; wi 1-3 ai s o i e a e a se ae; su ma gi a convex simple pores: present in a line just dorsad to setae sparse, with 4-5 pairs posteriorly on abdomen, 2-3 marginal spines, with about 1 pore per 3 or 4 spines; with anteriorly on head, and none on thorax. Antennae 6- or 7- 31-47 around head between anterior stigmatic areas; (iii) segmented (segmentation of long 3rd segment indistinct), large simple pores/macropores, much larger than simple 3 segme o ges ; o a e g 111-139 μm; e g o pores: present sparsely throughout median part of dorsum; terminal seta 7-17 pm, much shorter than stout fleshy and (iv) round, simple pores, similar to (iii) but smaller: setae on apical segment; length of lateral flagellate setae present sparsely throughout. Two large, membranous, e y s o 1 -1 μm C y eo a a s ie 117-1 μm cribriform-like areas present approximately dorsad to long. Spiracles quite small, each with a distinct, me a o acic egs; si e o eac a ea 9 -1 3 53- μm sclerotised spiracular plate; width of peritremes: 23-30 each with about 60 unevenly round pores with a granulate μm egs a e sma ; wi i ia a a sus use surface, distributed mainly around edge. Anal plates as for although on some specimens a slight indication of ge us; e g o a es -5 μm com i e wi 73- pseudosegmentation present; lengths (metathoracic): coxa μm; eac a e wi 5 o g s i ose se ae o i e ma gi 5-77 μm oc a e + emu 1-9 μm i ia + a sus and on apex, plus 1 shorter spinose seta set slightly onto 1 -131 μm; c aw 1 -15 μm; oc a e wi o y 1 o g dorsum near inner margin about 1/3rd from posterior seta. a e ; e g o se ae a e io s i es 7-31 μm; e s i es o i e ma gi 39- 3 μm; a ica s i e 3 -51 Material examined: LECTOTYPE (here designated): μm a s i e o o sum 1 - 1 μm; a e s i e o 1 o NEW ZEALAND: Inglisia patella, from Drimys marginal spines can be club-shaped (similar to club- [=Pseudowintera sp.], two adult females, July 1878, shaped marginal setae), all others strongly spinose with W.M.M.; CMNZ: l/2[f] ad. The lectotype female has an parallel sides and a sharp point. everted anal tube, is nearest the original Maskell label, Margin: marginal setae spinose, of 2 types: (i) club- and is clearly marked. s a e s i es e g 1 - μm; u a ica y a PARALECTOTYPES: (i) the second ad on the lectotype narrowing abruptly at base, with a large basal socket: slide (above). (ii) same data as lectotype, except "tests on approximately evenly spaced, with 46-56 around head leaf', = whole mount slide, NZAC: 1/1 ad, 9[m] 2nd. between anterior stigmatic areas; and (ii) sharply spinose s i es e g 1 -1 μm wi a qui e a ge asa socke Other material: NEW ZEALAND: Ex Maskell's dried usually present all around margin, alternating with club- material, mounted by C.F. Butcher: 1/1 [f] ad. Maskell coll., shaped setae but sometimes most frequent at corners of #46 USDA: 1/1 ad. DSIR #331, no coll, data, remounted decagon; with 25-59 around head between anterior by C.J. Hodgson: 4/4[ff] ad. BP: Lottin Point, Otanga, stigmatic areas; with a few sometimes on margins of anal Hedycarya arborea, 27 Apr 1993, RCH, #93-171: 1/l [m] cleft; a club-shaped spine also occasionally present on 2nd. As previous, except 29 Sept 1993, #93-314a-c: 3/ dorsal surface of each anal lobe. 3[f]ad. As previous, except 3 Nov 1993, #93-351a: 1/ Venter: pregenital disc-pores as for genus. Spiracular 1[f] ad, 1[f] 2nd, 1 1st + 4 neonates. Onepoto Bay, disc-pores in complete bands of 16-33 pores between Hedycarya arborea, 15 Mar 1994, RCH, #94-048: 1/1[m] margin and each anterior spiracle only; posterior bands ad. GB: Taikawakawa, Hedycarya arborea, 2 Feb 1993, only represented by a group of 2 or 3 pores just anterior to RCH, #93-048: 1/1[f] ad. As previous, except 18 Mar 1993, peritreme. Ventral microducts quite distinctive, with a #93-089: l/1 [m] 2nd, 1/1 2nd (+ bit). As previous, except long inner ductule swollen distally; scattered among 1 May 1993, #97-067: 1/1[m] 2nd. TO: Rangitoto Station, submarginal band of tubular ducts, in a band on Mangatutu, Pseudowintera colorata, 9 Nov 1996, RCH: mediolateral folds of abdominal segments and with a few 1/1 [m] 2nd. Ohakune,Pseudowintera[asDrimys], colorata medially on thorax and head. Ventral tubular ducts as for 6 June 1924, G. Brittin, #107: 2/2[m] 2nd. NN: Motueka, genus: in a narrow submarginal band, and with a few Pseudowintera colorata [as Drimys], G. Brittin coll. 1924, sometimes present more medially near antennae, Brittin #107: 1/1 [prob][m] 2nd [without coverslip], USDA. mouthparts and spiracles. Ventral setae as for genus, but Motueka, Hedycarya arborea, 9 Jan 1938, G. Brittin, #107: with 3 pairs of anterior anal cleft setae laterad to pregenital 2/2[ff] ad. Motueka,Elaeocarpussp., 2 Feb 1938, G. disc-pore groups; with bands of short, stout setae present Brittin, #107, USDA, #107: 2/l[f] ad, 2 pupae. Whangamoa n f l nd 4 Saddle, d nt r x ll r , 18 Sept 1968, J.A. de Biology. Normally located on the underside of leaves of Boer, #438: 1/2[m] 2nd, [+3[f] 2nd, prob.sp.].Ct n h t n its favoured host plants: species of d r , BR: Buller Gorge, d nt r l r t , Mar 1971, tt p r , andThe d nt r . number of J.A. de Boer, #724: 1/1 [f] ad (split dorsoventrally) + 1/1[f] generations per year is uncertain but, as the immature ad l h t n, fl v (split dorsoventrally). Fletcher's stages have been recorded in September and March, it is Creek, nr In n h , d nt r l r t , 7 Mar 1972, likely that there are at least two generations, one in the J.A.K. Farrell: 1/1 [f] ad (v. poor). Redman's Creek, nr. spring and the other in the autumn. Reefton, d nt r l r t , 6 Nov 1972, J.A. de Boer, #952: 1/1 [f] ad. Murchison, nr upper Matakitaki, Pathogens and parasitoids. Hymenopterous parasitoids d nt r l r t , 5 Jan 1998, C.J. Hodgson, #98- recorded are: Aphelinidae: rd l sp.; Platygastridae: 074: 1/1[f] ad. MC: [On hill above Lyttelton], on Err l sp. (undescribed); Pteromalidae: Aph b t d nt r [as r ] sp., June 1881, W.M.M., n n ( ouček NZAC: 2/1 1st, l/rostrum, antenna, foot and spines (poor). Riccarton Bush, d nt r l r t [as r ], 31 Distribution. In lowland forest throughout New Zealand Dec 1916, G. Brittin, #107: 1/1[f] ad. Christchurch, (Map 21). tt p r sp., 13 Aug 1921, J.G. Myers, BMNH: 1/ 3[ff] ad. Christchurch, d nt r l r t [as r ], no date, G. Brittin #107, BMNH: 1/1[f] ad. FD: Thompson Sound, Bauza I, d nt r l r t , Mar 1984, C.F. Butcher, #98-072: 1/2[ff] ad. Charles Sound, Elaenor 1, tt p r l n underside leaves, 26 Jan 1996, RCH: 11/9[ff] ad, [f] (settled) 1st, 5 crawlers. Remarks. The main characters of adult female In l p t ll are: (i) the decagonal, conical shape of the test; (ii) the decagonal shape of the body margin; (iii) the shape of the anal plates and the shape and distribution of the anal plate setae; (iv) the presence of 2 types of marginal spines; Ge us KA ASI IS e e so & o gso , (v) the presence of 2 `cribriform'-like areas medially on the abdomen; ew ge us (vi) the absence of a posterior spiracular disc-pore band; Type species: Ct n h t n p rf r t Maskell (here (vii) the position of the vulva, towards the middle of the designated) abdomen; (viii) the presence of a spiracular plate around each Diagnosis. Adult female. Test: glassy, roundish, thin at spiracle. first, thickening with maturity. Ben-Dov (1993) included 17 species in the genus Shape: mature adults fairly small (less than 4.5 mm In l . The senior author has seen most of these species long), oval to almost round, moderately convex when and he agrees with Morrison & Morrison (1922) that none mature, shrinking towards anterior end during larviposition are congeneric with I. p t ll . Indeed, a morphological so that posterior end of test forms a brood chamber; anal study of all the life stages of I . p t ll , including the adult cleft shallow; colour either translucent or light greenish, male, suggests that In l is a plesiomorphic genus and sometimes with a reddish stripe when immature. shares few characters with other known soft scales. Dorsum: derm membranous. Dorsal setae absent. In l , therefore, appears to be a monotypic genus Dorsal pores forming a distinct reticulate pattern endemic to New Zealand. delineating reticulation areas (which underlie wax plates In l p t ll is an unusual and distinctive species of test), in 5 or 7 longitudinal rows, with 5-8 reticulation and is quite unlike any other known from New Zealand. areas between anal plates and anterior margin and 26-29 Note: the measurements of the limbs, etc. given above are areas around margin. Dorsal pores of 3 or 4 types: (i) about 10% smaller than those given by Hodgson (1994a); small, dark microductules, each with a long inner filament the reasons for this are unclear. with a small balloon-like proximal end: throughout but 1 d n & nd r n (2000 : C d (In t : pt r : C d mainly in lines of reticulation; (ii) simple pores, flat: of 2 posteriorly on segment VII. sizes: (a) largish pores laterad to anal plates and/or in a submarginal band, and (b) smaller pores within Remarks. This genus contains three species: K l r reticulation lines; and (iii) macropores, rather flat, round d pr (Maskell), n. comb., K. p r d pr Henderson to oval, smaller than pregenital disc-pores (absent on Κ. & Hodgson n. sp. and K. p rf r t (Maskell) n. comb. p r d pr when present, restricted to reticulation lines and most abundant medially. Preopercular pores, The main diagnostic characters of the species in this genus dorsal tubercles and dorsal tubular ducts absent. Anal are: plates together widest at anterior 1/5th to 1/4th, tapering to (i) marginal setae particularly abundant, with some a narrow apex, often with a small pointed protrusion extending up margins of anal cleft (few there on Κ. apically; each plate with 4 finely spinose setae, 2 along p rf r t inner margin, 1 apically and another posteriorly on upper (ii) at least a few larger simple pores dorsally on either side surface near apex. Anal plates without supporting bars or of anal plates, usually common and forming a group; plates. Anogenital fold with 2 pairs of long setae along (iii) 2 types of ventral tubular ducts, a larger type anterior margin and a single seta on each lateral margin. submarginally and a smaller type medially and Anal tube quite long; anal ring with 6 setae. mediolaterally on posterior abdominal segments and Margin: marginal setae abundant, strongly spinose laterad to anal cleft, where common; with narrow basal sockets, in a single marginal line which (iv) ventral tubular ducts present in a submarginal band also extends at least part-way up anal cleft (few at distal near or on margin, the ducts often lying radially; end only on K. p rf r t reticulation setae usually larger; (v) groups of ventral tubular ducts never present near marginal setae on anal lobe undifferentiated. Stigmatic mouthparts. clefts shallow but distinct, without stigmatic sclerotisations; Two types of ventral tubular duct also occur on each cleft with a single spinose, stigmatic spine; rather Ct n h t n h l n, but they are very few and do not short, never longer than about 2 x length of longest have this distribution. marginal setae. Eyespots present. Species in the genus K l r are similar to those in Venter: pregenital disc-pores with mainly 8-10 outer Ct n h t n and Cr t ll t t in having abundant pre- loculi and a single oval inner loculus; distributed across genital disc-pores medially across the abdominal most or all abdominal segments medially and with groups segments, each pore with mainly 8-10 loculi. In addition, on most mediolateral folds. Spiracular disc-pores with species in the genus Cr t ll t t also resemble those in mainly 5 loculi, in narrow bands between spiracles and K l r in having: margin; with 0-5 extending medially past peritreme. (i) 2 types of ventral tubular duct; Preantennal pores present or absent. Ventral microducts (ii) small spiracles; in species-specific distributions. Ventral tubular ducts of (iii) abundant spinose marginal setae approximately 2 types: (i) a larger duct, forming a distinct submarginal evenly spaced around margin; band, some ducts opening very close to margin; most ducts (iv) large simple pores on the dorsum, generally associated near margin lying radially; and (ii) a smaller duct, either with the anal plates. without a terminal gland or with only a small gland, However, Cr t ll t t present medially on posterior abdominal segments and species differ in having laterad to anal cleft. Ventral setae: with 1-2 pairs of ventral tubular ducts present medially on the thorax and in anterior anal cleft setae; hypopygial setae absent; with broad submarginal bands, with the ducts not tending to lie pairs of long setae restricted to either segment VII or radially. segments VI and VII; other setae distributed as for family. Ct n h t n species differ from those of K l r in: Antennae well developed, 6- or 7-segmented (3rd segment (i) having very large spiracles; (or 3rd+4th in 7-segmented antennae) subequal to or (ii) having spinose marginal setae rather unevenly spaced longer than other 5 segments); setal distribution as for around the margin; family. Mouthparts normal and not displaced. Spiracles (iii) the distribution of the ventral tubular ducts. typical of family. Legs well developed, each with a The species currently placed in K l r are all separate tibia and tarsus but no tibio-tarsal articulatory endemic to New Zealand. sclerosis; distribution of leg setae as for family; claws without a denticle; claw digitules similar and broad; tarsal Name derivation. From l r (Gr., f.) meaning digitules dissimilar, 1 shorter and narrower than other; "garment fringed at the base", referring to the beautiful tarsal campaniform pores absent. Vulva opening marginal wax fringe in K. p rf r t (Maskell). n f l nd 4 2 ig. . K l r d pr (Maske , . com ., a u ema e. 22 d n & nd r n (2000 : C d (In t : pt r : C d ig. 8. K l r p r d pr e e so & o gso , . s ., a u ema e. n f l nd 4 2 ig. . K l r p rf r t (Maske , . com ., a u ema e. 24 d n & nd r n (2000 : C d (In t : pt r : C d Key o a u ema e K l r by Maskell (1887, p. 66) is here believed to refer to old Pairs of long pregenital setae only present on abdominal females attacked by fungus: "Test of adult female flat, segment VII; ventral microducts of 2 sizes, largest nearly circular, thin, waxy, greyish coloured; fringe forming a marginal band between margin and inconspicuous or sometimes absent. No perforations or submarginal band of ventral tubular ducts, smaller air-cells. Diameter about 1/7 in." "Adult female filling the microducts abundant medially on thorax .... depressa test, but shrivelling to the anterior end of the test at —Pairs of long pregenital setae present on 2 abdominal gestation. Colour brownish to grey." segments ( II and VI); ventral microducts of 1 size, with none present between margin and submarginal Mou e ma e ia : body oval to almost round; length band of ventral tubular ducts 2 2.0-4.5 mm; breadth 1.9-4.1 mm; anal cleft about 1/8th body length; stigmatic clefts shallow. 2 Ventral tubular ducts in a distinct narrow marginal band about 1 duct wide, most ducts lying radially but often o sum: dorsal pores in a reticulate pattern, with with a few also scattered submarginally away from reticulation areas in 7 longitudinal rows, with 8 marginal band; with, at most, 2-3 marginal setae reticulation areas between anal plates and anterior margin extending up anal cleft; ventral microducts absent and with 29 areas around margin. Dorsal pores of 3 types: medially on thorax perforata (i) microductules: present throughout, but most frequent associated with lines of reticulation; (ii) simple pores: —Ventral tubular ducts in a submarginal band distinctly rather variable in size: (a) small, flat pores, slightly larger several ducts wide, with a clear marginal area free than microductule pore: frequent associated with lines of from ducts but sometimes with a few ducts scattered reticulation; and (b) rather larger simple pores: in a sparse more medially on submargin; marginal setae band near margin; and (iii) large, heavily sclerotised extending part way up anal cleft; ventral microducts macropores, slightly sunken and convex: about equally frequent medially on thorax paradepressa abundant throughout except near margin; with 7-10 in os e io me ia mac o o e i e A a a es 137-1 μm o g com i e wi s 135-173 μm; wi - mi u e o es o o sa su ace o eac a e; e g o se ae i e K l r d pr (Maske ew com i a io ma gi 1 1 - μm; i e ma gi 1 - μm (some imes a ge a s ou e a a ica se a ; a ica 9- Figs C60, C61, 117 μm a ou e ma gi 1 μm A oge i a o wi - Ct n h t n d pr Maskell, 1884: 132; –Maskell, 3 ai s o se ae a o g a e io ma gi a 1 se a o eac 1887: 66 [description]; –Maskell, 1892: 19 [taxonomy]; – a e a ma gi ; o ges 3 - μm Maskell, 1895a: 12 [checklist]; –Cockerell, 1896, 330 Ma gi : ma gi a se ae s i ose 9- μm o g wi [checklist]; –Fernald, 1903: 159 [world catalogue]; – 1 - 7 a e a y e wee s igma ic c e s; e icu a io se ae Hutton, 1904: 226 [checklist]; –Myers, 1922: 199 o y s ig y a ge ; ma gi a se ae e e i g a ou a i [checklist]; –Wise, 1977: 104 [checklist]; –Deitz and o a a -way u a a c e ma gi S igma ic s i es a e Tocker, 1980: 28 [checklist]; –Ben-Dov, 1993: 100 s ou a u some imes s ig y cu e wi qui e we [world catalogue]. e e o e asa socke s; e g 1 - 7 μm Eyes o Ct n h t n d pr (small form) Maskell, 1892: 19. ese Ct n h t n d pr forma n r Maskell, 1895a: 12 e e : pregenital disc-pores with mainly 10 loculi [checklist]; –Cockerell, 1896: 330 [checklist]. (range 7-11); number per segment (medially/mediolaterally Ct n h t n d pr n r Maskell; –Fernald, 1903: on each side): VII, 10-24/24-32; VI, 26-50/6-11; V, 26- 160 (synonymy); –Wise, 1977: 104 [checklist]; –Deitz & 42/3-7; IV, 22-43/2-7; III, 2-16/1-4; and , 0/0-1; with Tocker, 1980: 30 [checklist]. none laterad to metacoxae. Spiracular disc-pores in bands Ct n h t n d pr n r –Ben-Dov, 1993: 100 l-4 pores wide; with 13-26 in each band; each anterior [mis-spelling of d pr ]. band with 0-5 pores present a short distance mesad to peritremes. Ventral microducts of 2 sizes: (i) larger U mou e ma e ia : mature female moderately convex microducts: present in a distinct submarginal band, lying and rounded like a shell; when larvipositing, test mainly between ventral tubular ducts and margin or within appearing bicoloured, greenish in anterior half due to band of tubular ducts; and (ii) smaller microducts: present colour of underlying female and fawn in posterior half due medially from labium to abdominal segment II. With 1-2 to colour of nymphs (Fig. C61). The following description preantennal pores/antenna. Ventral tubular ducts of 2 n f l nd 4 2 types, as for genus: smaller duct rather short, with a fine 1891, W M M "; AC 1/1[f] ad. inner ductule, lacking a terminal gland; frequent on either side of anal cleft and extending around anogenital area but PARALECTOTYPES: ex Maskell's dry material #201, rarely extending onto more anterior abdominal segments mounted 2 Mar 1972 by J.A. de Boer: 2/4[ff] ad. Mounted except a single duct usually present laterally on segment from W.M. Maskell's dry material Ct n h t n d pr VI; very occasionally, a few also present in submarginal small form, by RCH, NZAC #95-020a—f: 6/3[ff] ad, 2[f] band of larger ducts. Ventral setae: ventral anal lobe setae 3rd, 1 [m] 2nd.Mask.,Ct n h t n no data, d pr n r 1 - 3 μm o g; wi 1 ai o a e io a a c e se ae; New Zealand, Mask. coll. no.201, USNM: 1/1[f] ad. with long pregenital setae restricted to a pair medially on abdominal segment VII; number of short setae medially on Other material: NEW ZEALAND: ND: Omahuta SF, Kauri each abdominal segment: VII, 4-8; VI, 6-8; V, 4-10; IV, Sanctuary, C pr sp., 6 Oct 1980, J.S. Noyes, #81- 2-8; III and II, 0-4; with 0-2 setae near each metacoxa, 0- 40c: 1/[m] 2nd. AK: Huia Dam,sp.,C pr 31 Oct 1980, 2 near each mesocoxa, and with 1-3 near each procoxa C.F. Butcher, #80-311e: l/1[f] ad. Waitakere Ra, (often rather spinose); length of setae associated with each Karamatura V, C pr rb r underside leaves, 21 oco a qui e o g 1 - μm some imes e y s i ose; Oct 1994, RCH, #94-093a—f: 6/6[ff] ad. As previous, ex- with 3-4 pairs of interantennal setae laterally plus 1-2 cept 1 Dec.1994, #94-125a—b: 2/1[f] ad, 11 1st. As previ- setae medially; with 5-6 submarginal setae on each side ous, except 31 July 1995, 1/1 [m]2nd. As previous, except between each stigmatic area. Antennae 6-segmented, with 8 Oct 1995, #95-098a-e: 5/2[f] 3rd, 2[m] 2nd, 2 pupae. As a constriction between 2nd and 3rd segment; total length previous, except 11 Sept 1995, G. Hall & L.H. Clunie, 7 -3 3 μm; e g o a ica se ae 3 -5 μm; es y se a #95-099: 1/1 [m] 2nd. Waitakere Ra, 179 Laingholm Drive, near apex of each antenna particularly stout. Length of C pr rb r ea es 13 Se 1997 Ν A Ma i 97- c y eo a a s ie 131-1 μm Wi o s i acu a 1 9a-c 3/ [ ] 3 1[m] icks ay W a eka ika e i emes a e io 5- 5 μm os e io 5-7 μm egs i e C pr rh n d , 20 Sept 1992, RCH, #92- e g s (me a o acic co a 5- 7 μm oc a e + 304a—b: 2/4[ff] ad, 1[m]2nd.C pr TK: Mt Egmont, emu 11 -151 μm i ia 1-119 μm a sus 3-9 μm sp. leaves, 24 Feb 1983, C.F. Butcher, #83-067g: 1/2[ff] c aw 1 -1 μm ad. NN: Golden Downs, Gordons Creek, C pr sp., 4 ec 19 3 W A o oway ΝΖ o (a / [ ] a Material examined: LECTOTYPE [f] (here designated): Wai oa Go ge C pr ? l n , 13 Dec 1967, J.A. de NEW ZEALAND: labelled "Ct n h t n d pr Mask., Boer, no. 290: 1/1 [f] ad (split dorsoventrally). FD: Breaksea ex Mask, dry material #36, mounted 2.III.72". NZAC; I, (no host), 26 May 1982, C.F. Butcher, #82-175h: 2/1[f] mounted (2 March 1972) by J.A. de Boer: 1/1 [f] ad. ad (old), 1[m] 2nd. Dusky Sound, Seal I,C pr f t d , 8 Mar 1983, C.F. Butcher, #83-332g: 3/4[ff] PARALECTOTYPES: NEW ZEALAND: (i) labelled ad. Resolution I, Disappointment Cove, C pr "Ct n h t n d pr , from l nth , adult male and l n , 0 Mar 1983, C.F. Butcher, #83-293i, & 83-320g: female 2nd stage, May 1883, W.M.M."; gold label: Ento- 4/4[ff] ad. mology Div., DSIR, NZ, W.M. Maskell Collection, NZAC: l/1 [m] ad, 1 [m] 2nd; the "female 2nd stage" is actually a 2nd- Remarks. Maskell's original material was collected off instar male. (ii) as above, except "antenna and foot of l nth sp. and C th sp. and was sent to him from female" (actually those of a 2nd-instar male with part of Hawke's Bay by Rev. Colenso (Maskell, 1884, p. 133). the body margin attached); CMNZ: 1/1 [m] 2nd. (iii) mounted Two of the original slides of Ct n h t n d pr from W.M. Maskell's dried material of Ct n h t n Maskell in the syntype series available for study include a d pr by RCH, no date or host, NZAC #95-019a—b: 2nd-instar male, a piece of a 2nd-instar male and an adult 2/(part of)1 [f] ad, 1 [m] 2nd. (iv)Mask.,Ct n h t n d pr male. None of these specimens is ideal for designation as New Zealand, no data, Mask. Coll. no. 36, USNM: 1/1 [f] lectotype. In addition, a third slide listed by Deitz & ad. [Paralectotype: collection data as for paralectotype (ii) Tocker (1980) is clearly a misidentification by Maskell, as except "adult female", CMNZ: 1/1 [f] ad, a misidentification, it is actually an adult female of Ct n h t n h l n now considered to be Ct n h t n h l n described as (described as new above). However, we are confident that new above]. all the dry material in Maskell's collection #36 represents some of the original material and that it agrees with his Ct n h t n d pr n r MaskelI (small form): concept of C. d pr . We therefore consider it to be LECTOTYPE [f] (here designated): NEW ZEALAND: lab- syntypic and have designated one of the two adult females elled "Ct n h t n d pr small form, adult female. mounted from the dry material collection #36 as lectotype 1 d n & nd r n (2000 : C d (In t : pt r : C d of Ctenochiton depressus. The only remaining unmounted unfolded (unlike the folds on K. p rf r t post- Maskell dry material of C. depressus are nymphs. ovipositional females). The adult female lectotype of Ctenochiton depressus minor Maskell (what Maskell originally described as the Pathogens and parasitoids. Hymenopterous parasitoids "small form") agrees both with further adult females recorded are: Encyrtidae: Adelencyrtoides inconstans mounted from Maskell's dry material collection #201 Noyes and A. variabilis Noyes; Pteromalidae: Aphobetus relating to his "small form" description and with the above nana ( ouček females of C. depressus (#36). We therefore uphold their synonymy by Fernald (1903). Distribution. Throughout, from Northland to Fiordland, Adult females of K. depressa are characterised by the but not recorded from eastern South Island (Map 22). following combination of characters: (i) a submarginal band of tubular ducts, separated from the margin by a marginal band of larger ventral microducts; K l r p r d pr e e so & o gso ew (ii) smaller ventral microducts restricted to medially on s ecies thorax, posterior to labium and abdominal segment II; Fig. 118 (iii) a pair of long pregenital setae on abdominal segment VII only; Unmounted material: unknown. (iv) dorsal macropores flat and simple, present throughout except on radial reticulation lines near margin; Mounted material: body oval, with very shallow (v) a distinct constriction present between the 2nd- and stigmatic clefts and a shallow anal cleft, about 1/9th body 3rd-antennal segments (also present on K. length; length 3.0-5.0 mm; breadth 2.1-3.6 mm. paradepressa); ( ii a e s o a e ae ( 7 -3 3 μm Dorsum: dorsal pores in a reticulate pattern, with K. depressa is close to K. paradepressa, but the latter reticulation areas in 7 longitudinal lines, with 5-6 has no large dorsal macropores, and to K. perforata, but reticulation areas between anal plates and anterior margin the latter species has 7 longitudinal rows of reticulation and with 28 areas around margin. Dorsal pores of 4 types: areas on the dorsum rather than 5. For further differences, (i) small, dark microductules: present throughout but most see under these species. frequent within lines of reticulation; (ii) slightly larger, Maskell (1884) states that the original material was dark, granular, flattish simple pores: frequent in all collected off Plagianthus sp., although all subsequent reticulation lines; (iii) similar-sized simple pores which collections have been from Coprosma sp. As the type appear pale (possibly a paler type (ii)): scarce, mainly series appears to consist of adult and 2nd-instar males, laterad to anal plates; and (iv) rather larger simple pores, Plagianthus may not be a true host of K. depressa as male similar to (ii) but restricted to around the margin and stages are known to settle on many plant genera apparently laterad to anal plates; macropores absent. Anal plates unsuitable for the females. a e e o ga e 1 9-19 μm o g com i e wi s 1 3- 1 μm; wi - mi u e o es o o sa su ace o eac a e; se ae i e y s i ose e g s i e ma gi o - Biology. K. depressa favours Coprosma spp. with μm; a ica se a 3 - 3 μm; ou e ma gi se ae a smallish leaves as host plants. It apparently overwinters as missi g A oge i a o wi ai s o se ae a o g nymphs, becoming adult and larvipositing in the spring to a e io ma gi ( o ges a ou 5 μm a a s o e ai early summer. Post-ovipositional females have a o a e a ma gi s distinctively contracted abdomen, which is folded into a Margin: ma gi a se ae s i ose 1 - 5 μm o g; wi wide-mouthed pocket; the innermost fold is formed at 3 - o eac si e e wee s igma ic c e s; e icu a io about abdominal segment II, which comes to lie beneath se ae is i c y a ge a usua y se s ig y o o o sum; the anterior end of the thorax, about level with the ma gi a se ae e e i g a ou a i o a a -way u prothoracic legs. Another sharp fold is formed across the ma gi o a a c e S igma ic s i es a e s o s ou body approximately between the posterior stigmatic clefts, a u wi a qui e we e e o e asa socke ; e g forming the outer margin of the pocket; and the anal cleft 1-3 μm Eyes o usua y o scu e o ossi y a se and abdominal body margin are also retracted anteriorly, Venter: pregenital disc-pores with mainly 8 loculi pulling part of the abdominal dorsum round to the ventral (range 7-9); number of pores per segment (medially/ surface; the derm between the pocket margins remains mediolaterally on either side): VII, 22-30/19-30; VI, 42— n f l nd 4 2 56/5-9; V, 46-64/4-8; IV, 28-42/3-4; III, 16-22/2-4; (ii) ventral tubular ducts in a submarginal band 4-5 ducts and II, 2-10/0-2; with none laterad to metacoxae. wide (narrower on the other two species); Spiracular disc-pore bands widest near peritreme and (iii) pairs of long pregenital setae on abdominal segments margin; with 18-26 in each anterior band and 16-38 in VI and VII (as on K. perforata); each posterior band; each anterior band with 1 pore (iv) presence of ventral microducts medially on the thorax present a short distance mesad to peritremes. Ventral (as on K. depressa); microducts of 1 type, present in a distinct submarginal (v) extension of the marginal spinose setae up the anal cleft band just mesad to submarginal band of ventral tubular (as on K. depressa); ducts and also frequent medially on all thoracic segments, (vi) reticulate pattern in seven longitudinal rows (as on K. rather less frequent medially on abdominal segments; also depressa); present posterior to mouthparts. Preantennal pores (vii) with a fairly distinct constriction between the pedicel apparently absent. Ventral tubular ducts as for genus: with and segment III on the antennae (as on K. depressa); a distinct marginal area free from ducts; submarginal band (viii) the presence of distinct segmentation or pseudo- of ducts fairly narrow but also extending medially along segmentation in the long `3rd' segment, giving 7- margins of each spiracular disc-pore band; absent segmented antennae (absent on the other two species). elsewhere; smaller ducts, rather short, with a fine inner ductule appearing to arise from side of each cup-shaped Biology. Unknown. invagination and lacking a terminal gland; frequent on either side of anal cleft and extending around anogenital Distribution. This species has only been collected off two area on segments VII and VI. Ventral setae: ventral anal species of Hebe on the one occasion (at Lincoln, South o e se ae a ou 5 μm o g; wi o y a si g e ai o Island) (Map 23). a e io a a c e se ae; wi a ai o o g ege i a se ae me ia y o a omi a segme s II a I; um e o Name derivation. This species was initially mistaken for s o se ae me ia y o eac a omi a segme II 1- ; K. depressa and so the name refers to the similarity: para I - ; -III -9; a II - ; wi se ae ea eac (Gr.) meaning near and depressa, the species it resembles. me aco a -3 ea eac mesoco a a - ( a e s i ose se ae ea eac oco a e g o se ae associated with coxae all short, those near procoxae, 7-9 μm; wi 3- ai s o i e a e a se ae a e a y us 1 K l r p rf r t (Maske ew com i a io se a me ia y; wi -3 su ma gi a se ae o eac si e e wee s igma ic a eas A e ae 7-segme e igs Μ C C 3 119 occasio a y -segme e wi a i is i c Ctenochiton perforatus Maskell, 1879: 208; —Maskell seu osegme ; wi a co s ic io e wee a 3 1884: 130 [taxonomy]; —Maskell 1887: 72 [description]; — segme ; o a e g 39 - 3 μm; e g o a ica se ae Riley & Howard, 1893: 282 [poss. Californian record]; — 5- 1 μm; es y se a ea a e a icu a y a ge e g Cockerell, 1894a: 32,34 [checklist]; —Maskell, 1895a: 13 o c y eo a a s ie 1 -1 9 μm Wi o s i acu a [checklist]; —Cockerell, 1896, 330 [checklist]; —Howard, e i emes a e io 5 - μm os e io 1-7 μm egs 1897: 81 [record correction]; -Fernald, 1903: 161 [world e g s (me a o acic co a 7 - 3 μm oc a e + catalogue]; -Hutton, 1904: 226 [checklist]; -Myers, emu 1 9-19 μm i ia 151-175 μm a sus 7 -1 μm 1922: 199 [checklist]; -Green, 1929: 377 [record]; — c aw 1 - m Gourley, 1930: 7, 10 [parasitoid]; —Fulmek, 1943: 30 [parasitoid]; -Wise, 1977: 104 [checklist]; —Deitz & Material examined: HOLOTYPE [f]: NEW ZEALAND: Tocker, 1980: 31 [checklist]; —Ben-Dov, 1993: 102 MC: Lincoln, Hebe odora, 16 Nov 1967, B.P.J. Molloy, [world catalogue]; —Henderson, 1995: 105 [lectotype DSIR #283, NZAC: 1/1 [f] ad. designated]. PARATYPES: (i) as for holotype, DSIR #283 NZAC: 1/ 1[f] ad. (ii) as previous, but offbrachysyphon,#281,Hebe Unmounted material: young adult female thin, nearly NZAC: 2/2[ff] ad. flat; mature female more convex; test glassy, each plate with lines of air cells radiating from centre; fringe of long Remarks. Adult female K. paradepressa have the curved wax plates rather like the wing feathers of a bird. following combination of characters: Colour transparent to greenish. "Adult female filling the (i) no large macropores on dorsum (present on the other test, shrivelling to the anterior end of the test at gestation." two species); (Maskell, 1887, p. 72). 28 d n & nd r n (2000 : C d (In t : pt r : C d Μou e ma e ia o y o a ; a a c e a ou 1/ o y marginal ducts abundant as a narrow marginal band, length. Old females shrivelled (as above), with margin mostly lying very close to margin; also occasionally characteristically rather sclerotised and scalloped (folded); present on submargin and then more numerous on length 2.0-4.0 mm; breadth 1.75-4.0 mm; posterior abdomen; smaller tubular ducts usually lacking a terminal gland but more anterior ducts sometimes with a Dorsum: dorsal pores in a reticulate pattern, with gland present: abundant on either side of anal cleft and reticulation areas in mainly 5 longitudinal rows (but with extending around anogenital area and with a few two additional reticulation areas split from submedian row anteriorly, particularly mediolaterally on segments IV to on each side, 1 on thorax and 1 on abdomen) and with I e a se ae e a a a o e se ae 31-9 μm o g; probably 7 areas between anal plates and anterior margin with l-2 pairs of anterior anal cleft setae; with pairs of and with 26 areas around margin. Dorsal pores of 4 types: long pregenital setae restricted to medially on abdominal (i) microductules: present throughout, but most frequent segments VII and VI; number of short setae medially on within reticulation lines; (ii) simple pores, slightly larger each abdominal segment: VII, 4-12; VI–III, 4-8; and II, than microductule pore and flat: frequent, within 0-4; with 2-6 setae near each metacoxa, 4-5 near each reticulation lines and near margin; (iii) larger, slightly mesocoxa, and 1-4 near each procoxa; length of setae convex, simple pores: present in an elongate group around associa e wi eac oco a a icu a y o g 5- μm; anal plates but most abundant on either side, with from 8- with 2-4 pairs of interantennal setae laterally plus 0-1 seta 25 laterad to posterior margin of each anal plate; and (iv) medially; with 1-4 submarginal setae on each side heavily sclerotised macropores, convex and slightly between stigmatic areas. Antennae 6-segmented; total sunken; rather distinctive, sometimes rather square and e g 3 -5 μm; e g o a ica se ae - μm often appearing to have a dumbbell-shaped area medially e g o c y eo a a s ie 135-1 μm Wi o when viewed from above under light microscope, but s i acu a e i emes a e io 5- 7 μm os e io 3 -5 bilocular when viewed with the scanning electron μm egs qui e oa ; i io- a sa segme a io o e mic osco e ( ig Μ mos a u a i me ia a i is i c ; e g s (me a o acic co a 7 -11 μm submedian lines of reticulation, becoming less common trochanter + emu 17 -3 μm i ia 115- 7 μm a sus laterally; with 11-35 in posterior medial macropore line. 7 -135 μm c aw 1 - 5 μm A a a es 117-159 μm o g com i e wi s 1 -1 μm; wi - mi u e o es o o sa su ace o eac a e; Material examined: LECTOTYPE [f]: NEW ZEALAND e g o se ae i e ma gi 1 1 -1 μm; i e ma gi [Hawke's Bay]: "Ctenochiton perforatus, from Parsonsia, 11-1 μm; a ica 19- 5 μm a ou e ma gi 1 -1 female-2nd stage without fringe, Sept. 1877, W.M.M.", μm A oge i a o wi ai s o se ae a o g a e io CMNZ: 1/l [f] 3rd moulting into ad; designated by ma gi o ges 3 -5 μm a 1 s o e ai o a e a Henderson, 1995. margins. PARALECTOTYPES: NEW ZEALAND: (i) ex Ma gi ma gi a se ae s i ose 11- 5 μm o g; wi Parsonsia, five females-2nd stage without fringe June 17-40 on each side between stigmatic clefts; marginal 1877, W.M.M., NZAC: 1/5[m] 2nd. (ii) exPanax setae absent along margins of anal cleft or, if present, with [=Pseudopanax], female-2nd stage with fringe, June only 2-3 along outermost margin. Stigmatic spines 1877, W.M.M., CMNZ: 1/1[f] 2nd. (iii) ex Coprosma, head tapering slightly, rather blunt and slightly curved, with a & antenna of male, Nov. 1877 & Dec 1877, W.M.M.: 3/ qui e we - e e o e asa socke ; e g 1-5 μm 3[mm] (2 CMNZ, 1 NZAC). (iv) ex Pittosporum, adult fe- Venter: pregenital disc-pores with mainly 10 loculi; male, Feb. 1878, W.M.M., NZAC: 1/1 [f] ad. (v) ex number per abdominal segment (medially/mediolaterally Coprosma, two tests with old shrivelled females, May on each side): VII, 14-70/9-51; VI, 38-98/8-24; V, 44- 1878, W.M.M., CMNZ: 1/2[ff] ad. 112/3-21; IV, 22-8613-14/; III, 2-22/2-11; and II, 0-18/ 0-5; with 0-4 laterad to each metacoxa. Spiracular disc- Other material: NEW ZEALAND, no host or date, pores in bands 1-4 pores wide; with 17-73 in each band; W.M.M., #35 USNM: 1/1 [f] ad. Mounted from W.M. each anterior band with 0-3 pores present a short distance Maskell's dry material, Box 27, by C.J. Hodgson (1996), mesad to peritremes. Ventral microducts of 1 type: no host or data: 2/2 1st. Mounted from W.M. Maskell's scattered in a broad submarginal band, near mouthparts dry collection #34 & #35 by J.A. de Boer (2 Nov 1972): 3/ and occasionally near each meso- and metacoxa and 4 [ff] ad, 4[m] 2nd, 1 1st. ND: Poor Knights Is, Tawhiti medially on more posterior abdominal segments; absent Rahi, Coprosma sp., 10 Dec 1980, C.F. Butcher, #81-42g: medially on thorax. With a single preantennal pore/ 3/9 [ff] ad. AK: Southern Waitakere Ra, Twin Peaks Tk, antenna. Ventral tubular ducts as for genus but with 440 m, Pittosporum kirkii leaf, 21 Nov 1976, A.R. n f l nd 4 2 Ferguson, #76-334: 1/1[f] ad. CL: Little Barrier I, Hamil- The main features of adult females of K. p rf r t are: ton track, tt p r b ll t , Sept 1994, RCH, (i) the well-defined, narrow, marginal band of ventral #94-078: 1/2[ff] ad. BP: Rotorua, FRI Nursery, tubular ducts, these ducts mainly lying very close to tt p r n d , 30 Oct 1961, R.Zondag, FRNZ the margin, but with a few also generally present on the R(a)71,72: 9/9[ff] ad. Rotoehu S.F., C pr l d , 8 submargin (other two species with a clear marginal Feb 1959, R. Zondag: 4/4[ff] ad. GB: Karakatuwhero V. area free from ducts); Rd, Waipiata, fl r t tr ndr , 28 Sept 1993, RCH, (ii) the absence of ventral microducts from medially on the #93-316a-b: 2/1[f] ad, 1[f] 3rd (+l [m] 2nd unknown sp.). As thorax (present there on both other species); previous, except M l p pl x, #93-317a: l/1[f] ad. (iii) the rather complex form of the dorsal macropores As previous, 1 May 1993, #93-322b, 1/1[f] (pharate). As (absent on K. p r d pr , simple on K. d pr previous, 4 Nov 1993, #93-348: 1/1[f] ad. TO: Pureora (iv) the rather abundant larger simple pores on either side SF, tt p r t rn r (associated hymenoptera card- of the anal plates dorsally (fewer on K. d pr , mounted), Nov 1984, C.F. Butcher, #85-024f: 2/2[ff] ad. smaller on K. p r d pr WN: Wellington, tt p r sp., Oct. 1920, J.G. Myers, (v) pairs of long pregenital setae on the posterior two ΒΜΝΗ 1/3[ ] a 1 [ ] 3 NN:Motueka, Fearons Bush, pregenital segments (as on K. p r d pr tt p r sp., 6 Dec 1931, [no collector], #90-214a-c: (vi) the very Iong 3rd-antennal segment (shorter on K. 3/3[ff] ad (poor), 1 pupa. Motueka, tt p r sp., 21 d pr , with a pseudosegment on K. p r d pr Feb 1932, G. Brittin, #26: 1/1[f] ad. As previous, 28 Jan (vii) long setae medially near procoxae (short on other two 1935, 1/1[f] ad. As previous, 24 Dec 1937: 1/2[ff] ad. species). Whakapuaka, C pr sp. leaves, 3 Dec 1941, no coll., It is close to K. d pr but the latter has: Νo 5 3-3 3/3[ ] a o Mo ueka tt p r (i) 2 types of ventral microduct, the larger forming a ea es 1 Oc 19 3 W A o oway ΝΖ t n f l marginal band between the ventral tubular ducts and (a 1 1/1 [ ] a As e ious ec 19 3 ΝΖ the margin (but absent submarginally); (a 5 11/11 [ ] a e so tt p r (ii) an abundant group of smaller ventral microducts n d , 1 Dec 1967, E. Valentine, #288: 1/3[ff] ad. medially on the thorax Wairoa Gorge, C pr p th l t , 13 Dec 1967, J.A. (iii) marginal setae extending some way up the margins of de Boer, #291: 1/1[f] ad. Maitai R, C pr sp., 19 Dec the anal cleft; 1967, J.A. de Boer, #314: 1/1[f] ad (split dorso-ventrally). the dorsal macropores appearing rather simple and Maitai R, C pr sp., 17 Jan 1968, J.A. de Boer, #342: (iv) lacking the dumbbell-shaped markings; 1/1[f] ad. Nelson, C pr sp., 29 Sept 1968, E.W. Val- entine, #441: 1/2[f] 3rd.BR:Reefton,Gr l n l tt r l , (v) with a distinct clear submarginal area between the band 1914, G. Brittin, #8: 1/19 ad. MC: Christchurch, of ventral tubular ducts and the margin, with no ventral tt p r sp., R.G. Hamilton, 17 May 1939: 2/1[f] 3rd, tubular ducts on the marginal areas; 4[m] 2nd. Christchurch, Orchard belonging to Hamilton, (vi) with a pair of long pregenital setae on abdominal tt p r sp. leaves, 17 May 1939, G. Brittin dry col- segment VII only; lection, #94-057a-d: 4/8[m] 2nd, 5[ff] 2nd, 1 [f] 3rd. (vii) small ventral tubular ducts on either side of anal cleft Christchurch, Riccarton Bush, tt p r n d and anal plates much less frequent than on K. underside leaves, 26 Sept 1997, RCH, #97-135a--j: 10/1[f] p rf r t . ad, 2[f] 3rd,6[m]2nd, 11 pupae, 3[m] ad. Lyttelton, Corsair Bay, 48 Park Tce., C pr r p n , 0 Oct 1976, T. Hay: For a comparison with K. p r d pr see under that 1/1[f] ad (good). As previous, 14 Mar 1977, #77-89a: 2/ species. 8[ff] ad (poor). DN: Totara, r t t , Nov K. p rf r t (as Ct n h t n p rf r t was record- 1926, G. Brittin, #26: 1/1[f] ad. Oamaru, r ed as having been introduced into California by Riley & t t ., 12 Aug 1913, G. Brittin, #26: 1/1 [m] ad.FD: Howard (1893) but this record was later corrected by Manapouri, tt p r sp., 17 Mar 1984, no coll., #94- Maskell who indicated in a letter to Howard (1897) that it 054: 1/2[ff] ad. was probably a dactylopiid! Remarks. In addition to the above hosts, Maskell (1887) Variability. This material is very variable in size and this also recorded K. p rf r t off b sp. (Rubiaceae) and also affects the frequency of the pores. Some specimens r [= d nt r ] sp. (Winteraceae) and from the off C pr are easily the smallest (e.g., #81/42), whilst following localities: WN: Wellington; NN: Nelson; MC: the largest are off tt p r (e.g., Brittin, #26); Riccarton Bush; DN: Dunedin. however, some specimens off C pr are only slightly 0 d n & nd r n (2000 : C d (In t : lpt r : C d smaller than those off Pittosporum sp. Nonetheless, the iag osis. A u ema e. Note: this genus contains two basic characters of the species given above remained groups of species, (i) the L. actites spp.-group: L. actites n. constant. It is clear that there is considerable intra-specific sp. and L. metrosideri Maskell , and (ii) the L. minor spp.- variation in some of the numerical values, probably due to group: L. minor Maskell and L. scutellaris n. sp. These host-induced or climatic effects. two groups differ in a number of significant ways (see under Remarks below) but combined, they form a taxon (a a oge s a a asi oi s. Hymenopterous parasitoids single genus) that is very different from all other New recorded are: Aphelinidae: Euxanthellus philippiae Zealand genera. In the following generic diagnosis, the L. Silvestri; Encyrtidae: Adelencyrtoides blastothrichus actites spp.-group and the L. minor spp.-group refer to the Noyes; A. variabilis Noyes; Pteromalidae: Aphobetus species pairs above. nana ( ouček Test: either absent or represented by a thin glassy wax covering. io ogy. K. perforata is rather polyphagous, although S a e: convex but flat on top, "having the general Pittosporum species appear to be favoured hosts. appearance of an overturned basket" (Maskell, 1882); Probably with one generation a year, overwintering as fully mature female rather round but with a distinct anal nymphs. Post-ovipositional females have a distinctively cleft; dorsal surface sclerotised, with a central, flat, oval contracted abdomen, which is folded in a concertina area which represents true dorsum, surrounded by a wide fashion towards the anterior thorax, while the body margin formed from venter (the L. actites spp.-group: margins become differentially sclerotised and lateral margin <2.5 x wider than true dorsum; the L. minor characteristically folded in scallops. spp.-group: lateral margin <1/8 total width of true dorsum). Mounted material of young, pre-reproductive is i u io . In lowland forest throughout (Map 24). adult oval, with no stigmatic clefts and a fairly shallow anal cleft; length 0.5-l.0 mm, width 0.4-0.8 mm. As adult female matures, size of true dorsum remains constant and lies medially, whereas venter expands laterally so that its submargin lies dorsally and becomes the de facto margin of dorsal surface; mature specimens up to 2.0 mm long and 2.2 mm wide, with a deep anal cleft (Fig. 33). o sum: derm of true dorsum in central area of young adult sclerotised, lateral dorsal surface at first membra- nous, becoming uniformly sclerotised on older specimens, with an anal sclerotisation. Dorsal setae entirely absent. Dorsal pores of 2 ( the L. minor spp.-group) or 3 (the L. actites spp.-group) types: (i) small simple pores: present or absent; (ii) rather larger microductules (possibly absent in the L. minor spp.-group); and (iii) large, sunken, macro- pores, referred to as `pocket-like macropores': present in 2 Ge us ECA OC I O Maske distinct lines extending anteriorly from anal plates towards a point either halfway between the anal plates and the head Lecanochiton Maskell, 1882: 221; —Atkinson, 1886: 278 or nearly dorsad to the clypeolabral shield (the L. actites [taxonomy]; —Maskell, 1887:62, 64 [description] — spp.-group) and also in 8-10 radial lines (the L. minor Cockerell, 1894b: 1053 [distribution]; —Cockerell, 1896: spp.-group). Anal plates elongate, narrow, with outer 330 [checklist]; —Cockerell, 1899c: 332 [key]; - margin more or less rounded; each plate with 1-2 small Cockerell, 1900: 368 [mention]; —Fernald, 1903: 147 apical setae, a longer seta on outer margin and another near [world catalogue]; -Hutton, 1904: 226 [checklist]; - middle of inner margin; also generally 0-3 minute pores MacGillivray, 1921: 172, 178 [catalogue]; -Morrison & medially on posterior third of each plate. Anogenital fold Morrison, 1922: 69, 71 [redescription]; -Lindinger, 1932: with a pair of supporting bars and 3-4 pairs of setae along 197 [mention]; -Borchsenius, 1957:48 [mention]; -Wise, anterolateral margins. Anal tube of moderate length; anal 1977: 105 [checklist]; -Ben-Dov, 1993: 155 [world ring with 6 setae. catalogue]; -Hodgson, 1994a: 306 [redescription type]. Margin: marginal setae entirely absent, although Type species: Lecanochiton metrosideri Maskell, 1882, ventral submarginal setae may briefly appear to be by monotypy. marginal during rapid expansion phase (the L. actites spp.- n f l nd 4 group). The . n r pp.-group with a distinct marginal segment apparently fleshy, some even slightly swollen row of quinquelocular disc-pores or simple pores; these apically and distorted. Mouthparts typical of family. absent in the . t t spp.-group; however, both the . Spiracles very small, posterior pair pointing posterolaterally t t pp. -group and the . n r spp.-group with a on young adults and posteromedially on mature adults and marginal row of microductules. Stigmatic clefts and opening into abdominal concavity formed by medial stigmatic spines absent (except a single spine on one invagination of abdomen. Legs entirely absent. Vulva specimen of . t ll r . Eyespots possibly present in clearly visible on some young adults, apparently lying . t ll r , otherwise absent. between 6th and 7th visible abdominal segments. Venter: derm of young adult lightly sclerotised and ema ks. This genus contains four p : n h t n expanding very considerably outwards and therefore with t t Henderson & Hodgson, . s ., . tr d r some ventral structures coming to lie dorsally; that part of Maskell, . n r Maskell, and . t ll r Henderson derm that lies dorsally becoming moderately sclerotised at & Hodgson . s . Species in this genus are characterised maturity. On young adults, venter finely folded along by: band of ventral tubular ducts, so as to allow for expansion (i) the way that the venter swells during development, so — shown by arrow in Fig. 122. A cavity is formed that the lateral margins come to lie on the dorsal medially beneath abdomen and thorax, enclosed laterally surface, laterad to the true dorsum which does not by a large pair of distinct mediolateral abdominal folds, itself change in size; which extend from anogenital fold to each posterior (ii) the complete absence of legs; spiracle (the . t t spp.-group) or to each anterior (iii) the reduction in the size of the antennae; spiracle (the . n r spp.-group). Pregenital disc-pores (iv) the restriction of the spiracular disc-pore bands to the with 5 loculi: present in a mediolateral line extending from anterior spiracles only, and this band extending onto lateral margins of anogenital area to each posterior dorsal surface on mature females; spiracle, with 1 pair (the . n r spp.-group) or 2 pairs (v) the distribution of the pregenital disc-pores in a line (the . t t spp.-group) of disc-pores per abdominal segment and with a small group laterad to anogenital area between the anogenital fold and the posterior spiracle; (the . t t spp.-group). Spiracular disc-pores with 5 (vi) the formation of a large brood chamber ventrally loculi: in broad bands (in young adults) extending laterad beneath the thorax and abdomen, with the from anterior spiracles towards true margin on dorsum, development of large mediolateral folds; thus lying on ventral surface and partially on dorsal (vii) the absence of marginal and stigmatic setae; surface on older adults when band becomes stretched and (viii) the presence of dorsal pocket-like macropores. narrow; posterior bands of spiracular disc-pores absent, As indicated in the generic diagnosis above, species in but with a small group of 1-3 near peritreme of each the genus n h t n can be divided into two species posterior spiracle: in the . n r spp.-group, anterior groups based on structure. The . t t spp.-group bands of spiracular disc-pores also with a few simple contains . t t and . tr d r , while the . n r pores. Simple pores present in the . n r spp.-group, spp.-group contains . n r and . t ll r . These both submarginally and within anterior spiracular pore two groups differ in the following: band; absent in the . t t pp.-group. Ventral (i) the proportion of the dorsal surface composed of the microducts of 2 sizes in the . t t spp.-group, small expanded venter — broad in the . t t spp.-group, microducts medially and large microducts submarginally; narrow in the . n r spp.-group; in the . n r spp.-group only small ventral microducts (ii) the structure of the dorsal macropores: large, heavily present, possibly throughout venter. Ventral tubular ducts sclerotised pocket-like macropores, with sclerotised of 1 type, each with an elongate outer ductule and a thin margins in the . t t spp.-group, rather more inner ductule with a large terminal gland; present in a wide shallow, with a sclerotised margin and with a granular submarginal band which spreads laterally onto dorsal inner surface in the . n r spp.-group; surface on older females; more medial ducts slightly (iii) the distribution of the dorsal macropores: restricted to broader than those nearer margin. Ventral setae small, the two median longitudinal lines of pores in the . with 4-5 on each posterior lobe and 1-2 submarginal rows t t spp.-group, quite widespread throughout much in the . t t spp.-group but not thus in the . n r of dorsum in the . n r spp.-group; spp.-group; present medially in rows across abdominal (iv) presence of a pair of eyespot-like areas (occasionally 1 segments, and very few elsewhere; usually with 1-2 pairs or none) on dorsum near anterior margin in the between antennae. Antennae reduced, segmentation . n r spp.-group, absent in the . t t pp. obscure but probably 3-or 4-segmented; all setae on apical group; 2 d n & nd r n (2000 : C d (In t : pt r : C d (v) the number of pairs of pregenital disc-pores on the simple pores or quinquelocular disc-pores forming a mediolateral folds of the abdomen: 2 pairs on the . distinct marginal row 2 t t spp.-group, a single pair on the . n r spp.- —In mature adult, venter expands to <2 x width of true group; dorsum; dorsal pocket-like macropores present in (vi) the pregenital disc-pores forming a group on either radiating lines from median area of dorsum; each side of anogenital fold in the . t t spp.-group, dorsal macropore not heavily sclerotised; pregenital absent there in the . n r spp.-group; disc-pores in a single line between anogenital fold and (vii) without a distinct marginal row of pores in the . each posterior spiracle; with either simple pores or t t spp.-group, but with either 5-locular disc-pores quinquelocular disc-pores forming a distinct marginal or simple pores forming a distinct marginal row in the ow 3 . n r spp.-group; (viii) with only 5-locular spiracular disc-pores in the 2 Median bands of dorsal pocket-like macropores anterior band in the . t t spp.-group, but with extending from anal plates anteriorly to near dorsad of both 5-locular disc-pores and simple pores in the clypeolabral shield; antennae about 3 or 4 times as anterior spiracular pore band in the . n r spp.- long as broad, 3rd segment about half total length; group; spiracular disc-pores in anterior band abundant and (ix) ventral microducts of 2 sizes in the . t t spp.- clearly extending to true margin tr d r group but 1 size in the . n r spp.-group. —Median bands of dorsal pocket-like macropores Fig. 33 shows the comparative sizes of some of the extending anteriorly from anal plates, but only stages in the life cycle of . t t . This shows that the reaching to about half-way point dorsad to 1st-instar crawler expands to 4-5 times its original size clypeolabral shield; antennae short, about twice as before moulting, and that the 2nd-instar female only grows long as broad, 3rd segment as long as broad; spiracular a relatively small amount, so that the teneral adult female disc-pores rather few, band sometimes not reaching is only marginally bigger than the largest 1st-instar true margin on dorsal surface t t crawler. However, by the time the expansion of the venter is complete, the final size of the insect is several times 3 With a single row of 5-locular disc-pores all round true larger. Thus the size of the mature adult female is about margin between true dorsum and expanded venter; `normal' for a coccid. The reason for this behaviour is a e ae 7 μm o g n r unclear. That the female 2nd stadium may be quite short is —With a single row of simple pores all round true margin suggested by the very few specimens available compared between true dorsum and expanded venter; antennae with 2nd-instar males. In addition to the above lateral expansion, the mid- s o 5 μm o g t ll r ventral part of the abdomen and thorax withdraws upwards to form a large concavity which acts as a brood chamber. This chamber opens, often by means of a distinct tunnel- n h t n t t e e so & o gso ew like opening, beneath the anal plates and between the anal s ecies lobes. The posterior spiracles and the vulva, which appears to be rather further forward than on most other Figs M9, M11, M13, C64, 120-121 Coccidae, open into this space. The species currently placed in n h t n are all Unmounted material: young female light brown and endemic to New Zealand and are all host-specific to the covered in thin wax plates which make insect look hairy; genus M tr d r . old female dark brown, shiny, largest n n n h t n species. Body almost round but with a distinct anal cleft posteriorly, with an opening into brood chamber beneath abdomen. Dorsal surface rather flat; central true dorsum Key o a u ema e n h t n with indistinct radial ridges, only about l/5th total width 1 In mature adult, venter expands to several times width of and about 1/3 total length. Lateral areas of dorsal surface true dorsum; dorsal pocket-like macropores restricted formed from venter apparently without other markings. to 2 lines extending anteriorly from anal plates; each macropore with a heavily sclerotised margin; Mounted material: as for generic diagnosis. Width of pregenital disc-pores forming a double line between true dorsum 0.34-0.48 mm; final size of mature, fully anogenital fold and each posterior spiracle; without expanded female up to 2.30 mm wide. n f l nd 4 ig. 20. n h t n t t e e so .a u ema e. s ., & Mai o gso , awi g you g ema e Α o sa iew o ma u e mou e a u ema e s owi g e e a i e si es o o sum a a e a ma gi s o e a e e e Β os e io s i ac e a C a e io s i ac e. 4 d n & nd r n (2000 : C d (In t : pt r : C d s 2 ma u e ma e 00μ 0μ ig. 2 . iag am o s ow e amou o g ow e wee eac i s a o n h t n t t e e so & o gso . o g i e 00 μm ( o a u ema e , s o i e 0 μm (a o e s ages . n f l nd 4 ig. 22. n h t n tr d r Maske , a u ema e. Mai awi g you g ema e Α o sa iew o ma u e mou e a u ema e s owi g e e a i e si es o o sum a a e a ma gi s o e a e e e Β a e io s i ac e. 6 d n & nd r n (2000 : C d (In t : pt r : C d ig. 2 . n h t n n r Maske , a u ema e. Α os e io s i ac e. n f l nd 4 . 24. n h t n t ll r e e so s ., . a u ema e. & o gso , A c oss sec io oug a u ema e o s ow ow e a e a ma gi s o e e e e a o ecome a o e o sa su ace, w e e icke e a k i e e ese s e ue o sum. A so o e oo c am e e ea e e . 8 d n & nd r n (2000 : C d (ln t : pt r : C d o sum as for genus, with an anal sclerotisation NZAC: 6/6[ ] ad; BMNH: 1/3[ ] ad; USNM: 2/2[ ] ad around anal plates: dorsal pocket-like macropores large, [Brittin manuscript name . l nd ]. Kawau I su ke a ea i y sc e o ise ( igs Μ9 M11 Μ13 M tr d r x l [as pohutukawa], 25 Aug 1955, R.H. o y ese o os e io a o eac me ia o gi u i a Harrison: 2/24[ ] ad. Rangitoto I M tr d r x l o e a ; wi 9-3 (usua y ess a 5 i eac a ; [as pohutukawa], 12 Sept 1976, J. Cox: 2/9 [ ] ad. ese o es e ace a e io y y mo e a e-si e Rangitoto I, M tr d r x l [as pohutukawa] & mic o uc u es w ic a so occu a e s a se y e sew e e M tr d r sp. [as rata] leaves, 19 Dec 1990, C.F. Mo- o o sum o a y i a ia i es; sim e o es a so rales, #90-357a–c, -358a–d: 7/5 [ ] ad, 8[m]2nd, 13 1st. ese i associa io wi me ia i es o o sa Wattle Bay, M tr d r x l [as pohutukawa] stems, mac o o es; sim e o es a mic o uc u es o mi g a 22 Aug 1981, C.F. Butcher, #81-245: 7/21[ ] ad. is i c ma gi a a (mos o ious o you g Cornwallis, Spraggs Monument, M tr d r x l leaf, s ecime s A a a e e g 91-1 μm ea o 28 May 1983, P. Dale, #83-168b: 2/4 [ ] ad. C : Colville si g e a e 3 -3 μm; eac a e wi a si g e mi u e Hill, M tr d r r b t stem, 6 Aug 1957, J.M. Hoy: o e; se a o ou e ma gi oi i g ou wa s a qui e 2/14[ ] ad (l parasitised). : Ohope, M tr d r o g 1- μm; a wi ai s se ae ea a e 9-1 μm x l , 22 Feb 1979, R.J.M. Mckenzie, FRNZ 15 & 16: o g A oge i a o wi ai s o se ae a o g a e a 2/9[ ] ad. Otanga, Lottin Point, M tr d r x l ma gi s os e io ai o se ae mo e s ou a o e ai s twigs, 3 Nov 1993, RCH, #93-366a: 1/1 [ ] ad. G : -33 μm o g Kakanui, base of coastal cliff, M tr d r x l , 22 Ma gi as for genus. Sept 1992, RCH, #93-292: 1/1[ ] ad. I:, [Pohangina e e as for genus, but pregenital disc-pores in a V], Totara Reserve, M tr d r sp. [as rata], 17 Apr 1966, small group of only 1-4 on either side of anogenital fold no collector: 3/8[ ] ad. and with 2 mediolaterally on each abdominal segment. Spiracular disc-pores in broad bands, extending from each ema ks Wi tr d r , this species belongs to the anterior spiracle often only part way to true margin, with . t t spp.-group. Adult females of . t t , 18-35 in each band; posterior disc-pore bands absent but therefore, are similar to, although generally larger than, with a small group of 1-3 disc-pores near each peritreme. those of . tr d r . . t t differs in having: Ventral microducts present medially on head, thorax, and (i) much smaller antennae; anterior abdominal segments, and also with 1 mediolaterally (ii) fewer large, dorsal pocket-like macropores in the 2 on most abdominal segments. Ventral tubular ducts: median longitudinal lines; present in a wide submarginal band which extends (iii) dorsal microductules present within the dorsal lines of laterally onto dorsal surface on mature specimens. sclerotised pores and elsewhere on the dorsum; Antennae reduced to a short stump, only about twice as long as width of scape, segmentation obscure; total length (iv) ventral tubular ducts rarely present medially on 9- 3 μm C y eo a a s ie oa e a o g e g prothorax; 3-5 μm wi 5 - 3 μm S i ac es wi o eac (v) fewer spiracular disc-pores in each pore band. e i eme 11-1 μm egs a se These characters are best seen on young unexpanded females. Ma e ia e ami e . O O Y E [ ] EW EA A AK Okura R, 25 Oct 1993, R.C. Henderson, M tr d r io ogy Nymphs are found on the undersides of leaves x l midrib undersurface leaves & stems, #93-336b, positioned alongside the midribs, but young females move NZAC: 1/3[ ] ad, 1 1st; the youngest [ ]ad next to the 1st- to, and are nearly always found on, the young green stems. instar nymph is designated holotype and clearly marked. Sometimes very old dead females from a previous PARATYPES: (i) remaining 3 specimens on holotype slide. generation persist on old brown stems further from the (ii) as for holotype: NZAC, #93-336a, c–q: 17/12 [ ] ad, host plant's growing tips. There may be more than one 4[m][mm]2nd,ad, 3 pupae, 2 8 1st. generation per year, with the main generation in early summer. Most favoured host plant is M tr d r Other material: n h t n tr d r , New Zealand, x l , a coastal tree often found growing on seaside Maskell coll. #31, USNM: 4/3[ ] ad, 1 1st. : Poor cliffs and beaches and, in fact, . t t rarely found on Knights Is, Tawhiti Rahi I M tr d r x l stem, 6 trees away from the coast, suggesting an ability to tolerate Dec 1980, C.F. Butcher, #81-1 34a: l/1 [ ] ad. Poor Knights salt spray drift. . t t often found in association with Is, M tr d r x l , Dec 1980, #83-327e: 1/1[ ] . t ll r which inhabits the upper leaf surfaces — for ad. AK Auckland, no host, - Feb 1921, G. Brittin #9, example, the Poor Knights Is record of 6 Dec 1980. n f l nd 4 139 is i u io Generally found on North Island eastern anterior margin of anal plates; absent elsewhere. Anal coasts from Northland to the East Cape region but has also plates elongate and narrow, outer margin rounded, length been collected inland, from Pohangina Valley (Map 25). o a es -1 7 μm ea o si g e a e 3 - 5 μm; some specimens with an additional, fifth, seta on dorsal ame e i a io From the noun t t (Gr., m): a surface near apex; outer margin setae quite short, 7-14 dweller near the shore or coast; this species of μm; wi a ica se ae 9-1 μm o g A oge i a o n h t n appears to favour coastal areas. a e a ai o se ae eac 1 -1 μm o g Ma gi as for genus. e e as for genus but pregenital disc-pores in a small group of 5-9 on either side of anogenital fold and in n h t n tr d r Maske groups of 2 mediolaterally on each abdominal segment. Spiracular disc-pores on young, unexpanded adults in Figs C65, 122 broad bands extending from each anterior spiracle to true n h t n tr d r Maskell, 1882: 222; -Maskell, margin but becoming narrow as venter expands laterally; 1884: 129 [male description, distribution]; -Maskell, with 51-75 in each band; posterior spiracles each with 1- 1887: 64 [description]; —Maskell, 1895a: 11 [checklist]; — 3 disc-pores near peritreme; on 1 young adult, a single Cockerell, 1896: 330 [checklist]; —Fernald, 1903: 147 pore is present on 1 side near margin in posterior stigmatic [world catalogue]; —Hutton, 1904: 226 [checklist]; - area. Ventral microducts restricted to medially on head Myers, 1922: 199 [checklist]; —Morrison & Morrison, and thoracic segments. Ventral tubular ducts: in a wide 1922: 69 [redescription]; —Miller, 1925: 32, 64 [host]; — submarginal band, extending laterally onto dorsal surface Hoy, 1958: 185, 187, 199 [hosts; distribution]; -Wise, on older females; also with a sparse band extending 1977: 105 [checklist]; -Deitz & Tocker, 1980: 30 medially just posteriorly to anterior spiracles. Ventral [checklist]; —Ben-Dov, 1993: 155 [world catalogue]; - setae: as for genus. Antennae reduced, probably 4 Hodgson, 1994a:306 [redescription]. segmented, 3rd segment rather long, 3-4 times as long as oa ; o a e g 1-7 μm a ium oa e a o g U mou e ma e ia shiny medium-brown to dark e g 37- 7 μm wi 57- 3 μm S i ac es wi o brown, with two lateral white wax stripes associated with e i emes 13-1 μm egs a se anterior spiracular pore bands and sometimes with a pair of white wax ribbons on anal plates; apparently lacking Ma e ia e ami e LECTOTYPE [ ](here designated): any distinct waxy test. Shape rather round but more NEW ZEALAND: " n h t n tr d r , from Rata, pointed at anterior end and widest medially, with a distinct tests of adult, Jan. 1881, W.M.M.". Maskell coll, NZAC: anal cleft; at anterior end of anal cleft beneath anal plates 1/ 2[ ] ad (old); one specimen is split into dorsum and is a distinct vertical opening through which 1st-instar venter and this is here designated lectotype and is clearly crawlers emerge from brood chamber beneath concave marked. abdomen. True dorsum centrally placed, plate-like and PARALECTOTYPES: (i) remaining specimen on lecto- flat, about ½ total length and 1/3 total width. With type slide. (ii) as for lectotype, "adult female", NZAC: 1/ concentric ridges on lateral areas of dorsally located 1 [ ]2nd; "four females" [remounted by C.J. Hodgson], venter; outermost `pseudomargin' marked by a darker NZAC: 4/2[ ] ad (pharate), 1 [ ]2nd, 1 1st. line. Mainly found on lower leaf surface. Other material: NEW ZEALAND: AK Hunua Ra, Mou e ma e ia as for genus. Width of true dorsum M tr d r sp., Nov 1982, C.F. Butcher, #83-293e: 3/ 0.48 — 0.75 mm; final size of mature, fully expanded 4 [ ] ad. NN: near Karamea, Kohaihai Bluff, M tr d r female up to l.96 mm wide. sp. (rata), 19 Jan 1983, C.F. Butcher, #84-024d & #90- 215b—c: 4/7 [ ] ad, 1 [m]2nd,M tr d r 1 1st. Denniston, o sum as for genus but with a darker, heavily sp. leaves, 19 Jan 1983, C.F. Butcher, #84-024k: 2/4 [ ] sclerotised anal sclerotisation around anal plates. In ad. Garveys Creek Rd, M tr d r sp. leaves, 21 addition, dorsal macropores highly sclerotised and Jan 1983. C.F. Butcher, #83-285b: 2/2[ ] ad. Maruia, pocket-like, in 2 longitudinal lines extending anteriorly to M tr d r sp., Dec 1915 [as Xmas], G. Brittin #93: 2/ a point dorsad to bases of antennae, each line l-3 pores 2[ ] ad. Sewell Peak, = 2500 m (TV Repeater Mast), ex wide and with 20-48 (generally more than 30) pores in M tr d r sp. [as rata], 22 Nov 1984, C.F. Butcher: 19/ each line. Dorsal microductules and simple pores 9 [m]W [ ][ ]ad,2nd,Otira,M tr d r 3 1st. 51 restricted to a submarginal band near margin and around b ll t [as l d ], Jan 1914, G. Brittin #93, USNM: 40 d n & nd r n (2000 : C d (In t : pt r : C d 1/2[ ] ad. Otira, M tr d r sp., 28 Dec 1915, G. Brittin n h t n n r Maske #93, USNM: 1/1[ ] ad. Otira, M tr d r sp., Dec 1915 [as Xmas], G. Brittin #93: 2/2[ ] ad. As previous, mounted Fig. 123 by C.J. Hodgson from Brittin dry collection: 3/3[ ] ad (l n h t n n r Maskell, 1891: 12; —Maskell, 1895a: split) [+ 3/11 lst = uncertain species, population mixed 11 [checklist]; —Cockerell, 1896, 330 [checklist]; - with L. n r]. Franz Josef Glacier, M tr d r Fernald, 1903: 147 [world catalogue]; -Hutton, 1904: 226 b ll t , Se 19 1 A a ΝΖ 7 - 7 3/ [ ] [checklist]; —Myers, 1922: 199 [checklist]; —Morrison & ad, 3[m]2nd. Franz Josef Glacier, Roberts Point Track, Morrison, 1922: 71 [redescription]; —Miller, 1925: 32, 64 M tr d r sp. [as rata], 6 Feb 1983, J.M. Cox, #235: 2/ [host]; —Hoy, 1958: 185, 188 [hosts; distribution]; —Wise, 2[ ] ad. Cook Saddle, M tr d r sp. [as rata], no date, 1977: 106 [checklist]; —Deitz & Tocker, 1980: 30 J.M. Hoy:1l/3[ ] ad. Picnic Pt., M tr d r sp. [as rata], [checklist]; -Ben-Dov, 1993: 155 [world catalogue]. 6 Oct 1955, J.M. Hoy: 1/6 [ ] ad. FD: Breaksea I, M tr d r b ll t leaves, 29 Jan 1996, RCH, #96- Unmounted material: dried material light brown, very 077: 1/1[m]2nd.SI:StewartM tr d r I, Oban, Thule, flat; central true dorsum larger than in other known 20 Nov 1969, Kershaw, FRNZ nos 2-3: 2/12 [ ] b ll t , n h t n species, with faint radial lines; with a ad. narrow lateral margin formed from venter. Apparently restricted to lower leaf surface. Remarks. L. tr d r belongs to the L. t t spp.- group. Adult females of L. tr d r appear similar to Mounted material: as for genus but young, pre- those of L. t t but . tr d r has: reproductive adult roundly oval; length of dorsum 0.77- (i) much larger antennae; 0.90 mm, width of dorsum 0.84-0.92 mm; final size of (ii) many more dorsal macropores in the two medial mature, fully expanded adult up to 1.30 mm long and l.22 longitudinal lines; mm wide. (iii) dorsal microductules absent within the dorsal lines of sclerotised pores; Dorsum: as for genus but with 2 median longitudinal (iv) many more spiracular disc-pores in each anterior pore lines of dorsal pocket-like macropores fairly distinct and band. with radial lines sometimes rather short and not always These characters are best seen on the young, unexpanded extending to near true margin. Simple pores present on teneral females. dorsum within radial lines of macropores. With a pair of In addition to the above localities, Hoy (1958) pale areas near anterior margin anterior to antennal bases recorded it from AK: Auckland; Kawau Is; CL: Colville; on most specimens. Anal plates as for genus, length 86-91 RI: Pohangina Valley; BR: Maruia; WD: Wilberg Range; μm wi o si g e a e 3 -3 μm; some wi a Franz Josef; Cook Saddle; Kokatahi Gorge; OL/CO: additional fifth seta on dorsal surface near apex; setae on Devils Staircase, Lake Wakatipu; SL: Waimea Forest; ou e ma gi s o 1 -1 μm o g; se ae ea a e e y Tautuku Bay; FD: Alton Valley and Milford Sound. Also s o 5-9 μm o g A oge i a o a e a ai o se ae from M tr d r x l and M. r b t , in addition to i e 7-1 μm o g M. b ll t . However, some of these records could refer Margin: as for genus. With a complete row of to L. t t , particularly the record from Auckland off M. numerous 5-locular disc-pores around margin of true x l . dorsum, these pores generally about 1.5 x size of 5-locular spiracular disc-pores. Simple pores and small Biology. According to Hoy (1958), this species is more microductules also present, scattered within this row; on likely to be found on the smaller stems when there are large sclerotised specimens, ductules of microductules appear- populations of the diaspidid An pl p tr d r ing as clear areas in a dorsal submarginal row. Maskell, and on the leaves when these diaspidids are less Venter: as for genus, except brood chamber clearly abundant. extends into thorax and head as there are distinct dermal folds laterad to spiracles and antennae. Spiracular disc- Distribution. L. tr d r is common, especially on pores: with 26-35 disc-pores in each anterior band. southern rata (M tr d r b ll t , in lowland forest Simple pores present within anterior bands of spiracular throughout the West Coast of the South Island, including disc-pores and also scattered throughout. Ventral Fiordland and on Stewart Island. There is only the one microducts as for genus. With a submarginal row of collection in NZAC from the North Island (the Hunua minute setae, 2 pairs of anal lobe setae, other ventral setae Range) (Map 26). as for genus. Antennae reduced and segmentation very n f l nd 4 4 i is i c ; o a e g 73-97 μm a ium a e oa West Coast, South Island. Maskell's type collection site is wi 59- 7 μm S i ac es wi o eac e i eme 1 - uncertain, although amongst Hoy's (1958) locality records 1 μm egs a se he mentions Reefton, 1890, W. Maskell, on M tr d r robusta, and Reefton is close to the localities recorded by Brittin (Map 27). Ma e ia e ami e EC O Y E [ ] (here designated): NEW ZEALAND: " n h t n n r, adult female, 1889, W.M.M.". NZAC: 1/1[ ] ad. PARALECTOTYPES: as for lectotype, NZAC: 3/1 [ ] ad, 2 1st, (1 labelled "female 2nd stage") all uncleared. n hlt n t ll r e e so & o gso ew s ecies Other material: NEW ZEALAND: labelled " n h t n n r, New Zealand, Maskell collection No. 117 USNM": Figs C66, 124 2/1[m]W [ ]ad,2nd.labelled" n h t n tr d r 1 without a wax test. Colour brown. Mask.", Otira, M tr d r sp., 28 Dec 1915, G. Brittin U mou e ma e ia #93, USNM: 1/4[ ] ad. Otira Gorge, M tr d r sp., Dorsum represented by a flat, central plate with faint radial ridges; width of dorsal plate about 1/2 total width and no date, G. Brittin #93, BMNH: 1/6[ ] ad. Otira, length about 3/4 total length. Outer margin roundish but M tr d r b ll t [as l d ], 28 Dec 1915, G. more pointed at anterior end and with slight indentations Brittin, #320: 1/1[m]2nd.sp.M tr d r [as Otira, rata], - Dec 1915, G. Brittin, #93: 2/2 [ ] ad. Otira, M tr d r where spiracular disc-pore bands extend onto dorsal sp., 21 Dec 1915, from G. Brittin dry coll. #93, #90-221: surface. Found only on upper leaf surface. 8/10[ ] ad. Maruia, M tr d r sp., [as rata], 28 Nov as for genus but young, pre- 1935, G. Brittin #93: 1/1[m]2nd. Mou e ma e ia reproductive adult rather round; length of dorsum 0.90- l.00 mm, width of dorsum 0.59-0.74 mm; final size of ema ks . n r and . t ll r (described below), the two species which make up the . n r spp.-group mature adult up to 1.04 mm long and 0.91 mm wide. within n h t n, appear to be rather uncommon, . n r having only been collected on about four occasions o sum as for genus; median longitudinal lines of (and none recently) and . t ll r from only four dorsal pocket-like macropores indistinct but with localities but on several occasions. . n r differs from reasonably distinct radial lines which extend to near margin. Simple pores absent from dorsum apart from a . t ll r in having: (i) a complete band of quinquelocular disc-pores around few in a line on either side of anal cleft. With a pair of pale the margin (rather than simple pores as on . areas near anterior margin on most specimens, perhaps t ll r e ese i g eyes o s A a a es e g 79- μm wi o si g e a e -3 μm; ou e ma gi se ae 1 -1 (ii) the dorsal pocket-like macropores forming two fairly distinct median longitudinal lines, from which the μm o g; se ae ea a e ge e a y o isce a e a ou radial lines of pores usually extend only about half- 5 μm o g A oge i a o a e a ai o se ae ge e a y way to margin (median lines less distinct, but radial o isce a e ecause a a a es s ai a k 1 -1 μm long. lines usually nearly reaching the margin on . as for genus, but with a complete marginal t ll r Ma gi row of simple pores and small microductules; on (iii) in the relatively larger antennae. sclerotised specimens, ductules of microductules appear In addition to the above records, Hoy (1958) also as clear areas in a dorsal submarginal row. One specimen recorded this species from CL: Amodeo Bay; Colville; RI: has a small, curved, distinctly spinose, stigmatic spine in 1 Pohangina Valley; Reefton; W Wilberg Range; anterior stigmatic area. a ose ; Cook Sa e a Koka a i Go ge Α so om e e as for genus, except brood chamber clearly o M tr d r x l , M. r b t and M. b ll t . extends into thorax and head as there are distinct dermal However, some of these records could refer to . folds which appear to be mesad to spiracles and antennae; t ll r , described below. also anterior margin of anal cleft heavily sclerotised. io ogy Nothing known. Spiracular disc-pores: with 27-49 pores in each anterior band. Simple pores few, in a wide submarginal band; is i u io All the collections represented in NZAC by otherwise present in (i) a submedian band, (ii) within verified slides are from localities within a small area of the anterior spiracular pore band and (iii) commonly between 142 d n & nd r n (2000 : C d (In t : pt r : C d groups of ventral setae laterad to anogenital area; Ge us UMIC I O e e so & o gso apparently absent medially. Ventral microducts as for genus. With 1 pair of ventral anal lobe setae. Antennae ew ge us reduced with very indistinct segmentation; total length 55- 1 μm a ium a e oa wi -5 μm y e s ecies l h t n p ll n Henderson & S i ac es wi o eac e i eme 1 -13 μm egs Hodgson, ew s ecies (here designated). a se iag osis A u ema e es of thick, glassy plates with a broad fringe of thick white wax plates and with very Ma e ia e ami e O O Y E [ ] NEW ZEALAND: thick blocks of wax covering median dorsum. Colour Lottin Point, Otanga, 27 Apr 1993, R.C. Henderson, varying from green-, red-, or gold-browns to black. M tr d r x l , #93-277b, NZAC: 1/3 [ ] ad (middle S a e oval to almost round, convex when mature; specimen designated holotype and clearly marked). stigmatic clefts shallow, with long or medium-short PARATYPES: as for holotype: (i) other 2 females on holo- stigmatic spines; anal cleft rather wide; venter of female type slide, and (ii) NZAC, #93-277a, c-h: 7/5 [ ] ad, 3[m] becoming rather concave between mediolateral lobes of 2nd, 2 pupae. abdomen at maturity, to form a brood chamber for neonate larvae. Other material: NEW ZEALAND: ND: Poor Knights Is, o sum derm membranous. Dorsal setae absent. Tawhiti Rahi, M tr d r x l leaf, 6 Dec 1980, C.F. Dorsal pores usually delineating a distinct pattern of Butcher, #81-134a: 1/1[ ] ad (teneral). C Amo eo Bay, reticulation areas, with 5 rows of reticulation areas across M tr d r x l , 6 Aug 1957, J.M. Hoy: 2/11 [ ] ad, thorax and abdomen (discernible only on submargin on . 2 1st [as . n r]. Lottin Point, Otanga, upper sur- l rp , median and submarginal rows broad and face of leaves f M tr d r x l , 3 Nov 1993, RCH, between these submedian row narrower, and with 2, 4, or #93-366b-1 & #95-149: 11/3[m][ ]ad,2nd, 3 5 lst, 2 pu- 7 plates between anal plates and anterior margin. Dorsal pae, 1[m]ad. As previous, but dated 15 Mar 1994, #94- pores of at least 3 types: (i) microductules, with a dark pore 045a—d: 4/2[ ] ad, 4[m]2nd, 13 1st. and long inner filament with a small balloon-like proximal end: most common in reticulation lines; sometimes larger ema ks . t ll r is in the . n r spp.-group and near margin; (ii) small, flat, simple pores: within is closely related to . n r. For a comparison of the two reticulation lines and often enlarged near margin, forming species, see under . n r. a distinct submarginal band (but few on submargin between reticulation points in . l rp also present io ogy Found only on the upper surfaces of leaves of in a line laterad to anal plates; (iii) large simple pores, M tr d r x l growing close to sheltered sea equal in size to a macropore; and (iv) large, concave shores. Apparently has more than one generation per year. macropores, inner base heavily sclerotised, with a broad membranous tube opening to derm surface (absent on . a oge s a a asi oi s Hymenopterous parasitoid fl v . Preopercular pores, dorsal tubercles, and dorsal recorded: Pteromalidae: Aph b t n n ( ouček tubular ducts absent. Anal plates rather elongate, dorsal surface always folded or wrinkled, at least along inner is i u io Has been collected from eastern coasts of margins of plates (wrinkled over most of dorsal surface of the North Island, from Northland to the East Cape region plates on . l rp each plate with 3 long, thick, (Map 28). usually blunt-tipped, spinose setae near apex, and with a short, sharp spinose seta on outer margin. Anogenital fold ame e i a io From t ll (Latin, n. dim.) with 2 large internal supporting plates and with 4-6 pairs meaning a small shield, combined with the Latin adjectival of setae along anterior margin and a single pair laterally. suffix r meaning relating to. The unmounted material Anal ring with 3 pairs of setae. looks as though it is covered by a round shield. Ma gi marginal setae abundant and strongly spinose, rather elongate, short and straight, with narrow basal sockets; reticulation setae usually enlarged, with more pronounced basal sockets; often with a pair of broader and blunter setae laterad to each stigmatic spine; marginal setae absent from margins of anal cleft, except on . fl v where l-2 are present at distal end of cleft; marginal anal lobe setae not differentiated. Stigmatic n f l nd 4 143 clefts shallow but distinct, without stigmatic sclerotisations; (iv) presence of large, wide, concave dorsal macropores stigmatic spines from 2 to 5+ times as long as marginal (absent from . fl v spines. Eyespot present just dorsad to margin. (v) dorsal reticulation areas in 5 longitudinal rows; Venter: pregenital disc-pores with mainly 7-10 outer (vi) with 1 type of ventral tubular duct; loculi (mainly 7 on . d d , shape of central loculus (vii) abundant marginal spinose setae. variable between species; present mainly in groups on Species in the genus l h t n resemble those in mediolateral folds of each abdominal segment, thus U b n h t n n having pregenital disc-pores more or less forming a line between anogenital area and metathoracic restricted to a mediolateral-lateral line on the abdomen spiracles but with a few medially on abdomen on some between the posterior spiracles and the anogenital area. species but never as frequent there as laterally (also present However, U b n h t n species differ from those of medially on thoracic segments one. d d . Spiracular l h t n in having: disc-pores mainly with 5 loculi, in narrow bands between (i) convex (non-sunken) dorsal macropores; spiracles and stigmatic clefts, not extending medially past (ii) pointed spinose setae on the anal plates; peritremes, except on . d d . Preantennal pores (iii) dorsal surface of the anal plates without folds or present or absent. Ventral microducts of 1 type, usually wrinkles; scattered throughout and with some always present near (iv) few setae ventrally on abdomen and these mainly mouthparts but distribution often distinctive for each short; species. Ventral tubular ducts of 1 type; abundant in (v) fewer marginal spinose setae. submarginal band and generally present just posterior to The species currently placed in l h t n are all mouthparts and medially on thorax associated with coxae endemic to New Zealand. (exceptP. p n t t with some ducts opening very close to margin on some species. Ventral setae: with a distinct From pl (Latin, f.) mean- group of anterior anal cleft setae on either side of anal cleft; Ge e ic ame e i a io . ing soft and feathery, referring to the waxy extrusions from hypopygial setae absent; usually with several pairs of long pregenital setae present on segment VII, and often on VI the middle of the dorsum, and h t n (Gr. m.) a tunic or and sometimes also V; with only 1 submarginal seta on garment worn close to the skin. each side between stigmatic clefts; other setae as for subfamily. Antennae well developed, 6-segmented, third segment particularly long, with 0-2 pseudosegments; setal distribution as for subfamily. Mouthparts normal and not displaced. Spiracles normal. Legs well developed, with a separate tibia and tarsus but no articulatory sclerosis; setal distribution as for subfamily; tarsal digitules unequal, 1 Key o a u ema e l h t n broader and longer than other; tarsal campaniform pores 1 With very long stigmatic spines, about 10 x longer than absent; claws without a denticle; claw digitules equal and marginal spines 2 broad. Vulva opening on abdominal segment VII. —Stigmatic spines relatively short, < about 3 x longer than marginal spines 3 ema ks. This genus contains six species: l h t n d d Henderson & Hodgson . s ., . l rp 2 Wide, concave dorsal macropores present; on h p l (Maskell), . fl v (Maskell), . n Henderson & t l nikau Hodgson . s ., . p ll n Henderson & Hodgson . —Wide, concave dorsal macropores absent; on various s ., and . p n t t Henderson & Hodgson . s . hosts other than h p l t l flavus Species in the genus l h t n have the following combination of characters: (i) anal plates with characteristic folds on their dorsal Dorsal macropores about as numerous as dorsal simple surface; pores; dorsal macropores in reticulation lines (ii) anal plate setae spinose, stout and bluntly tipped, throughout dorsum, reaching to near margin resembling a 3-fingered comb on the apex; punctatus (iii) pregenital disc-pores with 7 or more outer loculi, —Dorsal simple pores far more numerous than dorsal mainly present in a line on the mediolateral folds on macropores; dorsal macropores never near margin, the abdomen; mainly present submedially on abdomen. 4 1 d n & nd r n (2000 : C d (In t : pt r : C d Each anal plate nearly rectangular; anal plates with most o sum dorsal pores distributed in a reticulate of upper surface obviously deeply folded, including pattern, as for genus but with 7 reticulation areas between laterally on anterior third of each plate; with pairs of anal plates and anterior margin; number of reticulation long pregenital setae present on abdominal segments areas around margin uncertain; median and submarginal VI and VII; reticulation lines not distinguishable reticulation areas much wider than intervening submedian medially because dorsal pores apparently placed areas. Dorsal pores of 4 types: (i) microductules: most randomly; pregenital disc-pores absent medially from numerous pore within reticulation lines; a few much larger abdominal segments elaeocarpi microductules present submarginally; (ii) small simple —Anal plates broadest anteriorly and tapering towards pores, about size of microductule pores: present within apex; anal plates with deep folding on upper surfaces reticulation lines; (iii) much larger simple pores (Fig. restricted to near inner margins and posteriorly, never Μ ese wi i e icu a io i es i a su ma gi a laterally on anterior third; pairs of long pregenital band close to marginal spines and in a row along margins setae present only on abdominal segment II but with of anal cleft; and (iv) concave macropores, each with a some other setae at least half their length present on deep, heavily sclerotised, base and a wide, non-sclerotised segment VI; reticulation lines clearly present medially tube: restricted to reticulation fines submedially on on dorsum, although less clear laterally; pregenital abdomen; none in posterior medial macropore line. Anal disc-pores present medially on at least some plates: broad, often indented along posterior margin, abdominal segments 5 a e i g o a e ; e g 1 5-19 μm com i e wi s 1 5- 1 μm; wi -9 mi u e o es o eac u e 5 Pregenital disc-pores present on all abdominal segments, surface; dorsal surface along inner margins often folded; medially on all thoracic segments and with a large inner margin and apical setae long, spinose, with rounded group laterad to each metacoxa; with several pairs of i s; e g s i e ma gi -3 μm i e ma gi long pregenital setae on abdominal segment VII; -31 μm a ica se ae o ges 31-55 μm; ou e se ae ventral microducts forming a distinct line near margin s o a s a y s i ose 19- μm o g Wi ai s o diadema anogenital setae along anterior margin and 1 pair laterally, —Pregenital disc-pores only present medially on o ges 5 - 1 μm posterior-most abdominal segments, absent medially Ma gi marginal setae spinose, 11- 3 μm o g; wi from thorax and with few laterad to each metacoxa; 18-26 spines between stigmatic clefts; reticulation setae with only a single pair of long pregenital setae on slightly longer and displaced slightly onto dorsum; abdominal segment VII; ventral microducts not in a marginal setae on margins of stigmatic clefts sometimes distinct line but sparsely distributed in a broad band slightly broader than other marginal setae. Stigmatic marginally pollicinus s i es s ig y o se o o o sum; s o 7-5 μm o g e e pregenital disc-pores with mainly 5-9 loculi; distributed as for genus; number of disc-pores on each segment (medially/mediolaterally): anal cleft: VII, 0-6/ l h t n d d e e so & o gso ew 12-18; VI, 1-5/6-9; V, 2-8/6-10; IV, 5-10/5-11; III 7- 20/7-12; and II 9-15/6-12; metathorax with 7-16 disc- s ecies pores medially and 4-19 laterad to each coxa; mesothorax with 1-8 medially and 2-7 laterad to each coxa; prothorax igs Μ C7 1 5 with 0-4 medially and 0-2 near each coxa. Spiracular U mou e ma e ia test circular or slightly elliptical in disc-pores: with 21-31 in each anterior band and 25-36 in outline, convex, of thick glassy wax plates, with a fringe of each posterior band, each band extending well past broadly triangular segments around edge, extending peritremes. Ventral microducts in a submarginal row but beyond body rather like petals on a daisy; submarginal ring absent from within submarginal band of tubular ducts; also of small, thinner plates allowing reddish colour of insect to scattered throughout rest of venter. Tubular ducts as for show through; median plates very wide, appearing as genus but with a few on head between antennae and near regular transverse bands on young specimens but mouth, and usually with a few near each coxa and coalescing and becoming very thick in old specimens. sometimes with 1 or 2 medially on abdomen. Ventral Colour white and dark red. se ae e a a a o e se ae 3-3 μm o g; wi - pairs of flagellate anterior anal cleft setae; with 1-2 pairs Mou e ma e ia body round-oval. Length 1.48-l.86 of longer setae amongst rows of moderately long setae on mm, breadth 1.03-1.55 mm. a a omi a segme s o ges 53- 5 μm; o a um e n f l nd 4 4 . 2 . l h t n d d nd r n & d n, n. p., d lt f l . 46 d n & nd r n (2000 : C d (In t : pt r : C d . 26. l h t n l rp (Maske , . com ., a u ema e. n f l nd 4 4 . 2 . l h t n fl v (Maske , . com ., a u ema e. 48 d n & nd r n (2000 : C d (In t : pt r : C d . 28. l h t n n e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . n f l nd 4 4 ig. 2 . l h t n p ll n e e so & o gso , . s ., a u ema e. 0 d n & nd r n (2000 : C d (In t : pt r : C d ig. 0. l h t n p n t t e e so & o gso , . s ., a u ema e. n f l nd 4 of setae medially on each abdominal segment: II 5-7; species have long stigmatic spines), while . fl v differs I 8-10; V, 7-10; I 9-13; III 5-14; and II 1-8; with from . d d in having no dorsal macropores. . 3-10 setae near each metacoxa, 1-6 near each mesocoxa d d shares the presence of long claws with . and 3-4 near each procoxa; with 4-6 (total) interantennal p n t t . se ae A e ae -segme e 3- 57 μm o g; a ica se a 3-3 μm o g C y eo a a s ie 13 -1 5 μm io ogy . d d appears to favour montane habitats. o g Wi o s i acu a e i emes a e io 3 -3 μm; All the host plants have small leaves that remain on the os e io 3 -3 μm egs e g s (me a o acic co a plant during the winter and all collection sites have periods 1 -157 μm; oc a e + emu 177- 3 μm; i ia 13 - of snow coverage. The type series was found above the 15 μm; a sus 9 -115 μm; c aw a e o g 19- 5 μm tree line between about 1200 and 1350 metres asl, on a prostrate plant of r h b growing amongst rock scree. Ma e ia e ami e O O Y E [ ] EW EA A Young adult females and empty male tests were present in Kaweka Range, Makahu Spur track, r h b l n , midsummer (January) and there was little change by late upperside leaves, 25 Jan 1998, R.E. Beever, #98-003h, summer (March), although the females' wax tests had NZAC: 1/1[ ] ad. grown thicker. The only known collection with juvenile PARATYPES: collection data as for holotype: NZAC #98- stages is from Desert Road in late September. From this, 003a—g, i &j: 9/999 ad. it is concluded that the adult female overwinters and reproduces in the spring. Other material: NEW ZEALAND: Kaweka Range, Makahu Spur track, 1250m, r h b l n upperside is i u io The seven lots of material available were leaf, 14 Mar 1998, Herman, #98-051: 1/1[ ] ad. TO: collected in the southern half of the North Island and Tongariro National Park, Whakapapanui Stm, Tawhai Falls, northern half of the South Island (Map 29). Oz th n l pt ph ll [as C n l pt ph ll ], 25 Jan 1982, C.F. Butcher, #98-004: 1/1 [ ]ad. Desert Road, near ame e i a io The name is from d d (Greek, n.) Waiouru, Ol r n l r f l , 22 Sept 1958, G.B. meaning royal circlet or crown, reflecting the shape and Rawlings & R. Zondag, FRNZ no.R(a)67: 1/2 [ ] ad, 1[m] jewel-like ornamentation of the wax test. As e ious e ce 3 Se 195 ΝΖ os 5 - 5 1 / [ ] ad, 1[ ] 3rd, 1[ ] 2nd, 1 [m]NN:2nd,Lake 3 1sts. Sylvester, Oz th n l pt ph ll [as C n v v ll r ], 29 Apr 1969, J.A. de Boer, no. 514: 1/2[ ] l ht n l rp (Maske ew com i a io ad (one split dorsoventrally). MC Porters Pass, G lth r sp., 7 Feb 1982, C.F. Butcher, #83-320c & #90-254a— Figs C74, C75, 126 b:3/2[ ] ad, 4 1sts. Ct n h t n l rp Maskell 1885: 26; -Maskell, ema ks . d d n be differentiated from all other 1887: 42 [description]; -Maskell, 1892: 17 [adult male]; - l h t n species by: Maskell, 1895a: 12 [checklist]; —Cockerell, 1896: 330 (i) the numerous pregenital disc-pores present medially on [checklist]; -Fernald, 1903: 160 [world catalogue]; - the thorax and on all the abdominal segments, as well Hutton, 1904: 226 [checklist]; -Myers, 1922: 199 as in the 2 mediolateral bands on abdomen; [checklist]; -Miller, 1925: 33, 63 [host]; —Wise, 1977: (ii) the extension of the spiracular disc-pore bands past the 104 [checklist]; —Deitz & Tocker, 1980: 28 [checklist]; — peritremes onto the median thorax. Ben-Dov, 1993: 101 [world catalogue]; -Henderson . d d is similar to . n and . p ll n in 1995: 106 [lectotype designation]. having the dorsal macropores mainly restricted to the abdomen. U mou e ma e ia "Test of adult female oval, nearly . d d closest to . p ll n , with which it shares: circular, black in colour, divided into hexagonal and (i) 7 reticulation areas between the anal plates and the pentagonal segments which are not conspicuous, and of anterior margin; which the median series forms a very slightly elevated (ii) the medium length of the ventral anal lobe setae; ridge somewhat lighter in colour. Test is only slightly (iii) the large dorsal simple pores in the reticulation lines convex. Fringe is very long and conspicuous, segments and in a submarginal row. toothlike" (Maskell, 1887, p. 67). Young adult female . d d differs from . l rp and . n in almost round, although often fitting over curve of branch; having medium length stigmatic spines (the latter two colour speckled grey-brown, apart from a fringe of quite 2 d n & nd r n (2000 : C d (In t : pt r : C d long white wax plates and blocks of dirty-white wax 10;III, 6-11; and II, 6-13; also with 6-15 in a fine laterad medially on dorsum; at first with curls or cones of wax on to metacoxae. Spiracular disc-pores: with 19-40 in each top but these soon wear off. Mature female convex, black, anterior band and 21-42 in each posterior band; bands 1 with only marginal fringe plates and small amounts of pore wide, except near peritremes and at margin. median dorsal wax blocks remaining, otherwise mature Preantennal pores sometimes present. Ventral microducts females devoid of wax plates on elevated sides but with a as for genus. Tubular ducts as for genus but with a few thin wax coating over sclerotised derm. On twig or small medially on head, thorax (mostly near coxae) and branch of host plant. occasionally on abdomen. Ventral setae: ventral anal lobe se ae o g 37-5 μm; wi 7-1 ai s o age a e Mounted material: body shape oval. Length 1.63-2.47 a e io a a c e se ae; wi ai s o o g ege i a se ae mm, breadth 1.02-2.0 mm. o a omi a segme s I a II a some imes a so ; o ges 5-1 μm; o e ege i a se ae o segme s I Dorsum: derm becoming sclerotised at maturity. a II mo e a e y o g; o a um e o se ae me ia y Dorsal pores distributed in a reticulate pattern laterally but o eac a omi a segme II -13; I 7-1 ; 3- reticulation lines obscure medially; possibly with 25-26 11;IV, 2-5; III, 1-6; and II, 1-6; with 3-6 setae near each areas around margin. Dorsal pores of 3 types: (i) metacoxa, 2-5 near each mesocoxa and 2-3 near each microductules: with a large sclerotised pore and very long procoxa; with 2-4 (total) interantennal setae. Antennae inner filament often appearing looped and twisted within wi seu osegme s o a e g 3 - 3 μm; a ica body of slide-mounted young females; present in se a 3 -5 μm o g C y eo a a s ie 173- 7 μm reticulation lines and scattered over dorsum but not on o g Wi o s i acu a e i emes a e io - 5 μm submargin; (ii) very small simple pores, about half size of os e io 55- μm egs e g s (me a o acic co a microductule pores: seemingly randomly distributed; with 1 7-157 μm; oc a e + emu 1 5- 15 μm; i ia 13 - a row of slightly larger pores of both (i) and (ii) in an arc 1 9 μm; a sus 5-1 μm; c aw 15- μm just anterior to anal plates; and (iii) large concave macropores, with a heavily sclerotised base with a Material examined: LECTOTYPE [ ]: NEW ZEALAND: granular surface and a lightly sclerotised tube wall: present labelled: "Ct n h t n l rp , antenna and foot of in a broad, loose band submedially on abdomen and adult female, from El rp , Oct 1884, W.M.M." thorax, sometimes extending in a single row onto head; NZAC: 1/1[ ] ad.; "Entomology Div., DSIR, NZ, W.M. with none in posterior medial macropore line. Anal plates: Maskell Collection"; designated by Henderson, 1995. each plate almost rectangular, with very deep folds over PARALECTOTYPES: (i) as lectotype, except labelled almost entire upper surface including laterally over "Female-2nd stage" (actually an early 2nd-instar male; a e io i ; e g o eac a e 173- 11 μm com i e NZAC: 1/1[m] 2nd). (ii) "Ctenochiton elaeocarpi, test of wi s 15 - μm; wi mi u e o es ese u a e y female 2nd-stage, .... El rp , W.M.M."; gold label isi e wi i o s o eac a e; wi 3 ai s o a a a e as above (actually a whole mount of a 2nd-instar male on se ae ea a e a s ou y s i ose wi ou e i s; a piece of leaf, in good condition and distinctive of the e g s i e ma gi 1 - 5 μm i e ma gi -53 species); NZAC: 1/1 [m]2nd. μm a a ica se a 53- 3 μm; ou e ma gi se ae s o e a e i g s a y s i ose 3 - μm o g A oge i a Other material: NEW ZEALAND: ex Maskell's dry mate- o usua y wi 5 ( a ge - ai s o se ae a o g a e io rial (round box) labelled Ct n h t n l rp , mounted ma gi a 1 ai a e a y; o ges 77-1 5 μm by J.A. de Boer: 2/3 [ ] ad. ND: Tangihua Range, Margin: marginal setae spinose, slim and sharply Horokaka, l h d t underside leaf, 6 Apr 1993, oi e eac 15-3 μm o g; e icu a io se ae s ig y RCH, #98-090a-c: 3/1 [ ] 3rd, 1[ ] 2nd, 1 1st. AK: Waitakere o ge ; wi - 3 s i es o eac si e e wee s igma ic Range, Fairy Falls Tk, l h d t underside leaf, c e s; ose ea s igma ic s i es some imes s ig y 18 Oct 1994, RCH, #98-087: 1/1 [ ] 2nd. Hunua Range, a ge S igma ic s i es o me ium e g 5 -73 μm Mangatangi Reservoir, Workman Tk, W n nn lv l o g leaves, 31 Jan 1998, C.J. Hodgson, #98-088: 19 3rd, 1/ Venter: pregenital disc-pores with mainly 10 loculi 1 [ ]2nd.W n nn BP: Mamaku, Aquarius Rd, (range 8-12), each with a distinctively offset inner loculus r , 1 Se 19 1 M McKe ie ΝΖ 7 1/ associated with a crescent-shaped dark area; restricted to a 1 [m]2nd.27 t Lottin l r , Point,Jan Otanga, line on mediolateral folds on abdomen, extending to 1993, RCH, #93-265: 1/1[ ] 3rd. As previous, except posterior spiracles: number of disc-pores on each side of d r rb r , 27 Apr 1993, #93-0175a—b: 2/1 [ ] ad segment: anal cleft/VII, 15-30; VI, 2-11; V, 4-8; IV, 5— (pharate), 2[m]2nd. As previous, except 3 Nov 1993, #93- n f l nd 4 153 365: 1/1 [ ]ad (old). Te Koau, Bush Walk, twig of (iii) the absence of a clear dorsal reticulation pattern; l h d t r , 15 Mar 1993, RCH, #93-087a-b: (iv) the sclerotisation of the dorsum at maturity. 2/1[ ] ad (old), 4 1sts. As previous, except 30 Sept 1993, . l rp differs from . fl v in possessing #93-320: 1/1 [m] 2nd. As previous, except underside leaves, concave dorsal macropores. It also differs from . fl v 14 Mar 1994, #97-054: 1/1[m] 2nd (split dorsoventrally). and . n in having medium-short stigmatic spines As previous, except Hovell's Watching Dog, #97-053: 1/9 (long on the latter two species) and from . d d , . 1sts. Hicks Bay, Onepoto Bay creek, d r rb r , p ll n , and . p n t t in the absence of pregenital stem and underside leaves, 31 Oct 1994, RCH, #94-108a disc-pores medially on the abdomen. (stem): 1/3[ ] ad, & b-c (leaves), 3/1 [ ] ad (pharate), 1[ ] 3rd (pharate), 1 [m][ ]2nd,2nd. 5 As previous, except #97- io ogy Young females were collected in spring, 055a-b: 2[m][ ]3rd2nd (dorsoventrally split), 2/2 (dorsoven- larvipositing females in summer through autumn, and very trally split). Orete Forest, Te Puia Hut, l h d old, sclerotised females in autumn through winter, the t r leaves, 26 Apr 1993, RCH, #97-052: 1/5 1sts. latter remaining on the host even when dead. The nymphs Rereauira Swamp, Beech Ridge, W n nn r , were most often collected from autumn through winter to 29 Jan 1993, RCH, #93-064a—b: 2/2 [m]2nd (parasitised). late spring, but sometimes also in midsummer. Adult Waiaroho, l h d t r , 29 Apr 1993, RCH, females appear to have been collected in nearly all months #93-121: 1/1 [m]2nd. d r As previous, except of the year but, when the wide span of their ages is taken rb r , 29 Sept 1993, #93-315a—c: 3/2 [ ] ad, 1[m]2nd, into consideration, it possibly signifies a long period of (+1 r p z d r d [m][ ]3rd,ad. pharate); #93-338: 1/1 adulthood and an extended larval production period, As previous, except 3 Nov 1993, underside leaf, #93-359: perhaps resulting in non-synchronous single generations 1/1 [m]ad. As previous, except twig at node, #93-364a—b: per year rather than being multivoltine. At any onetime, a 2/2[ ] ad. GB: Karakatuwhero V Rd, Waipiata, few neonate larvae shelter in the brood chamber beneath l h d t r twig, 28 Sept 1993, RCH, #93- the female's abdomen before emerging from beneath her 319a—c: 3/2[ ] ad (old), 1[ ] 2nd, 1[m]2nd. Pohutu, posterior end. d r rb r , 17 Mar 1993, RCH, #93-305: 1/1 [ ] 2nd (parasitised). As previous, except 28 Sept 1993, #93- is i u io From Northland to Christchurch (Map 30). 318: l/3[m] O 2nd.Rangitoto Station, Mangatutu, l h d t underside leaf, 9 Nov 1996, RCH, #98- 091: 2[m]WI 2nd.Bruce Park, 5km south Hunterville, (Maske ew com i a io d r rb r , 18 Jan 1994, RCH, #98-089: 1/1[ ] l h t n fl v 3rd. Motueka, (no host), 27 Jul 1924, G Brittin, no. 110: 2/1[m][ ]ad,2nd.El rp Nelson, h r n , 1 igs Μ 7 Μ C73 1 7 8 Aug 1968, J.A. de Boer, no. 403: 2/1 [m][ ]3rd,2nd (split 2 Ct n h t n fl v Maskell 1884: 130; —Maskell 1885: 26 dorsoventrally). As previous, except 19 Dec 1968, no. [host]; -Maskell, 1887: 68 [description]; -Maskell, 465: 1/1[ ] ad. Nelson, Ruby Bay, Il t l [as r nth ] 1895a: 13 [checklist]; -Cockerell, 1896: 330 [checklist]; — sp., 20 Nov 1969, J.S. Dugdale, no. 608: 2/2[ ] ad (old). Fernald, 1903: 160 [world catalogue]; —Hutton, 1904: 226 Nelson, Garden's Valley, El rp h r n , 6 Apr [checklist]; —Froggatt, 1921: 23 [taxonomic remark]; - 1973, J.A. de Boer: 2/2[ ] ad (old). MC Lyttelton, MacGillivray, 1921: 178 [catalogue]; —Myers, 1922: 199 C pr sp., 13 Jan 1917, G Brittin, no. 113 (Brittin [checklist]; —Miller, 1925: 33, 63 [host]; -Costa Lima, original slide) & ex dry material Brittin Collection, #94- 1936: 167 [doubtful Brazilian record]; -Lepage, 1938: 060a-e: 6/6[ ] ad. Lyttelton, C pr sp., (no date), G. 351 [doubtful Brazilian record]; —Lindinger, 1954: 619 Brittin, #113, [Brittin manuscript name `Ctenochiton [Brazilian record incorrect]; —Hoy, 1954: 601 [distribu- bicornutum']: 4/3[ ] ad, 1[m] ad. tion]; —Hoy, 1958: 197 [host; distribution]; —Hoy, 1961: 58 [distribution]; —Wise, 1977: 104 [checklist]; -Deitz & ema ks . l rp can be differentiated from all Tocker, 1980: 28 [checklist]; -Ben-Dov, 1993: 101 other l h t n species by: [world catalogue]. (i) the large, deep folds on the anterior dorsal surface of the anal plates (whilst other l h t n species have U mou e ma e ia "Female test golden, waxy, flat folds on their anal plates, these are never present on the beneath, convex above; outline circular or slightly anterior third); elliptical, with a fringe of broadly triangular segments (ii) the lack of a submarginal band of simple pores around the edge. Apex of test an irregular elongated mass dorsally; of wax, remainder divided into two concentric series of 15 d n & nd r n (2000 : C d (In t : pt r : C d plates, inner series pentagonal with sharp angles, outer 7 loculi; with 19-39 in each anterior band and 21-42 in pentagonal with rounded angles and with outer sides each posterior band; bands 1 pore wide except near forming base of segments of the fringe. The inner series peritremes and margin. Preantennal pores present. often forms irregular lumps of wax" (Maskell, 1884, p. Ventral microducts and ventral tubular ducts as for genus. 130). Colour varies from dark and light brown to yellow- e a se ae e a a a o e se ae 3 -53 μm o g; wi brown. The thick fringe of wax plates resemble petals on 5-11 ai s o age a e a e io a a c e se ae; wi -3+ a daisy. ai s o o g ege i a se ae i a i io o se ae o me ium e g o segme s – II o ges 5 -9 μm; Mou e ma e ia body round-oval. Length 1.48-2.76 um e o se ae me ia y o eac a omi a segme mm, breadth 1.0-2.05 mm. II -11; I 5-11; -7; I 3-7; III 0-5; and II 1- 4; with 1-8 setae near each metacoxa, 1-4 near each o sum dorsal pores distributed in a reticulate mesocoxa, and 1-2 near each procoxa; with 2-6 (total) pattern, delineating 4 reticulation areas between anal i e a e a se ae A e ae -3 μm o g; a ica plates and anterior margin and probably with 21 areas se a 17-35 μm o g C y eo a a s ie 1 -173 μm a ou ma gi o sa o es o y es ( ig Μ 7 (i o g Wi o s i acu a e i emes a e io 3 -5 μm mic o uc u es ( ig Μ a ou a si e o sma es os e io -5 μm egs e g s (me a o acic co a sim e o es ese wi i e icu a io i es a wi a 11 -1 μm; oc a e + emu Ι 5 -17 μm; i ia 119- ew o su ma gi a e icu a io a eas a (ii sma 13 μm; a sus -9 μm; c aw 15- μm sim e o es o si es sma es a ou wice si e o mic o uc u e o e ese wi i e icu a io i es; Ma e ia e ami e EC O Y E [ ](here designated): s ig y a ge sim e o es i a su ma gi a ow c ose o NEW ZEALAND: labelled "Ct n h t n fl v , from ma gi a s i es a a o g ma gi s o a a c e ; r h l tt , June 1882, W.M.M."; gold label, "Ento- e icu a io i es wi ma y o es o o y es o e mology Div., SI W M Maskell Collection", o es wi e u a owi g o 1 o e wi e ea ma gi s NZAC: 1/4[ ]ad — lectotype female clearly marked. A a a es e g 1 - 19 μm com i e wi s 15 - PARALECTOTYPES: (i) on lectotype slide above, NZAC: μm; wi -1 mi u e o es o eac u e su ace; 1/3[ ] ad. (ii) as for lectotype slide, NZAC: 3/3 [ ] ad "in u e su ace o eac a e wi ee y w i k e o o e puparia", 1 [m][ ]"puparia",3rd, stained "female 2nd stage" = a o g i e i (a occasio a y o os e io a u and remounted by RCH. e e a e a y o a e io i ; i e a ou e ma gi se ae s o a s a y s i ose a ica se ae s i ose Other material: NEW ZEALAND: from pt p r , longer and thicker, usually with a rounded tip; length of Oct 1884, W.M.M.: 1/1[m]AK ad.Waitakere Ra, Sharp se ae i e ma gi 1 1 -31 μm i e ma gi 1-33 Bush, El rp d nt t leaf, 29 Apr 1995, C #95- μm a ica se ae 33- 3 μm a ou e se ae 1 - 1 μm 044: 1/1 [ ]ad. Riverhead Forest, Barlow Rd reserve, Wi 5- ai s o se ae a o g a e io ma gi o a oge i a M r n tr l leaf, 22 July 1998, C #98-085: 1/ o a 1 ai a e a y; o ges 57-1 μm pupa. Waenga Bush, Otanga, (Lottin Pt Road), Ma gi marginal setae spinose, narrow, straight and r n p t f rr n stems, 3 Nov 1994, C #94- s a y oi e eac 11- 3 μm o g; e icu a io se ae 104a–c: 3/1[ ] ad, 2[m][ ]3rd,2nd,2nd. Waiaroho, 12 s ig y o ge ; wi 1- ai s o o icea y s ou s i es l h d t r , 26 Apr 1993, RCH, #93-185: 1/[ ] e e i g u ou e mos ma gi s o a a c e ; wi 19-35 2nd. Ruatahuna, Whakatane R, (N 44.35.42 [NZFS no.1), s i es e wee s igma ic c e s; ose o ei e si e o d nt r x ll r leaves, 9 Apr 1961, S Dugdale, s igma ic s i es a ge wi a u i 3-3 μm o g FRNZ R12,R13: 2/12[ ] ad. G Pohutu, Awatere R S igma ic s i es o g 13 -19 μm is ace s ig y bridge, d r rb r stem, 1 Nov 1994, C 9 - o o o sum 102: 1/1[ ] ad. Kakanui E-W saddle, head of Waipohatuhatu e e pregenital disc-pores with mainly 10 loculi Stm, p n nd n , 22 Sept 1992, C #92-31 2a– (range 8-10), most abundant on mediolateral folds on b: 2/1 [m][ ]ad4 p r ll (infested d pl x ,with fungus abdomen in a line between genital area and posterior 2nd. Kakanui, 300 m, M r n l n , 16 Mar 1993, spiracle but with a few also present medially on abdomen: J.W. Marris, #93-083: 1/1[ ] ad. As previous, except 30 number of disc-pores (medially/mediolaterally on each Apr 1993, C #93-083b: 1/1[m] K ad.Awakau Road, side) on each segment: anal cleft/VII, 0/13-40; I 0-5/3- W n nn r leaf, 12 Dec 1993, C #93-372: 9; V, 0-3/5-10; IV, 1-2/6-9; III 1II -3/3-9;0-2/3- and 1/1[ ]ad.22W n nn TO: Pureora r , Forest, 9; metathorax with 0-2 disc-pores medially and 3-9 Mar 1960, J.M. Hoy: 1/1[ ]ad.M l t NN: Tarakohe, laterad to each coxa. Spiracular disc-pores with mainly 5– r fl r , 14 Aug 1925, G. Brittin, no. 215: 4/4 [ ] ad, n f l nd 4 4[ ] 3rd, 2[m]2nd.sp., tt p r As previous, ex except the warmer months of the year and . fl v probably dry material Brittin collection, #94-058a—c: 3/2 [ ] ad overwinters as the adult female. Much honeydew is (pharate), 3[ ] 3rd, 1[m] 2nd. Buller, near Millerton, produced by colonies of this insect; this was particularly W n nn p., 19 Jan 1983, C.F. Butcher: 1/1[ ] ad. Α: notable in Fiordland, where the host trees were blackened Kaikoura, Half Moon Bay, d r rb r stems, 15 with sooty mould living on the honeydew. Jan 1998, C.J. Hodgson, #98-092: 1/1[ ] ad. BR: Buller Gorge, d nt r l r t , 17 Mar 1971, J.A. de Pathogens and parasitoids. . fl v is susceptible to Boer, no. 724: 1/1[ ] ad (+Inglisia p t ll [ ]ad). Punakaiki, attack by the orange-coloured parasitic f n p r ll b ( ll pt ? , 25 Oct 1970, J.A. de Boer, no.650: 1/ d pl x (Berk.) Petch. 2[ ] ad (split dorsoventrally). MC: Riccarton Bush, d nt r l r t [as r ], 30 Dec 1916, G. Distribution. Throughout, from Auckland in the north to Brittin, no. 108: 1/1[ ]ad. As previous, except 17 Mar Fiordland in the south (Map 31). 1917: 1/1 [ ]ad.FD,Bligh Sound, Wild Natives R, W n nn r , stems and leaves, 21 Jan 1996, RCH, #96-048a—n: 14/11 [ ] ad (2 pharate), 3[ ] 3rd, 1[ ] l h t n n e e so & o gso ew s ecies 2nd, 2[mm] ad, 4 pupae, 3 prepupae, 6[m]2nd, 5 1sts. Bradshaw Sound, Precipice Cove, W n nn r , 28 Jan 1996, RCH, #96-085a—d: 4/1[ ] ad, 4[ ]3rd,2nd, 2 Figs C78, 128 1 [m] 2nd. Unmounted material: glassy wax test convex and oval, with thick plates, fused into 3 longitudinal rows Remarks. . fl v can be differentiated from other dorsomedially; raised sides formed from submarginal row l h t n species in: of plates with fringe plates apparently fused together; base (i) the absence of concave dorsal macropores; of test broad with a marginal footplate of gum-like wax (ii) in having only 3-4 reticulation areas between the anal which sticks test to leaf substrate; colour variable, from plates and the anterior margin; light green and light brown. Positioned along length of (iii) the presence of long stigmatic spines (shared with . leaf on either leaf surface. n . . fl v shares with . l rp the character of Mounted material: body shape oval. Length 2.13-2.90 long ventral anal lobe setae, and with . d d , . mm, breadth 1.84-2.49 mm. p ll n , and . p n t t the presence of pregenital disc-pores medially on the abdomen. Dorsum: dorsal pores distributed in a reticulate This species was also recorded off W n nn pattern, delineating 2 reticulation areas between anal r from Pohangina Valley (RI), Tamaki River plates and anterior margin (one very large) and with (RI), and Waituhi Saddle (TO) by Hoy (1958). As no perhaps 21 areas around margin. Dorsal pores of 3 types: other species of l h t n has been recorded off (i) microductules: present in reticulation lines and W n nn , these records seem likely to be . fl v , submarginally; (ii) small simple pores, slightly larger than although no material is available to confirm them. Hoy microductule pore: also present in reticulation lines and (1958) also mentions another species ("near fl v " off submarginally; reticulation lines with many pores of both W. r which had an "unusually small number of types, often 2 pores wide, but narrowing to 1 pore wide pores in the spiracular canallae". It was possibly a 3rd- towards margin; and (iii) large concave macropores, with instar female C. fl v , although again no material is a heavily sclerotised base which appears granular with thin available. walled tube; this pore appears to become more heavily . fl v (as Ct n h t n fl v has also been re- sclerotised with age; restricted to median and submedian corded from São Paulo, Brazil (MacGillivray, 1921; Costa reticulation lines on abdomen; with none in posterior Lima, 1936; Lepage, 1938). As pointed out by Lindinger me ia mac o o e i e A a a es e g 19 - 3 μm (1954), the hosts quoted by these authors are those from com i e wi s 19 - μm; wi 1-9 mi u e o es o New Zealand NOT Brazil and so nothing more is known eac u e su ace; o sa su aces o eac a e about this record. It seems highly improbable that it was . distinctively folded or wrinkled but never laterally on fl v . anterior third; each plate with 3 spinose setae near apex, of rather uniform width and usually with rounded tips, Biology. It is not known whether . fl v is uni- or e g i e ma gi 1 1 - μm i e ma gi -33 multivoltine, but early- instar nymphs are present mainly in μm a ica se ae - μm; ou e ma gi se ae se o 6 d n & nd r n (2000 : C d (In t : pt r : C d dorsal surface of plates, short and sharply spinose, length sapida leaves, 31 Jan 1993, RCH, #93-060a j: 10/8 [ ] 1 -33 μm A oge i a o wi usua y 5 ai s o se ae ad, 7 1sts. As previous, except 15 Mar 1993, #95-154a—c: a o g a e io ma gi a 1 ai a e a y; o ges se a - 3/1 [m][ ]ad,2nd. 4 As previous, except 30 Sept 1993, #95- 115 μm 155, l/1 [ ]ad. Waenga Bush, Lottin Point Road, Margin: marginal setae spinose, straight, sharply Rhopalostylis sapida, 27 Jan 1993, RCH, #95-152a—b: 2/ oi e 11-15 μm o g; e icu a io se ae s ig y o ge 1[ ] ad, 6 1sts. a wi a ai o a ge u i e s i es o ei e si e o s igma ic s i e e g 1 - 3 μm; wi 5-3 s i es o Remarks. P. nikau is similar to all other Plumichiton eac si e e wee s igma ic c e s Wi a su ma gi a species other than P. flavus in having dorsal concave ow o sim e o es c ose o ma gi a s i es a a mos as macropores but it differs from all other Plumichiton ume ous a wi a ow o mo e ume ous species in: mic o uc u es c ose o a us o sa o sim e o es (i) having very short ventral anal lobe setae; S igma ic s i es o g 1 7-15 μm o g o se s ig y (ii) the near absence of pores medially on the dorsum; o o o sum (iii) only about two reticulation areas between the anal Venter: pregenital disc-pores with mainly 10 loculi plates and anterior margin. (range 6-10), restricted to mediolateral folds on abdomen It is similar to P. flavus in having very long stigmatic between posterior spiracle and genital area; absent spines, to P. elaeocarpi in lacking pregenital disc-pores medially; number of disc-pores on each side of segment: medially on the abdomen, and to P. pollicinus in having anal cleft/VII, 25-32; VI, 0-5; V, 3-7; IV, 3-9; III, 3-6; submarginal rows of dorsal microductules and simple and II, 3-9; with 2-12 pores in a line laterad to each pores. metacoxa. Spiracular disc-pores: with 21-38 in each anterior band and 22-42 in each posterior band; each band Biology. Collected only from Rhopalostylis sapida, a 1 pore wide except near peritremes and margin. native palm. The mature female is highly convex, her Preantennal pores present. Ventral microducts and abdomen retracting ventrally to form a brood chamber, tubular ducts as for genus. Ventral setae: ventral anal lobe where a few neonate nymphs are generally found se ae e y s o 7-15 μm o g; wi -7 age a e sheltering during larviposition, before emerging from a e io a a c e se ae; wi ume ous o g se ae o beneath the posterior end. segme s I— II o ges -9 μm a wi some mo e a e y o g se ae o segme ; o a um e o se ae Pathogens and parasitoids. P. nikau is commonly me ia y o eac a omi a segme II -1 ; I 1 - attacked by the orange-coloured parasitic fungus 1 ; -1 ; I -1 ; III -1 ; a II -1 ; wi -7 Hypocrella duplex (Berk.) Petch. se ae me ia y o eac me a o a ; - se ae me ia y o eac meso o a a 1- ea eac oco a; wi -5 Distribution. As this species is uncommon compared ( o a i e a e a se ae A e ae 15- 3 μm o g; with its host plant and then often not found alive (it is a ica se a 19-3 μm o g C y eo a a s ie 13 -15 impossible to identify when parasitised with fungus), it is μm o g Wi o s i acu a e i emes a e io -5 likely that the distribution is wider than records show, but μm os e io 5 -57 μm egs e g s (me a o acic it may be absent from the wetter areas of R. sapida's co a 9 -115 μm; oc a e + emu -1 μm; i ia distribution (Map 32). -1 7 μm; a sus 5 -73 μm; c aw 1 -15 μm Name derivation: after the Maori name for the host plant, Material examined: HOLOTYPE [ ]: NEW ZEALAND: nikau. BP: Te Koau, 243m, Rhopalostylis sapida, 4 Nov 1993, R.C. Henderson, #93-363a, NZAC: l/1 [ ] ad. PARATYPES: as for holotype: NZAC #93-363b—e: 4/9 l h t n p ll n e e so & o gso ew ad, 1 [ ] 3rd, 1 2nd, l ad. [m] [m] s ecies Other material: NEW ZEALAND: AK: Auckland, Titirangi, (Bruce Taylor's property), Rhopalostylis sapida igs Μ 1 C7 C77 1 9 leaf, 22 Nov 1977, A.R. Ferguson, #79-198b: 2/2 [ ] ad. Unmounted material: test of young female slightly CL, Kauaeranga State Forest, Rhopalostylis sapida [as elliptical in outline, convex, composed of thick glassy wax ikau a m ] ea es a 19 o ag ΝΖ plates, with a fringe of broadly triangular segments around (a & 7 /3[ ] ad, 6 1sts. BP: Te Koau, Rhopalostylis margin; mature female highly convex and hornlike, wax n f l nd 4 plates coalescing from margin to form a median rounded medially but with 2-4 laterad of each coxa. Spiracular point, with rough edges where marginal and submarginal disc-pores: with 8-23 in each anterior band and 8-29 in plates join. When on leaf of host plant, inhabits upper each posterior band, pore lines not extending past surface. peritremes. Preantennal pores present. Ventral microducts and tubular ducts as for genus. Ventral setae: Mou e ma e ia body oval. Length 1.04-1.64 mm, e a a a o e se ae 19-3 μm o g; wi - ai s o breadth 0.75-1.30 mm. age a e a e io a a c e se ae; wi a ai o o g ege i a se ae o segme II and sometimes on o sum dorsal pores distributed in a reticulate segment VI, but with some medium length setae on both pattern, delineating 7 reticulation areas between anal II a I o ges 1-9 μm; um e o se ae me ia y plates and anterior margin and with about 17 areas around o a omi a segme s II 7-1 ; I 5-9; 3- ; I margin. Dorsal pores of 4 types: (i) microductules of 2 -11; III 5-8; and II 1-7; with 1-2 setae near each sizes: smaller pores abundant in reticulation lines; larger metacoxa; 1-3 near each mesocoxa and 1-2 near each pores present as a submarginal row; (ii) small simple pores procoxa; with usually 2 pairs of interantennal setae. (subequal in size to larger microductule pore): present in A e ae 1 - μm o g a ica se a 3-3 μm o g reticulation lines; (iii) large simple pores (about size of a C y eo a a s ie 11 -1 μm o g Wi o spiracular disc-pores): present in reticulation lines, in a s i acu a e i emes a e io 3-3 μm os e io 7-3 submarginal row close to marginal spines, and in a row on μm egs e g s (me a o acic co a 9 -1 3 μm; either side of anal cleft and extending around anterior oc a e + emu 1 7-15 μm; i ia -1 3 μm; margin of anal plates; and (iv) concave macropores (Fig. a sus 5-9 μm; c aw s o 13-17 μm Μ 1 eac wi a ee ea i y-sc e o ise ase a a o -sc e o ise wi e ou e u e ese i e icu a io Ma e ia e ami e O O Y E [ ]: New Zealand: C i es su me ia y o a ome o e i a is i c i e Little Barrier I (upper) Valley k Leptospermum ig ag a e ; wi occasio a y 1 o i os e io me ia scoparium, upperside leaves, 6 June 1994, R.C. Henderson, mac o o e i e A a a es o e i e e us os e io NZAC #94-065b: /1 [ ]ad. o wi es a ; e g 1 -177 μm com i e wi s 1 3- PARATYPES: collection data as for holotype: NZAC: #94- 157 μm; wi 1-5 mi u e o es o u e su ace o eac 065a: 1/1[m][ ]ad;2nd; 94-065c, 94-065d, 1/1 1/1 pupa, 3 a e; u e su ace o eac a e s ig y o e o 3rd, 2 1st instars. w i k e a icu a y a o g i e ma gi s e e a e a y o a e io i ; i e ma gi se ae o g a s i ose wi Other material: NEW ZEALAND: C Little Barrier I, ou e i s e g 19- μm; a ica se ae o ges 31- Valley Tk ridge, Leptospermum scoparium upperside 3 μm o g s i ose a u ; ou e ma gi se ae se o o leaves, 17 Sept 1994, RCH, #94-084a-b: 1/1[ ] ad, 1 [m]ad. o sum s o a s a y s i ose e g 1 - 1 μm AK Waitakere Ra, Huia Dam, [no host], 23 Aug 1980, A oge i a o wi ai s o se ae a o g a e io ma gi C.F. Butcher, #80-242d: 1/1[ ]ad.Leptospermum Huia, a 1 ai a e a y; o ges - 9 μm o g scoparium, 19 Sept 1980, C.F. Butcher, #80-269h: 1/1 [ ] Ma gi marginal setae spinose, narrow, straight, ad. Waitakere Ra, Matuku Reserve, Taranga Tk, Kunzea apex mostly rounded to blunt, but some pointed, each 11- ericoides leaf, 27 May 1995, RCH, #95-045: 1/1 [ ] ad. 3 μm o g; e icu a io se ae o y s ig y o ge u Waitakere Ra, Sharp Bush roadside, Kunzea ericoides usua y is ace s ig y o o o sum; wi 15-33 s i es upperside leaves, 22 Mar 1998, RCH, #98-052a—b: 2/2 [m] e wee s igma ic c e s; wi a ai o s ig y wi e a 2nd, 2 prepupae, 2 pupae, 1 1st. As previous, except o icea y u se ae o ei e si e o s igma ic s i es Mountain Rd, lvs & twigs, 7 July 1998, #98-078a—d: 4/ eac 1 - 3 μm o g S igma ic s i es s o a ow o e 2[ ] ad, 1 [m] ad (pre-emergent), 1 [m] 2nd, 2 pupae. G s ig y cu e wi oi e i a s a ow asa socke Paoneone, Kunzea ericoides upper surface leaves, 15 Mar is ace s ig y o o o sum; eac 1 -3 μm o g 1994, RCH, #94-141 a—b: 1/1[ ] ad (pharate), 1[ ] 3rd, 2[m] e e pregenital disc-pores with a highly variable 2nd. Rotorua, Atiamuri Rd, Leptospermum number of loculi ranging from 5-10; mostly on scoparium, 12 July 1959, R. Zondag, FRNZ R94: 2/2[ ] mediolateral folds of abdominal segments extending from 3rd. WI: Flock House, Leptospermum scoparium, 30 Apr pregenital area to posterior spiracles but also with a few 1959, M.A. Stoodley, FRNZ R83-86: 4/22 [ ] ad. SD: medially on abdominal segments; number of disc-pores on Okiwi Bay, Leptospermum scoparium, 17 Oct 1955, J.M. each segment (medially/mediolaterally on each side): anal Hoy, #201: 1/1[ ] ad. Para Swamp, near Picton, cleft/VII, 0-1/4-9; VI, 2-4/2-6; V, 0-3/3-5; IV, 2-4/2-4; Leptospermum scoparium, 9 Aug 1948, T.G. Sewell, #78: III, 0-4/2-4; and II, 0-4/3-5; metathorax with none 3/2[ ] ad (one pharate), 3[ ] 3rd (one pharate), pupa. Rocky 15 d n & nd r n (2000 : C d (In t : pt r : C d Creek Bridge, Leptospermum scoparium, 9 Aug 1948, T.G. l h t n p n t t e e so & o gso ew Sewell, #74: 2/3[ ] ad (one pharate), 3[ ] 3rd, 2[m]M 2nd. s ecies Waihopai [as Waihopeu] Valley, Leptospermum scoparium, 5 Aug 1948, T.G. Sewell: 4/3[ ] ad (pharate), 3[ ] 3rd, 2[m] Fig. 130 2nd, pupa. NN: Nelson, Leptospermum sp., upperside leaves, 10 Apr 1925, Brittin dry collection #71, #98-068a— U mou e ma e ia inhabits upperside of leaf (waxy n: 14/3[ ] 3rd (one pharate), 9 [ ] 2nd, 11 [m]2nd, 2 1st. base pad remaining on leaf in dry collection); nothing else Collingwood, Kunzea ericoides, [as Leptospermum] 3 Aug known. 1948, T.G. Sewell: 5/4 [ ] ad (3 pharate), 4[ ] 3rd, 1[m]2nd. 5 miles north of Westport, Leptospermum scoparium, 23 Mou e ma e ia body round-oval. Length about 2.57 Mar 1960, G McBurney, FRNZ R(a)44-46: 3/5 [ ] mm, breadth about 2.1 mm. ad, 2[ ] 2nd, 4BR:[m]2ndInangahua (+some bits), 1 1st. Landing, Leptospermum scoparium, 9 Nov 1959, J.G.R. o sum dorsal pores distributed in a reticulate McBurney, FRNZ no. R(a)3: 1/1[m]ad. pattern, delineating 7 reticulation areas between anterior margin and anal plates and with perhaps 19 areas around ema ks P. pollicinus can be differentiated from all margin. Dorsal pores of 4 types: (i) microductules of 2 other Plumichiton species by: sizes: very small pores: in reticulation lines; much larger (i) the short stigmatic spines, with a very blunt marginal pores: in a submarginal row along with large simple pores; spine either side; (ii) small simple pores, slightly larger than small (ii) the characteristic zigzag pattern formed by the concave microductules: associated with reticulation lines and macropores dorsally on the abdomen. occasionally elsewhere; (iii) large button-shaped simple P. pollicinus is similar to all otherPlumichiton species pores, about size of spiracular disc-pores: present in other than P. flavus in having concave macropores reticulation lines, in a submarginal row alternating with dorsally on the abdomen. It is closest to P. diadema, with large microductules (about 1 opposite every 2.5 marginal which it shares: spines), in a sparse row along anal cleft margin and around (i) 7 dorsal reticulation areas between the anal plates and anterior margin of anal plates and rather randomly within anterior margin; reticulation areas, particularly medially on dorsum; and (ii) medium length ventral anal lobe setae; (iv) large concave macropores, each with a heavily (iii) some large dorsal simple pores medially within the sclerotised base and a short, wide, non-sclerotised tube; reticulation areas and in a submarginal row. rather common, present in all reticulation lines, and with It resembles P. nikau in having a submarginal row of 0-4 in posterior medial macropore line. Anal plates: dorsal microductules, and a short claw on the legs; and is e g a ou μm com i e wi s a ou 15 μm; similar to P. flavus and P. punctatus in having a few with 2-5 minute pores on each upper surface; upper pregenital disc-pores medially on the abdomen. surface of each plate slightly folded or wrinkled, particularly along inner margins but never laterally on io ogy Apparently restricted to New Zealand's two anterior third; inner margin setae spinose with blunt tips, most common shrubland plants, Kunzea ericoides e g -31 μm; a ica se ae simi a u mo e s a y (previously Leptospermum) and Leptospennum scoparium. oi e e g a ou μm a ou e ma gi se ae se o The main overlapping generation(s) seems to be from dorsal surface, short and sharply spinose, length about 24 autumn through winter to spring, with both nymphs and μm A oge i a o wi ai s o se ae a o g a e io adults collected then. All males, nymphs, and some young ma gi a 1- ai s a e a y; o ges a ou μm adult females are found on the upper leaf surfaces but the Ma gi marginal setae spinose and stout, with mature females are on the stems. ou e i s 11-19 μm o g; e icu a io se ae s ig y displaced onto dorsum and with distinctly different basal is i u io Lowland shrubland throughout (Map 33). sockets; with about 27 spines on each side between stigmatic clefts; with a pair of larger setae on either side of ame e i a io The name is taken from: pollex, pollicis stigmatic spines, differentiated from other marginal setae (L.) meaning thumb, pollicinus meaning thumb-like (the only in being blunter. Stigmatic spines short and sharply mature adult female test resembles a small, horny thumb- oi e wi e y s a ow se a socke s; eac - μm nail). long, displaced onto dorsum. e e pregenital disc-pores with mainly 1 (range 7-10) loculi; mainly present on mediolateral folds on n f l nd 4 159 abdominal segments from anogenital area to posterior Although only two adult females are known, it is felt spiracles; also present sparsely medially on abdomen; that they are sufficiently different from other l h t n number of disc-pores on each segment (medially/ species to warrant description. mediolaterally on each side): anal cleft/VII, 0/15-20; VI, 3/2-4; V, 4/2-3; IV, 1/4-5; III, 1/2-6; and II, 1/2-5; Distribution. This species is only known from the one metathorax with none medially but with 2-5 laterad to collection in central South Island (Map 34). each metacoxa. Spiracular disc-pores: with 25-37 in each anterior band and 27-40 in each posterior band, pore Name derivation. The specific n p n t t is a Latin bands not extending past peritremes. Preantennal pores adjective meaning dotted or marked with spots, referring present. Ventral microducts in a submarginal row and to the many dorsal macropores. very sparse medially. Tubular ducts as for genus except possibly of 2 types, a broader type forming the submarginal band and with a slightly narrower duct sparse medially on abdomen; with a single duct near procoxae. e a se ae e a a a o e se ae a ou 7 μm o g; with about 7 pairs of flagellate anterior anal cleft setae; with a pair of long pregenital setae on each of segments VI a II o ges a ou 77 μm o g wi o e se ae o O O E A e e so & o gso , ew ge us moderate length; approximate number of setae medially on each abdominal segment: VII, 8; VI, 6; V, 5; IV, 7; III, Type species: Ct n h t n d r d Maskell (here 4; and II, 4; with perhaps 2 setae near each metacoxa, 3-4 designated) near each mesocoxa, and 3-4 near each procoxa; with 2 Diagnosis. Adult female. Test: (known for . d r d pairs of interantennal setae. Antennae noticeably broad, only), composed of moderately thick glassy wax plates, with 2 pseudosegments in the long third segment, total which separate from each other rather easily, marginal wax e g 5-3 5 μm; a ica se a 3 -3 μm o g fringe short on mature females, more noticeable on young C y eo a a s ie a ou 155 μm o g Wi o adults. s i acu a e i emes a e io 3 -3 μm os e io - Shape: oval to round, sometimes wider than long, with μm egs e g s (me a o acic co a 15 -1 μm; nearly vertical sides and a flattish top; mature females very oc a e + emu 1 - μm; i ia 15 -15 μm; large, up to 10 mm long, with a concave brood chamber a sus 11 -1 μm; c aw o g 3 μm beneath abdomen. Dorsum: derm membranous, apart from a distinct anal Material examined: HOLOTYPE [ ]: NEW ZEALAND: sclerotisation. Dorsal setae absent. Dorsal pores either MC: Waipara, Ol r r d nt , 12 Dec 1915, G. apparently randomly distributed ( . l b t or Brittin, no. 76, NZAC: 1/1[ ]ad. arranged in a reticulate pattern ( . d r d dorsal pores PARATYPE: as for holotype, (i) on holotype slide: crawler; of at least 3 types: (i) small, dark microductules: present (ii) mounted from dry material, upperside leaf, NZAC: #90- throughout and/or forming reticulation lines; (ii) simple 214,1/1[ ]ad. pores of various sizes: present throughout or in reticulation lines and also in a submarginal band; and (iii) Remarks. . p n t t can be differentiated from all large, heavily sclerotised macropores, either bollard-like, other l h t n species by the numerous large concave or cone-shaped with base set in a derm pocket: most dorsal macropores found throughout the dorsum in all abundant medially on abdomen near anal plates, tending to dorsal reticulation lines. . p n t t is closest to . become less frequent anteriorly and marginally; present in d d and . p ll n in having: reticulation lines when bollard-like or randomly when (i) medium length ventral anal lobe setae; cone-shaped. Preopercular pores, dorsal tubercles, and (ii) dorsal simple pores as large as spiracular disc-pores, dorsal tubular ducts absent. Anal plates broad, widest at present in the reticulation lines and in a submarginal about 3/5 from anterior margin, inner margins diverging row. slightly posteriorly; apex rather blunt; each plate with . d d differs (in addition to the distribution of either a few small pores ( . d r d or many, very large the dorsal macropores) in having pregenital disc-pores convex pores ( . l b t each plate with 4-5 long medially on all thoracic segments, while . p ll n setae posteriorly. Anogenital fold with about 5 pairs of differs in having frequent ventral tubular ducts medially setae along anterolateral margin, lying in unusually on the thorax. posterior position on . l b t probably without 160 d n & nd r n (2000 : C d (In t : pt r : C d supporting bars or plates. Anal tube short, barely reaching (viii) presence of marginal setae along margins of anal anterior margin of anal plates; anal ring with 6 setae. cleft; Margin: marginal setae stoutly spinose, with well (viii) presence of ventral microducts throughout most of developed, deep basal sockets; in a band 1-3 setae wide venter. around margin, on margins of anal cleft and extending The two species currently included in r p z are onto posterior margins of anal sclerotisation on dorsum. both endemic to New Zealand. Stigmatic clefts absent; stigmatic spines present or not differentiated from marginal setae. Multilocular disc- Name derivation. r p z is a combination from poros pores forming a complete marginal band 2-5 pores wide (Gr.) meaning hole and peza (Gr.) meaning edge or border, ventrally, apparently becoming dorsal along part or all of for the many disc-pores around the margin. margins of anal cleft anteriorly and extending onto anal sclerotisation on dorsum. Eyespot present or absent. Venter: pregenital disc-pores similar to those around Key o a u ema e r p z margin: present medially on posterior abdominal Dorsal pores not forming a distinct reticulate pattern, but segments (and more anteriorly on . l b t . Spir- abundant and apparently evenly distributed; large acular disc-pores similar to disc-pores around margin: in simple pores abundant throughout dorsum, particu- short, broad bands between spiracles and margin and also larly near margin; each anal plate with a group of very extending medially past peritreme to near or even past large simple pores; similar large simple pores also associated coxae. Preantennal pores possibly absent. frequent ventrally, particularly posteriorly on abdo- Ventral microducts of 1 size: more or less present men; marginal spines abundant, with more than 100 on throughout. With a large type of simple pore forming a either side between lateral stigmatic areas; stigmatic complete submarginal band. Ventral tubular ducts of 1 spines undifferentiated from spinose marginal setae type: abundant throughout but absent from near-marginal l b t band and medially on abdominal segments VI (on . l b t or VI and VII (on . d r d . Ventral setae: —Dorsal pores forming a distinctly reticulate pattern; with 4-10 anterior anal cleft setae; setae present medially dorsum and venter lacking large simple pores similar on head, thorax, and abdomen rather long and flagellate; in size to macropores; each anal plate with a few i e a e a se ae o g e g 1 3 μm a e y i e y minute pores only; with less than 35 marginal spinose age a e o e is a a (o e oke ; o e se ae as setae on either side between lateral stigmatic areas; o su ami y A e ae - o -segme e somew a stigmatic spines differentiated from marginal spines; e uce o . l b t setal distribution probably as legs reasonably well developed d r d for family. Mouthparts typical of family. Spiracular e i emes a icu a y a ge wi u o 19 μm se a e c ose o ma gi egs a e sma o si e o ma u e ema es a icu a y o . l b t structure and setal r p z l b t e e so & o gso ew distribution varying depending on degree of size s ecies reduction; lacking a tarsal campaniform sensilla; claw without a denticle. Vulva opening on segment VII. Fig. 131 Unmounted material: unknown. Remarks. This genus contains two species: r p z l b t Henderson & Hodgson, n. sp. and . d r d Mounted material: body oval to round, almost as wide as (Maskell) n. comb. These two species share a number of long, with a shallow anal cleft about 1/9th body length. important characters: Length 9.7-9.9 mm; breadth 8.3-8.6 mm. (i) large size, when mature; (ii) a marginal band of multilocular disc-pores, which also Dorsum: dorsal pores apparently randomly distrib- extends up anal cleft onto anal sclerotisation on uted; pores of 4 types: (i) unusually large microductules, dorsum; with along, rather characteristically shaped, inner ductule: (iii) presence of abundant tubular ducts throughout venter; present throughout but most abundant posteriorly; (ii) (iv) large spiracles, set rather close to margin; slightly larger, convex simple pores: infrequent through- (v) broad bands of spiracular disc-pores; out; (iii) large, slightly convex pores with a granular (vi) rather distinctively shaped anal plates; appearance: abundant throughout but also in groups (vii) short anal tube; forming a broad disjointed band near margin; and (iv) n f l nd 4 6 ig. . r p z l b t e e so & o gso , . s ., a u ema e. 62 d n & nd r n (2000 : C d (In t : pt r : C d . 2. r p z d r d (Maske , . com ., a u ema e. n f l nd 4 6 large macropores, with a very blunt apex and with base a ou 55 μm o g; wi 5- ai s o i e a e a se ae sunken in derm: abundant near anal plates but becoming o ges a ou 55 μm; wi 7-11 su ma gi a se ae less frequent anteriorly and marginally; absent in a wide e wee eac s igma ic a ea; wi sma se ae is i u e su ma gi a a A a a es e g 7-3 3 μm a e a om y me io a e a y A e ae somew a com i e wi 91-3 μm; wi 3 -53 a ge o es o e uce -segme e ; e g - 93 μm; wi o g o sa su ace o eac a e simi a o o e y e (iii o se ae o sca e; e g o a ica se a e a s 1 μm o sum; wi 5 se ae o os e io a o eac a e C y eo a a s ie 5 - μm o g Wi o eac os e io y o o sa su ace 1 o i e ma gi a s i acu a e i eme a ge 153-1 7 μm egs oo y su a ica y; eac 7 -1 μm o g A oge i a o yi g e e o e a some imes segme a io o scu e o a se ; e ea os e io e o a a a es; wi 5-7 ai s o se ae e g s (me a o acic co a 55- 1 μm oc a e + a o g ma gi o ges 15 + μm o g emu 99 μm i ia 5 μm a sus μm a c aw 9 μm; Margin: marginal setae each stoutly spinose, straight a sa igi u es a ou as o g as c aw 1 s ig y s ou e or slightly bent, in a narrow basal socket; with 110-132 in a o e o wi sma k o e a ices; c aw igi u es a band 1-3 setae wide on each side between stigmatic e y s o e a s o s ig y i a e a ica y c e s; e g 5- 5 μm; wi - 5 a o g eac ma gi o a a c e a e e i g o o os e io e o a a Material examined. HOLOTYPE [ ]: NEW ZEALAND: sc e o isa io o o sum S igma ic s i es o (WN]: No. 553, Dacrydium cupressinum, Orongorongo i e e ia e om ma gi a se ae Ma gi a a o Valley, 25 Sept 1969, D. Campbell, J.A. de Boer Collec- mu i ocu a isc- o es as o ge us Eyes o oo y tion, NZAC: 1/1[ ]ad (split dorsoventrally and mounted e i e o a se under 2 coverslips). Venter: pregenital disc-pores with mainly 7 outer loculi (range 5-8) and a round central loculus; present Other material: NEW ZEALAND: AK: Waitakere Ra, medially on all abdominal segments, throughout segments Sharp Bush, Blechnum fraseri underside leaves, 23 Oct VI, V, and IV but only along posterior margins of 1995, RCH, #95-105a—c: 3/2[ ] 3rd, 2[ ]2nd. As previous, segments III and II; also present in groups of 22-29 mesad except 18 Aug 1996, #96-154: 1/3 [ ]2nd,2nd 1 1st (+1 to each meso- and metacoxa. Marginal band of unknown sp.). WO: Raglan, Bridal Veil Falls, Schefflera multilocular disc-pores as for genus but also extending digitata, 19 Sept 1981, C.F. Butcher, #81-282j: 1/1 [ ]3rd. onto segment VII ventrally, where they form a distinct BP: Waiaroho, Hedycarya arborea, 29 Sept 1993, RCH, band anterior to anal tube opening. Spiracular disc-pores #93-315b: 1/1[ ] 3rd (pharate). As previous, except un- with mainly 8 loculi: with more than 150 disc-pores in derside of leaf midrib, 3 Nov 1993, #93-346: 1/1 [ ]3rd each band; each band extending medially to near each (pharate). GB: Taikawakawa, litter 92/71, 18 Oct 1992, associated coxa, also onto median venter near each J.S. Dugdale, #96-001: 1/1 [ ]3rd. *No collection data ex- procoxa. Ventral microducts: throughout, most abundant cept, New Zealand, NZAC, #83-326c: 2/2 [ ] ad. in a fairly broad submarginal band but also very common elsewhere; least frequent medially. With large, slightly Remarks. This is a very distinctive species. It is close to convex pores (similar to type (iii) on dorsum): present P. dacrydii but is immediately recognisable by the sparsely in a complete, broad, submarginal band, presence of large simple pores on the anal plates and in the becoming very abundant posteriorly on either side of anal more abundant marginal spinose setae. cleft; also present in small groups of 3-4 and 4-5 mediolaterally on abdominal segments II and III Biology. Immature specimens of P. cologabata, despite respectively and with a few medially on head. Ventral being found on leaves of Blechnum fraseri, Hedycarya tubular ducts: abundant throughout except in a marginal arborea, and Schefflera sp., and in litter rather than on band; least frequent medially and absent medially on Podocarpaceae, seem to belong to this species. Although abdominal segments VI and VII. Ventral setae: ventral none of the records of adult females indicate the site of anal lobe setae poorly demarcated from other submarginal collection on the host plant, it is assumed to be under the se ae eac a ou 1- 9 μm o g; wi - ai s o bark as with P. dacrydii, in which case the life cycle is a e io a a c e se ae; wi ai y o g se ae ese probably also unusual. No male stages are known. me ia y o a a omi a segme s a mesa o eac co a; um e me ia y o eac a omi a segme II Distribution. Possibly widespread but rarely collected; -1 ; I - ; 1 -1 ; I -1 ; III -1 ; a II 1 - adult known from the southern tip of North Island; 1 ; wi -1 se ae ea eac me aco a 11-15 ea eac imma u es known from Auckland, Waikato, and the East mesoco a a - ea eac oco a; o ges o o a Cape region (Map 35). 64 d n & nd r n (2000 : C d (In t : pt r : C d Name derivation. The specific name refers to the mediolaterally; absent submarginally; with 26-39 in abundant pores found apparently randomly over the posterior medial macropore line. Anal plates: length of dorsum: l from the Latin l (n.) meaning a sieve, a es 37- 5 μm com i e wi s 5 -3 3 μm; wi and b t the Latin for a dish or platter. 3-15 minute pores on dorsal surface of each plate; with 2 long setae along inner margin, a long apical seta and a long se a o ou e ma gi a se ae 1 + μm Wi 5- se ae o anterolateral margins of anogenital fold; longest 108-150 μm Margin: marginal setae stoutly spinose, with a heavily r p z d r d (Maske ew com i a io sc e o ise ee asa socke ; e g 1 - 3 μm; wi 17- 9 a e a y e wee s igma ic c e s; wi -1 o ei e Figs C79, C82, 132 margin of anal cleft and extending onto posterior margins Ct n h t n d r d Maskell, 1892: 18; —Maskell, of anal sclerotisation on dorsum. With 0 or 1 stigmatic 1 895a: 12 [checklist]; —Cockerell, 1896: 330 [checklist]; — spine per stigmatic area, each apically blunt and slightly Coleman, 1903: 81 [host comment]; —Fernald, 1903: 159 cu e e g 1 - 9 μm Ma gi a a o mu i ocu a [world catalogue]; —Hutton, 1904: 226 [checklist]; - disc-pores with mainly 10 loculi. Eyespots each 14-20 Myers, 1922: 199 [checklist]; —Miller, 1925: 33, 63 μm wi e [host]; -Hall, 1926: 18 [mention]; —Deitz & Tocker, 1980: Venter: pregenital disc-pores with mainly 10 outer 28 [checklist]; -Ben-Dov, 1993: 100 [world catalogue]. loculi and an oval central loculus; mainly found in groups on each side of posterior abdominal segments; number of Unmounted material: "Test of adult female white, or with pores on each side: VII none, VI, 15-20; V, 7-20; IV, 1- a slight yellowish tinge, moderately thick, formed of a 10; III and II, 0. Similar multilocular disc-pores also number of subcircular segments, which seem to be very present in a band 2-5 pores wide round entire margin. brittle and apt to break off; the edge of each segment is Spiracular disc-pores each with 8-10 (mainly 10) outer irregular, and the surface marked with irregular lines, and loculi; with more than 100 disc-pores in each band; each also radiating lines, so that it has somewhat the appearance band extending medially almost to associated coxa. of a fish-scale; the circular lines are rather deeply indented. Ventral microducts as for genus. With a slightly convex Fringe [of wax plates] in the specimens seen fragmentary. simple pore forming a complete, narrow, submarginal The segments of the test are sometimes convex, sometimes band; with 20-40 between stigmatic areas. Ventral flattish" (Maskell, 1892, p. 18). Live colour pinkish- tubular ducts as for genus: less frequent medially and brown but dark brown when preserved. Young adults absent from abdominal segment VII. Ventral setae: broadly oval with anal plates clearly noticeable posteriorly e a a a o e se ae eac a ou -1 μm o g; wi on dorsum; mature female more convex, with a raised 4-10 pairs of anterior anal cleft setae; with long setae flattish top and nearly vertical sides, and with anal plates present medially on all abdominal segments and mesad to and anal cleft actually on posterior vertical margin. each coxa, rather abundant, many with an extremely long, fine, flagellate apex; number medially on each abdominal Mounted material: body oval to round, sometimes wider segment: VII, 7-14; VI, 18-26; V, 20-30; IV, 18-28:III, than long, with a shallow anal cleft, about 1/10th body 15-26; and II, 10-28; with 18-38 on metathorax, 30-50 length. Length 3.5-8.5 mm; breadth 2.7-9.5 mm. near each mesocoxa, and 4-24 near each procoxa, longest 135+ μm; wi 3- ai s o i e -a e a se ae; wi -13 Dorsum: dorsal pores distributed in a reticulate submarginal setae laterally between each stigmatic area; pattern, delineating reticulation areas each with a few small setae also present rather randomly mediolaterally. small microductules only; these reticulation areas in 7 Antennae well developed, 6- to 8-segmented; total length longitudinal rows, probably with 9-10 areas between anal 3 -511 μm; wi a o g se a o sca e; e g o a ica plates and anterior margin and with 30 areas around se a 135+ μm e g o c y eo a a s ie 3-5 μm margin; dorsal pores of 3 types: (i) microductules, Wi o eac s i acu a e i eme 1 9-1 9 μm egs probably with long inner ductules, although these could well developed but rather narrow and frequently not be seen: distributed throughout but most abundant in miss a e ; e g s (me a o acic co a 91-1 μm reticulation lines; (ii) slightly larger, convex simple pores: trochanter + emu - 1 μm i ia 13 -171 μm a sus common in lines of reticulations and also forming a band 91-135 μm c aw 13- μm; a sa igi u es o o ge submarginally; and (iii) macropores bollard-like; most than claw, 1 slightly stouter than other; claw digitules both abundant towards posterior end, medially and broad; claw without a denticle. n f l nd 4 6 Material studied: LECTOTYPE (here designated): NEW pr n , 16 Feb 1982, C.F. Butcher, #83-335e: 1/1[ ] ZEALAND: labelled "Ct n h t n d r d Mask., New ad. Zealand, Mask. Coll, No. 202", USNM: 1/1 [ ]ad (dorsum and venter split). Remarks. The original material was collected in the PARALECTOTYPES: "Ct n h t n d r d Mask., New Reefton district ( (Maskell, 1892). This species and P. Zealand, Mask. Coll. No. 202", NZAC: 1/1 [ ]ad (un- l b t (described as new above) appear to be closely mounted, dry). "Ct n h t n d r d , early 2nd stage fe- related, but . d r d is immediately separable in: male, 1891, W.M.M.", NZAC: 1/1 1st (i.e., a crawler rather (i) possessing a reticulate pattern of pores on the dorsum; than a [ ]2nd).1891,Ct n h t n W.M.M.: d r d l rv , (ii) lacking the large simple pores on the anal plates which 1/9 1st. Mask. coll., #202, USNM: 1/21 1st. are so distinctive on . l b t . Other material: NEW ZEALAND: ΚΕ Kermedec Is., io ogy Maskell (1892, p. 19) commented that "Mr Raoul Is. crater, litter 72/188, 11 Oct 1972, C.R. Veitch, Raithby first found the specimens under thick moss on the #17, 18: 3/2[ ] 3rd, 1[ ] 2nd. AK Riverhead SF, under roots just above the ground; later finds were under the bark bark of d rp t t r , associated with ants, 7 Nov on the trunk; and he thinks that the species may sometimes 1981, C. Mercer, #81-317d: 1/1[ ]ad. Riverhead Forest, be subterranean." All specimens in the NZAC from under Barlow Road reserve, d rp t t r under bark, 6 bark of various Podocarpaceae are either large mature Feb 1998, C.J. Hodgson & RCH, #96-007a-e: 5/4 [ ] ad, females, much smaller young adult females or still smaller 1[ ] 3rd. O Whirinaki, d rp t t r bark, 18 May 3rd-instar females, all of which have large mouthparts, 19 5 S ug a e ΝΖ ( / 7 / [ ] ad (poor con- with stout, long stylets, presumably adapted to feeding dition). Te Whaite area, d rp t t r , Jan 1970, R.J. through the outer tissues of the tree; 2nd-instar females McKenzie, FRNZ R(b)28-31: 4/4 [ ] ad (dorsums and have never been found under bark with adults. The venters split). Mimiha Strm, Ruatahuna, d rp crawlers that emerge from beneath the mature females do t t r , 17 Mar 1970, R.J. McKenzie, FRNZ R(b)32-36: not have particularly strong mouthparts and it is supposed 5/5[ ] ad (dorsums and venters split). Mangawiri Basin that they may leave the maternal site and migrate 125', S.F. 58, Whirinaki, ex d rp t t r , April 1972, elsewhere, perhaps feeding on the fine roots? Miller R.J. McKenzie, #844: 1/1 [ ]ad (dorsum and venter split). (1925) indicated that he had found . d r d infesting W Orongorongo Valley, r d pr n , 25 not only the bark but also the roots of rimu ( r d Sept 1969, D. Campbell, #553: 1/1[ ] ad (venter and dor- pr n . If the early instars do feed elsewhere, it is sum split). NN: Riwaka Valley, Motueka, Nelson, likely that they grow to the 3rd instar before migrating r rp d r d d [as d rp d r d d ], back beneath the bark. This suggestion is supported to 17 Nov 1918, G. Brittin: 1/1[ ] 3rd. Buller Gorge, some extent by the fact that three nymphs of . d r d , r d pr n , 17 Mar 1971, J.A. de Boer, #713: including the only known 2nd-instar female, were 4/3[ ] ad (all split, 2[ ] on single slides, 1[ ] dorsum and collected from litter (on the Kermedec Islands). It is also venter on separate slides). Mawhera S.F., nr Reefton, possible that the young and mature adults found together r d pr n , 19 Apr 1972, J.S. Dugdale, #846: under the bark may be from different generations as it is 1/1[ ]adsp., d rp (split). FD: Secretary I, Bauza I, unusual in New Zealand Coccidae to find such a huge size 26 Nov 1981, C.F. Butcher #82-148c: 4/3 [ ] ad, 1[ ] 3rd. difference in a given population - mature adults 3-4 x Breaksea Sound, Gilbert 6, r n p t f rr n [as length of young adult females - in one cohort; d rp f rr n ] bark, 12 Mar 1983, C.F. Butcher, alternatively the life cycle may be very long. No male #83-321 a, -326f: 2/2[ ] ad. As previous, except [as miro], stages are known and so this species is probably Gilbert Is. 6, #90-242a-c: 12/6 [ ] ad, 1 [ ]3rd, 31 1st. parthenogenetic. Bauza I, d rp t t r under bark & moss, 28 Jan . d r d is invariably attended by the native ant 1996, RCH, #96-080: 1/1[ ]ad (anal plates dissected). r l dv n (Fr. Smith) when under the bark of trees. Breaksea Sound, Gilbert 6, d rp t t r under thin This is the only New Zealand coccid known to have a bark, 2 Feb 1996, RCH, #96-046a-b: 2/4[ ]3rd. Dusky relationship with ants. Sound, Anchor I, by lake, r d pr n under bark, 10 Feb 1996, RCH, #96-047: 1/1 [ ]ad. Chalky Inlet, is i u io In lowland podocarp forest throughout Little I, d rp t t r under thin bark, 11 Feb 1996, (Map 36). RCH, #96-045a-d: 4/2 [ ] ad, 2[ ]3rd, 7 1st. Preservation Inlet, Cuttle Cove, d rp t t r under thin bark, 13 Feb 1996, RCH, #96-044a-o: 15/10[ ] ad, 7 [ ]3rd, 7 1st. SL: Bluff Hill, Glory Track, bark of r d 66 d n & nd r n (2000 : C d (In t : pt r : C d OU AMOCOCCUS e e so & o gso sparsely medially and mediolaterally on more posterior abdominal segments. Ventral setae: long pregenital setae n Henderson & Hodgson: Hodgson & on segment VII present or absent, otherwise distribution Henderson, 1998: 606 of ventral setae typical of family. Antennae well y e s ecies: n t b l Henderson & developed, 6- to 8-segmented, 3rd segment generally with Hodgson. l-2 pseudosegments; 3rd segment from apex with a flagellate seta and penultimate segment with 2 flagellate iag osis. A u ema e. es : of medium thick glassy setae in addition to the usual single fleshy seta. wax, with or without sutures. Mouthparts usually displaced to one side and labium o y: pyriform, narrower at head end, usually clearly twisted through about 90°. Spiracles as for family. Legs asymmetrical, with distinct stigmatic and anal clefts. quite well developed; each with a separate tibia and tarsus Asymmetry shown most obviously by mouthparts which but no articulatory sclerosis; tibia with 2 setae on distal are generally nearer 1 procoxa and with labium usually end; claws with or without a small denticle; tarsal digitules twisted through 90°. both fine and not extending past end of claw in length; o sum: derm membranous, even on old specimens; claw digitules dissimilar, one broader than other; tarsal with or without sclerotisations in the stigmatic clefts but campaniform pore present on all 3 pairs of legs. Vulva with no sclerotisations anterior to anal plates. Dorsal setae opening on segment VII. absent. Dorsal pores not in a reticulate pattern; dorsal pores of 3 or 4 types: (i) microductules, with a long inner ema ks. This genus is endemic to New Zealand and filament: fairly frequent throughout but not forming a contains two species: n n t Henderson distinct pattern; (ii) simple pores, varying in size, those in & Hodgson and . t b l Henderson & Hodgson. . n t medium size, while those in . t b l very Species in the genus n differ from all other small; (iii) macropores very large and slightly convex known soft scale genera in possessing tarsal campaniform (present on . n t and (iv) chimney-shaped pores (see Hodgson & Henderson, 1998). Other macropores (present on . t b l . Preopercular pores significant characters are : and dorsal tubercles absent. Dorsal tubular ducts present (i) the asymmetry of the labium, which is approximately at or absent, when present in 2 diverging lines anterior to anal right angles to the long axis of the body; plates. Anal plates together approximately quadrate; each (ii) the presence of 2 setae on the distal end of each tibia; with 2 setae along inner margin near apex, an apical seta (iii) the restriction of the ventral tubular ducts to medially and 1 small seta either on posterior margin or almost in and mediolaterally on the abdomen; discal position. Anogenital fold apparently without (iv) more than l stigmatic spine per stigmatic cleft, supporting bars; with 2 pairs of setae along anterior margin forming a distinct group at the base of the stigmatic and with 1 or 2 setae along lateral margins. Anal tube clefts; moderately long; anal ring with 3 pairs of setae. (v) presence of finely spinose marginal setae; Ma gi : marginal setae all finely spinose, with a blunt (vi) presence of 2 flagellate setae on the penultimate apex and a small basal socket: in a single row along antennal segment. margin, with 5-18 on each side between stigmatic clefts, On the basis of these characters, n is usually with fewer on side nearest mouthparts; not quite unlike any other New Zealand genus. All known extending up margins of anal cleft; anal lobe setae not stages of the two species currently included in differentiated from other marginal setae. Stigmatic clefts n were described by Hodgson & Henderson distinct and quite deep (shallower on older swollen (1998). specimens), with or without stigmatic sclerotisations. Each cleft with 2-8 stigmatic spines of distinctive shape. Eyespot probably absent. Key o a u ema e n e e : derm membranous. Pregenital disc-pores Stigmatic spines long and tubular, expanded apically, generally with more than 5 loculi; restricted to posterior 3 with 6-8 spines per cleft; tubular ducts absent from the pregenital segments. Spiracular disc-pores with mainly 5 dorsum; dorsal macropores heavily sclerotised, loculi: in broad bands between spiracles and margin. chimney-like, and rather scarce t b l Preantennal pores present or absent. Ventral microducts —Stigmatic spines wedge-shaped, with 2-3 spines per frequent throughout. Ventral tubular ducts of 1 type, with cleft; tubular ducts present dorsally in 2 diverging a long outer ductule, an almost equally long inner ductule lines anterior to anal plates; dorsal macropores large, and a well-developed terminal gland: present rather convex and frequent throughout n t n f l nd 4 6 ig. . n n t e e so & o gso , a u ema e. 68 d n & nd r n (2000 : C d (In t : pt r : C d ig. 4. n t b l e e so & o gso , a u ema e. n f l nd 4 1 9 n n t e e so a o gso extending medially past each peritreme. Preantennal pores present. Ventral tubular ducts similar to those on Figs M31, C80, C81, 133 dorsum but slightly narrower: very sparse medially and mediolaterally on abdominal segments IV—VI. Ventral Pounamococcus cuneatus Henderson & Hodgson; setae: with a pair of moderately long setae medially on all Hodgson & Henderson, 1998: 623 abdominal segments, particularly V and VI, but these Unmounted material: live specimens mottled green, absent from VII, and with a pair of shorter setae on often with a darker green submedian oval ring, extending segments VII—III; also generally with 1 long and 1 short from head to anal plates. Wax test formed of 7 median- seta near each meso- and metacoxa, and 1-2 short setae thick plates, 4 smallish plates on cephalothorax (1 near each procoxa; with 2-3 pairs of interantennal setae, anteriorly, 2 laterally, and 1 mediodorsally) and 3 long o ges 5 - 5 μm; wi a ou - su ma gi a se ae o plates posteriorly; internal structure of test composed of eac si e e wee s igma ic c e s A e ae we layered wax (Fig. M31); median plates separated from e e o e - o 7-segme e 3 segme ge e a y wi lateral plates by submedian suture which corresponds to - seu osegme s; o a e g 3 13 μm; e g o green stripe (when that is present) on underlying insect; eac a ica se a 1- 5 μm e g o c y eo a a s ie surface of test slightly dimpled. All sutures between plates 1 7-1 7 μm Wi o s i acu a e i emes a e io 9- are true sutures. Post-ovipositional females tend to shrink 5 μm; os e io 3 -5 μm egs e g s (me a o acic towards anterior end of test. co a 7 -95 μm; oc a e + emu 175- 3 μm; i ia 9 -1 μm; a sus 1 1-139 μm; c aw 3- 5 μm; c aws Mounted material: length 2.5-4.4 mm, width 1.5-3.8 wi a sma e ic e mm. Material examined: HOLOTYPE [ ]: NEW ZEALAND: Dorsum: dorsal pores of 3 types: (i) microductules: WD: Otira, 30 Dec 1915, on fern, ex dry material G. Brittin probably throughout but most frequent submarginally; (ii) collection, NZAC #90-213g: 1/1 [ ] ad (designated by medium-size simple pores present submarginally; and (iii) Hodgson & Henderson, 1998). large, slightly convex macropores: frequent throughout PARATYPES: as for holotype, NZAC #90-213a—f :6/6 [ ] except submarginally. Dorsal tubular ducts present in 2 ad; Brittin original slides labelled "on fern, Otira, Dec 1915, diverging lines just anterior to anal plates, each duct with Ctenochiton, no 96": 2/2[ ] ad. a fairly broad outer ductule, deep cup-shaped invagination and narrow inner ductule with a distinct glandular end; Other material: NEW ZEALAND: AK: Waitakere Ra, with 13-20 ducts in each line. Anal plates with apices not Fairy Falls, Blechnum fraseri, 21 Sept 1982, C.F. Butcher, i e gi g; e g o a es 133-1 7 μm wi o si g e #84-014h: 3/1[ ] ad, 2[ ] 2nd. Fairy Falls, Dec. 1982, ex a e 59-7 μm; a es wi ou mi u e o es; e g s fern, C.F. Butcher, #83-321d: 1/1[ ] ad and #95-148: 1/9 i e ma gi 1 1 -1 μm; i e ma gi 1 - μm; 2nd. Waitakere Ra, Sharp Bush, Blechnum fraseri under- a ica 1 - μm; se ae o eac os e io ma gi a mos i side leaves, 23 Oct 1995, RCH, #95-103a—c, f: 4/3 [ ] ad, isca osi io 1 - μm A oge i a o wi ai s o 1 1st. As previous except 18 Aug 1996, #97-155: 1/1 [ ] se ae su equa i e g (37-5 μm a wi 1 se a o ad. As previous except, underside of fronds, 5 Feb 1997, eac a e a ma gi e g 1 - μm #97-031 a—g: 7/72nd. As previous except, 14 Mar 1997, Margin: marginal setae finely spinose: length 12-36 [m] RCH, #97-058a—d: 4/4 pupae, 1 ad + cast pupal skin, 19 μm wi 1 -1 o eac si e e wee s igma ic c e s [m] 2nd, 1 2nd2nd. As previous+ pharate except 6 Feb Stigmatic clefts lacking stigmatic sclerotisations; each [m] 1998, RCH & C.J. Hodgson, #98-a-i: 9/1 [ ]ad (pharate), cleft with 2-3 stigmatic spines (occasionally with a 1 [ ]2nd,2nd (3 split dorsoventrally). 7 smaller spine on or near outer margin, which is usually [m] peg-like but can resemble typical stigmatic spines), each spine distinctively wedge shaped, although round to oval Remarks. Adult female P. cuneatus differ from those of at base, and with poorly-developed basal socket; length P. tubulus in the following: 11-1 μm (i) shape of the stigmatic spines, which are wedge-shaped Venter: pregenital disc-pores with 5-8 (mainly 6-7) and short (trumpet-shaped in P. tubulus); loculi; total number per segment: anal cleft/VII, 36-49; (ii) presence of dorsal tubular ducts in 2 diverging lines VI, 7-16; V, 1-13; IV, 0-2; III, 0-1; and II, 0; none laterad just anterior to the anal plates; to metacoxae. Spiracular disc-pores: with 28-58 in each (iii) a denticle on the claw; anterior band and 39-85 in each posterior band; with 0-4 (iv) pregenital disc-pores with mainly 5-8 loculi; 17 d n & nd r n (2000 : C d (In t : pt r : C d (v) moderate-sized, granulate pores throughout the o a es 15 -19 μm wi o si g e a e 7 -1 3 μm; dorsum; each plate with 0 or 1 minute pore; lengths of setae: inner (vi) absence of a pair of long setae on abdominal segment ma gi 1 5-3 μm; i e ma gi 1- 7 μm; a ica VII; 1 -1 μm a ou e ma gi se ae (o ou e ma gi ea (vii) absence of highly-sclerotised, tubular macropores on a e 7-9 μm A oge i a o wi ai s o se ae a o g the dorsum. a e io ma gi (1 ai a e o g ( 3-7 μm a o e For a further discussion, see Hodgson & Henderson very short) and with 2 pairs of setae on lateral margins, (1998). o ges 7-33 μm Ma gi marginal setae narrowly spinose: length 9- io ogy Presumed to be univoltine, with adult males and 17 μm; wi 5-15 o eac si e e wee s igma ic c e s females emerging in late summer, the adult females Stigmatic clefts with a slight stigmatic sclerotisation; each overwintering and the young stages appearing in late cleft with 6-8 stigmatic spines, each spine broadening at spring to early summer. Test not persisting on host plant apex (rather like a trumpet) and possibly with an apical after death. pore; with well-developed basal sockets; length 12-35 μm a oge s a a asi oi s Hymenopterous parasitoid e e pregenital disc-pores with mainly 10 loculi; recorded: Encyrtidae: Ad l n rt d v r b l Noyes. with 14-28 disc-pores on segment VII, 4-7 on VI , and 0- 2 on segments II— ; also with 0-3 pores laterad to each is i u io Originally collected from the middle of the metacoxa. Spiracular disc-pores: with 45-75 in each South Island just west of the Main Alpine Divide at Otira anterior band and 50-130 in each posterior band; with more than eighty years ago, although the more recent only 2-5 extending medially past peritreme. Preantennal collections have been from the forested hills west of pores absent. Ventral microducts with a characteristically Auckland City, North Island. It appears to be restricted to shaped inner ductule. Ventral tubular ducts: present ferns and possibly Blechnaceae (Map 37). medially and mediolaterally on all abdominal segments but most abundant posteriorly; also frequently with a single tubular duct just posterior to an antenna. Ventral n t b l e e so & o gso setae: with a pair of moderately long setae on all abdominal segments but those on pregenital segment Figs M 15, 134 much the longest; total number per segment: III 2; II 4-8; VI-lI, 4; with 2-4 setae near each metacoxa, 3-5 near n t b l Henderson & Hodgson; — each mesocoxa, and 3 near each procoxa; with 2-4 pairs of Hodgson & Henderson 1998: 612 interantennal setae, often asymmetrically distributed, U mou e ma e ia live females medium-dark green, o ges - 5 μm; wi -7 su ma gi a se ae o eac with 4 spiracular pore bands showing as white stripes. side between stigmatic clefts. Antennae well developed, Test composed of a medium-thick wax plate without 7- or 8-segmented, 3rd segment sometimes with a pseudo- sutures but with a faintly dimpled surface; older, post- segmentation when 7-segmented; total length 315-416 ovipositional females tending to shrink towards anterior μm; e g o eac a ica se a 3 - μm Mou a s end of test by concertina-like folds of abdomen. generally displaced to one side; labium almost invariably twisted through about 90°; length of clypeolabral shield Mou e ma e ia length 1.8-4.7 mm, width l.0-2.7 1 -195 μm S i ac es wi o e i eme a e io 3 - mm. 5 μm; os e io 5 - μm egs e g (me a o acic co a 3 -5 μm; oc a e + emu 1 1-1 μm; i ia o sum dorsal pores of 3 types: (i) microductules: 5- 1 μm; a sus 1-7 μm; c aw (wi ou a e ic e ; frequent throughout; (ii) simple (closed) pores, very small, 1 - μm which may be of 2 types: 1 type with a granulate surface, the other possibly without; both fairly frequent Ma e ia e ami e O O Y E NEW ZEALAND: throughout; and (iii) heavily sclerotised macropores, Doubtful Sound, Bauza 1 28 Jan 1996, RC Henderson, composed of a short chimney-like tube protruding above d p n x pl x [= pl x] upperside derm, a broad sclerotised ring on derm surface and an inner leaves, NZAC #96-042k: 1/1 [ ]ad (designated by Hodgson sclerotisation about equal in length to that above derm & Henderson, 1998). surface (Fig. M15): rather sparse. Dorsal tubular ducts PARATYPES: as for holotype, NZAC #96-042a-j, l-o: absent. Anal plates with apices slightly divergent; length 14/10[ ] ad, 2[m] 2nd, 1 pharate pupa, 17 1st (paratypes of n f l nd 4 171 nymphs omitted in Hodgson & Henderson (1998)). Ge us UM O IC I O e e so & Other material: NEW ZEALAND: FD: Fiordland, [no locality], Pseudopanax sp., Mar 1983, C.F. Butcher,#90- o gso ew ge us 207 (#83-339): 1/1 [ ]ad. Charles Sound, Elaenor I, Type species: Ctenochiton hymenantherae Maskell Hedycarya arborea. 26 Jan 1996, RCH, #96-041a-b; 2/ (here designated). 2[ ] ad, and #96-082a-c: 3/3 I,[m]2nd. Secretary Diagnosis: adult female. Test: glassy wax, convex, with Pseudowintera sp., May 1982, C.F. Butcher, #84-012c: 3/ 7 rows of plates, median row with six plates, each plate 3[ ] ad. Secretary I, Grono Bay Track, Pseudowintera convex like a knob (except on U. pellaspis where test is of colorata 24 Nov 1981, C.F. Butcher, #97-002: 1/1 [ ]ad. leathery wax with fibrous strands); insect beneath often As previous except, Pseudopanax sp., March 1983, #83- showing bicoloured through test. 332d: 3/4[ ] ad. Bauza I, Pseudopanax lessoni, Jan. 1984, Shape: elongate oval with rounded ends. Small, C.F. Butcher, 1/1[ ]ad. Dagg Sound, Anchorage Arm, length <3.5 mm; breadth <2.1 mm. Pseudowintera colorata, 1 Feb 1996, RCH, #96-083: 1/ Dorsum: derm membranous. Dorsal setae absent. 1 [ ]ad.Pseudopanax Breaksea Sound, Breaksea I, Dorsal pores distributed in a reticulate pattern, delineating arboreus, 26 Jan 1996, RCH, #96-074a-e: 5/3[ ] ad, 1 reticulation areas in 7 longitudinal rows across abdomen, pharate prepupa, 4 1sts. Dusky Sound, Cooper I, Sports- with 9 areas between anal plates and anterior margin and man Cove, Pseudopanax arboreus both sides of leaves, 7 29 areas around margin; dorsal pores of 3 or 4 types (i) Feb 1996, RCH, #96-040a—n: 14/12[ ] ad, 1 pharate [ ] small, dark microductules, with or without discernible ad, l[m][mm]2nd,pharate,ad. Dusky 1 pupa, Sound, 1 3 inner ductules; (ii) small simple pores, about same size as Girlies I, Raukaua simplex [as Pseudopanax simplex] microductule pore; both pore types most frequent in leaves, 9 Feb 1996, RCH, #96-084: 1/1 [ ] ad (pharate [ ] reticulation lines, occasionally with 1 or 2 between 2nd-[ ] ad). reticulation junctions on margin and along margins of anal cleft; (iii) large, heavily sclerotised macropores, shape Remarks. Adult female . tubulus differ from those of . probably diagnostic of species, either cone-shaped, cuneatus in the following: bollard-like or mushroom-shaped and apparently extend- (i) stigmatic spines trumpet-shaped; ing above dorsal derm surface between wax plates of test, (ii) dorsal tubular ducts absent; each with a round, heavily sclerotised, inner base; most (iii) pregenital disc-pores with mainly 10-loculi; abundant in median and submedian reticulation lines. (iv) presence of a pair of long setae on the pregenital Preopercular pores, dorsal tubercles, and dorsal tubular segment VII; ducts absent. Anal plates widest in anterior quarter, (v) presence of highly sclerotised macropores on the tapering to apex; with minute pores on upper surface of dorsum; each plate; surface relatively smooth; each plate with 2 (vi) absence of a denticle on the claw. inner margin setae, each short, sharp, and fine, 1 apical For further discussion see Hodgson & Henderson seta, spinose or setose and generally slightly longer than (1998). inner margin setae, and a posterior margin seta on upper anal plate surface, usually rather setose; without an anal Biology. Apparently oviparous and univoltine, with sclerotisation. Anogenital fold with 2 large sclerotised young produced in summer. Old tests not persisting on plates arising internally and extending anteriorly from host plant after death. anterior margin; with 2-4 pairs of setae along anterior margin and with 1 pair laterally. Anal tube moderately Distribution. Known only from Fiordland in the south- long; anal ring with 6 setae. west of the South Island. This mountainous region has a Margin: marginal setae small and finely spinose; with mean rainfall of about 5-6 metres per year (Map 38). 3-12 setae on each side between stigmatic clefts; in a single line not extending up margins of anal cleft; reticulation setae sometimes differentiated; marginal setae on anal lobes not differentiated. Stigmatic clefts shallow, without a stigmatic sclerotisation, each with 1 stigmatic spine of moderate length, about 6-10 x length of marginal spines. Eyespot present (but hard to discern on most specimens). Venter: pregenital disc-pores with 3-8 (mainly 5) outer loculi, on mediolateral folds of abdominal segments 2 d n & nd r n (2000 : C d (In t : pt r : C d in a line from anal cleft extending towards each posterior U b n h t n species by its lack of ventral tubular ducts. spiracle; occasionally 1-2 present medially on segments r p z d r d also has bollard-shaped macropores V–VI on U. d l a few sometimes present laterad to and can be distinguished by its marginal rows of disc- metacoxae. Spiracular disc-pores with mainly 5 loculi, in pores. narrow bands 1-5 pores wide between spiracles and Species in the genus U b n h t n resemble those in margin and with a few extending medially. Ventral l h t n in having pregenital disc-pores more or less microducts of 1 type, present in a submarginal band restricted to the mediolateral lobes of the abdomen in a line (throughout submargin on U. d l and in segmental between the anal cleft and the posterior spiracles. bands medially, except on posterior 1-2 abdominal l h t n species differ from those in U b n h t n in segments. Preantennal pores: generally with 1-2 pairs having: present. Ventral tubular ducts of 1 type, present in a broad (i) concave dorsal macropores; submarginal band and usually also present medially on (ii) spinose setae on the anal plates, with those nearest e head, thorax, and abdomen (absent medially on U. apex tending to be very blunt and parallel-sided; h n nth r . Ventral setae: with 1-4 pairs of anterior (iii)the dorsal surface of the anal plates folded or wrinkled, anal cleft setae; with a single pair of long pregenital setae particularly along the inner margins; on segment II only; hypopygial setae absent; with 4-9 (iv) many long setae ventrally, especially around the (total) (mainly 8) interantennal setae; other setae anogenital fold; distributed as for family. Antennae 6- to 8-segmented, (v) rather more abundant marginal spinose setae. with 0-2 pseudosegments on segment III when 6- segmented; setal distribution as for family, but longest seta Ge e ic ame e i a io Name refers to the knobbly test on apical segment shortest on U. j b t (3 - μm a and the convex dorsal macropores, thus: b , b n o ges o U. h n nth r ( -11 μm Mou a s (L., masculine) meaning a boss, knob, or shield and chiton occasio a y is ace o o e si e S i ac es y ica o (Gr.) meaning tunic or garment worn next to the skin. family. Legs well developed, generally with a separate tibia and tarsus but no articulatory sclerosis; tarsal campaniform pore absent; claws small and short without a Key o a u ema e U b n h t n denticle; tarsal digitules knobbed, unequal in length and Dorsal macropores approximately mushroom-shaped thickness; claw digitules expanded and equal, much when seen from the side, apex expanding to several longer than claw. Vulva present in segment II times the width of the basal `stalk' p ll p ema ks This genus contains five species: U. d l —Top of dorsal macropores approximately similar in Henderson & Hodgson, s U. b ll t Henderson & width to or narrower than their inner base 2 Hodgson s U. h n nth r (Maskell) com U. Dorsal macropores `bollard-like', with a broadened, Henderson & Hodgson and U. p ll p j b t s very blunt, apex; marginal reticulation setae larger Henderson & Hodgson s than other marginal setae b ll t Species in the genus U b n h t n are characterised by the following combination of characters: —Dorsal macropores `cone-shaped', with more or less (i) few spinose marginal setae, and moderately long pointed apices; marginal reticulation setae similar in stigmatic spines; size or only slightly larger than other marginal setae (ii) presence of very large sclerotised dorsal macropores of 3 either cone -, bollard-, or mushroom-shape; 3 Ventral tubular ducts absent medially from head, thorax (iii) very small simple pores and apparently ductless and abdomen (although 1-3 ducts occasionally microductules in dorsal reticulation lines; present between coxae) h n nth r (iv) a broad submarginal band of ventral tubular ducts; —Ventral tubular ducts present medially on head, thorax (v) pregenital disc-pores restricted to either side of the anal and abdomen 4 cleft and on the abdominal mediolateral folds, forming a line between the anal cleft and posterior spiracles. Marginal setae finely spinose and with 3-5 setae In addition, the genus is characterised by a knobbly between stigmatic clefts; dorsal macropores few, with glassy test (except for U. p ll p , which has a unique test none in transverse median reticulation line anterior to made up of strands of fibre incorporated with wax). anal plates; ventral tubular ducts sometimes present Aph n h t n n n p , with large bollard- medially on abdominal segments II–III or absent shaped macropores, can be distinguished from all medially on all abdominal segments d l n f l nd 4 173 —Marginal setae clearly spinose and with 5-10 setae present medially on each abdominal segment: VII, 4-7; between stigmatic clefts; dorsal macropores more VI, 4-7; V, 8–11; IV, 8-13; III, 6-10; and II, 3-12; with numerous, with several on transverse median (laterally near each coxa/medially between coxae), 1-4/3- reticulation line anterior to anal plates; ventral tubular on metathorax; 3-6/1-2 on mesothorax, and 3-4/0 on ducts scattered throughout abdominal segments II–V prothorax; with 6-7 (total) interantennal setae; with 2-7 at least j b t small submarginal setae on each side between stigmatic clefts. Preantennal pores present. Antennae 6- or 7- segmented, segment 3 with 1-2 pseudosegments when 6- segme e ; o a e g 5- 77 μm e g a ica se a U b n h t n d l e e so & o gso ew 3 - μm C y eo a a s ie 115-13 μm o g Wi s ecies o s i acu a e i emes a e io 7-3 μm os e io 3 - 3 μm egs e g s (me a o acic co a 5-1 μm; Figs C92, 135 oc a e + emu 11 -119 μm; i ia -9 μm; a sus U mou e ma e ia : test as for genus. 5- μm; c aw 1 -15 μm Mou e ma e ia : body elongate oval with rounded Ma e ia e ami e : HOLOTYPE [ ]: NEW ZEALAND: ends. Length 1.27-2.24 mm; breadth 0.76-1.40 mm. AK: Riverhead Forest, Barlow Road reserve, 14 August 1997, L.H. Clunie, Podocarpus totara leaves, NZAC #97- o sum: dorsal pores distributed in a reticulate 125a: 1/1[ ]ad. pattern as for genus; dorsal pores of 3 types as for genus PARATYPES: same collection data as holotype: NZAC except that dorsal macropores are heavily sclerotised and #97-125b-1: 11/11[ ]ad. cone-shaped, sometimes becoming detached from their sunken base during slide-mounting; most abundant in Other material: NEW ZEALAND: AK: Riverhead Forest, median and submedian reticulation lines, but with none in Barlow Road reserve, Podocarpus totara leaf by petiole, me ia a s e se i es A a a es e g 1 3-13 μm 31 July 1997, RCH, #97-113: 1/1 [ ] ad. As previous, ex- com i e wi s 9 -11 μm; wi 3- mi u e o es o cept 14 Aug 1997, #97-126a–j: 10/1 [ ] ad, 5[ ] 3rd, 2[ ] 2nd, upper surface of each plate; length of setae: inner margin 1, 4[m][ ]2nd,ad. As 11 previous, 1st; #97-142, except 1/1 17-1 μm i e ma gi 7-1 μm a ica se ae 7-1 stem and leaf, 22 July 1998, #98-084: 1[m]/12nd. Waitakere μm ou e ma gi se ae 1 -1 μm A oge i a o wi Ranges, Scenic Drive, under bark of Podocarpus totara, 3-4 pairs of setae along anterior margin and a single pair 15 Jan 1983, .M. Cox, 134: 1/1[ ] ad [note, this collection a e a y o ges se a 3-3 μm `under bark' seems at variance with most other collections Ma gi : marginal setae small and finely spinose, (from leaves) and this detail maybe incorrect]. Warkworth, occurring near to, or at, each reticulation point around Podocarpus totara, 5 Sept 1982, [no collector], #82-267i: margin; with 3-5 setae between stigmatic clefts, each 1/1[ ] ad. a ou 1 μm o g u wi se a us os e io o eac c e a e a ge 15- 3 μm o g S igma ic s i es o a e u i o m ick ess 9 -13 μm o g ema ks. Umbonichiton adelus shares with U. e e : pregenital disc-pores as for genus: number hymenantherae and U. jubatus large cone-shaped macro- present medially/mediolaterally on each segment: anal pores. U. adelus differs from U. hymenantherae in cleft/VII, 0/4-8; VI, 1-2/1-2; V, 1-2/1-2; I , 0/1-2; III, having: 0/1-2; and II, 0/1-3; with 0-3 laterad to each metacoxa. (i) ventral tubular ducts medially; Spiracular disc-pores: with 15-28 in each anterior band (ii) e a a a o e se ae s o (1 - μm o U. adelus and 21-36 in each posterior band. Ventral microducts a 5- μm o U. hymenantherae); present in a wide submarginal band and medially (iii) pregenital disc-pores usually present on all throughout except on posterior abdominal segments. mediolateral folds of the abdomen (in U. Ventral tubular ducts as for genus, except absent from hymenantherae, the pores are rarely present on abdominal segments IV–VII and only occasionally segments II & III). present on abdominal segments II–III. Ventral setae: U. adelus differs from U. jubatus in having: e a a a o e se ae 19-3 μm o g; wi a ai o (i) few finely spinose marginal setae (those of U. jubatus larger, stouter, marginal setae at base of anal cleft, 0 20 are both more spinose and more numerous); μm o g; wi 3- ai s o a e io a a c e se ae; e g (ii) few (0-3) tubular ducts on the median abdominal o o ges ege i a se ae 3 -5 μm; um e o se ae venter (U. jubatus has numerous tubular ducts there). 4 d n & nd r n (2000 : C d (In t : pt r : C d . . U b n h t n d l e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . n f l nd 4 . 6. U b n h t n b ll t e e so & o gso , . s ., a u ema e ci c e co ai s e icu a io i e o o sa o es u e e a ge . 6 d n & nd r n (2000 : C d (In t : pt r : C d . . U b n h t n h n nth r (M ll , n. b., d lt ema e. n f l nd 4 ig. 8. U b n h t n j b t e e so & o gso , . s ., a u ema e. Α si e iew o a u ema e mi us es , s owi g e ma o s. 8 d n & nd r n (2000 : C d (In t : pt r : C d . . U b n h t n p ll p e e so & o gso , . s ., a u ema e. n f l nd 4 io ogy U. d l has been collected only from e e pregenital disc-pores with 5-7 loculi, d rp t t r . The adult females have been found in restricted to mediolateral folds of abdominal segments III– winter through to spring (July–September) and also in the II; um e o eac si e a a c e / IΙ 9- 3; I 5-1 ; summer (January), perhaps suggesting two generations V, 2-9; IV, l-6; III l-2; and II 0; absent from medial per year, abdomen and thorax, and from laterad to metacoxae. Spiracular disc-pores: with 14-51 in each anterior band is i u io Only known from the Auckland region and 22-55 in each posterior band; each band l-5 pores (Map 39). wide near margin. Ventral microducts in a sparse submarginal row and abundant medially. Preantennal ame e i a io The name d l is based on d l pores: with 1-2 near each scape. Tubular ducts as for (Gr. = unseen, unknown, obscure), the scales are well ge us e a se ae e a a a o e se ae 5 - μm hidden at the base of the leaves of d rp t t r , long; with 1-2 pairs of anterior anal cleft setae; length of where the petiole twists a 1/ turn from the stem. o g ege i a se ae 57-77 μm; um e o se ae me ia y on abdominal segments: VII, 5-10; VI, 6-9; V, 10-18; IV, 11-20; III 7-15; and II 3-12; with 5-8 setae near each meso- and metacoxa and 2-5 near each procoxa; with 8-9 (total) interantennal setae. Antennae 6- or 7-segmented, U b n h t n b ll t e e so & o gso ew with 1 pseudosegment when 6-segmented; total length s ecies 15-35 μm; e g o a ica se a - 9 μm C y eo a a s ie 15 -1 μm o g Wi o s i acu a e i emes a e io 5- μm os e io 3 -53 Figs C93, 136 μm egs e g s (me a o acic co a -13 μm; U mou e ma e ia one teneral female known, oc a e + emu 11 -1 μm; i ia 9 -1 3 μm; positioned lengthwise on twig of host plant; colour when a sus 77-1 7 μm; c aw we e e o e 15 μm o g alive chestnut brown; test composed of knobbly wax plates as for genus; dorsal derm with 5 rows of knobbly Ma e ia e ami e O O Y E [ ] NEW ZEALAND: protuberances; shape of adult female elongate. Ο a e e i e ci "Ct n h t n, x W n nn r , Waithui [=Waitui] Saddle, J.M. Hoy, 8 Aug Mou e ma e ia body elongate-oval to oval, slightly 1957", NZAC: 1/2 ad (of two[ ]ad on slide, the holo- narrower at head end; convex, with a lengthwise row of type has complete anal plates, is nearest to the locality rounded derm folds medially and 2 submedian rows each label, and is clearly marked. side, which relate to dorsal knobs of live female; PARATYPE: as for holotype, the other adult female on submarginal row of folds absent. Body length 2.06-2.51 the holotype slide. mm, breadth 1.22-1.94 mm. Other material: NEW ZEALAND: Matakana I o sum dorsal pores distributed in a reticulate pt p r p r , 20 Apr 1961, A.E. Marsack, pattern as for genus; dorsal pore types as for genus except FRNZ R14: 1/1G [ ]ad.Paoneone,2K nz r d , dorsal macropores very large, sclerotised and bollard- Nov 1994, C #94-098: 1/1[ ]a Ο e W ai i a ea shaped: present in median and submedian reticulation d rp t t r , 19 Jan 1970, R.J. McKenzie: 1/1[ ] lines; with l-5 macropores in posterior medial macropore ad. i e A a a es e g 119-157 μm com i e wi s 1 -157 μm; wi - mi u e o es o eac u e ema ks Adult female U. b ll t are characterised by: su ace; e g o se ae i e ma gi 1 1 -1 μm i e (i) 5 rows of dorsal protuberances beneath the test in the ma gi 1 -1 μm a ica se ae 1 -19 μm ou e ma gi position of the reticulation areas, which appear as 5 se ae o g wi a wi e sc e o ise socke -31 μm rows of regular folds on slide-mounted specimens, Anogenital fold with 3 pairs of setae along anterior margin whereas only a median row of such folds can be seen a -1 ai a e a y o ges a ou μm o g on the slide-mounted material of other U b n h t n Ma gi marginal setae variable in shape and size, species; from lanceolate to spinose, with pointed tips; with 9-12 (ii) bollard-shaped dorsal macropores (U. d l , U. setae on each side between stigmatic clefts, length 9-23 h n nth r , and U. j b t all have cone-shaped μm; e icu a io se ae o icea e o ge a e sew e e macropores — U. p ll p has larger, stalked, and S igma ic s i es g a ua y a owi g 5-15 μm o g mushroom-shaped macropores — Aph n h t n 1 d n & nd r n (2000 : C d (In t : pt r : C d n n p also has bollard-like dorsal macropores anal cleft about 1/5 body length. Length 1.96-2.96 mm; but lacks ventral tubular ducts). breadth 1.50-2.10 mm. U. b ll t is close to U. j b t , both having: (i) distinctly spinose marginal setae; Dorsum: dorsal pores distributed in a reticulate (ii) numerous ventral tubular ducts medially; pattern as for genus; dorsal pores of 4 types: (i) small, dark (iii) prominent dorsal derm folds (although they differ in microductules, with a fine, rather short, inner ductule, only form as described above). slightly swollen near pore; (ii) minute simple pores, U. j b t differs, in addition to the shape of the dorsal smaller than microductule pore; both pore types most macropores, in having: frequent in reticulation lines; (iii) slightly larger, convex (i) pregenital disc-pores mediolaterally on abdominal simple pores: present in a row along anal cleft margins, segment II and laterad to the metacoxae (absent on U. and (iv) heavily sclerotised, cone-shaped macropores: b ll t most abundant in reticulation lines around median three (ii) the dorsal macropores very infrequent on the head and rows of reticulation plates; with 0-2 in posterior medial almost restricted to around the median three macropore line. Anal plates: widest in anterior fifth, reticulation areas (more widespread on U. b ll t . a e i g o a e ; e g 1 7-1 7 μm com i e wi s 1 -1 3 μm; wi 3-9 mi u e o es o u e su ace o eac a e; e g o se ae i e ma gi 1 1 -1 μm Biology. Nothing known. i e ma gi 1 -1 μm a ica se ae 1 -1 μm a ou e se ae 1 -1 μm A oge i a o wi ai s o Distribution. Currently only known from the eastern setae along anterior margin and 1 pair laterally, longest central North Island (Map 40). — 53 μm Eyes o o y o e o o e s ecime Margin: marginal setae small and finely spinose, Name derivation. The name b ll t is a Latin adjective mainly present approximately where dorsal reticulation from b ll meaning knobbed, describing the knobbly lines meet margin but with a few occasionally present appearance of the live female. e sew e e eac 7-1 μm o g; wi 3- (a e age se ae on each side between stigmatic clefts; with a rather larger marginal seta at posterior edge of each stigmatic cleft, U b n h t n h n nth r (Maske ew a ou 15 μm o g S igma ic s i es o a e u i o m com i a io ick ess -1 μm o g Venter: pregenital disc-pores with 4-6 loculi, as for Figs C94, 137 genus but usually present only on segments IV-VII Ct n h t n h n nth r Maskell, 1885: 25; - (occasionally on III); number mediolaterally on each side Maskell, 1887: 71 [description]; —Maskell, 1895a: 13 of abdominal segments: anal cleft/VII, 11-33; VI, 1-2; V. [checklist]; —Cockerell, 1896: 330 [checklist]; —Fernald: l-2; IV, 0-2; III, 0-1; and II, 0; none medially or laterad 1903: 160 [world catalogue]; —Hutton, 1904: 226 to metacoxae. Spiracular disc-pores: 20-43 in each [checklist]; —Myers, 1922: 199 [checklist]; —Deitz & anterior band, 26-53 in each posterior band. Preantennal Tocker, 1980: 29 [checklist]; —Ben-Dov, 1993: 102 pores generally present. Ventral microducts of 1 type, [world catalogue]. present throughout including submargin but not on margin. Ventral tubular ducts present in a broad Unmounted material: "Test of adult female waxy, submarginal band; absent medially except 1-3 ducts circular, convex, dirty-white, yellow or brownish, formed occasionally present near coxae, or mesad to spiracles. of a number of hexagonal or octagonal segments, which e a se ae e a a a o e se ae o g - 1 μm; wi are also convex, giving it a rough appearance. Fringe [of 2-4 pairs of anterior anal cleft setae; longest pregenital wax plates] not very conspicuous. Diameter of test, about se ae 1-1 μm; um e o se ae me ia y o a omi a l/12 in. [2 mm]" (Maskell, 1887, p. 71). No fully mature segments: VII, 6-9; VI, 4-7; V, 5-8; IV, 5-7; III, 4-7; and females seen; young adult females as above but not I I, 2-10; with 1-4 setae near each meso- and metacoxa, particularly convex. Colour due to adult female showing and 1-3 near each procoxa; with 5-8 (total) interantennal through thick, glassy wax test; often young female is setae; with 4-9 small submarginal setae on each side bicoloured yellow/brown. between stigmatic clefts. Antennae 6-segmented, with 2 seu osegme s o segme 3; o a e g 9 -3 5 μm; Mounted material: body broadly oval , often with anterior e g o a ica se a 55-73 μm C y eo a a s ie 155- margin nearly straight transversely, rather than rounded; 175 μm o g Wi o s i acu a e i emes a e io - n f l nd 4 8 3 μm os e io 3 -3 μm egs e g s (me a o acic (ii) fewer pregenital disc-pores on segment VII (3-10 as co a 1 -13 μm; oc a e + emu 13 -155 μm; i ia compared with 11-33 on U. hymenantherae); 1 7-1 5 μm; a sus 73-9 μm; c aw 1 -17 μm (iii) s o e a e ae ( 57- 9 μm as com a e wi 9 - 3 5 μm o U. hymenantherae). Material examined: LECTOTYPE [ ](here designated): U. hymenantherae has been recorded from several NEW ZEALAND: "Ctenochiton hymenantherae, adult broad-leaved host plants but not from any Podocarpaceae. female, from Hymenanthera, Aug 1884, W.M.M."; "[gold However, other material studied off Astelia species (a label] "Entomology Div., DSIR, NZ / W.M. Maskell Col- monocotyledenous genus) has a greater number of lection"; "stained & remounted in Canada Balsam, 2 Feb pregenital pores on the mediolateral lobes between the 1995, R.C. Henderson": NZAC: 1/1[ ] ad. anal cleft and posterior spiracles: number mediolaterally PARALECTOTYPE: "antenna and foot of female", Aug on each side of abdominal segments: anal cleft/VII, 20; VI, 1884, W.M.M., NZAC: 1/1 bits. 2-6; V, 7; IV, 6; III, 2; II, 1; none medially but 2 near metacoxae. (AK: Waitakere Ra, Huia, Twin Peaks Ridge, Other material: NEW ZEALAND: Mask. Coll. No. 38, Astelia banksii, 8 Dec 1972, B.M.May, No. 965 (J.A. de USNM: 1/1[ ] ad. "Ex Maskell's dry material, labelled Boer) + #97-145a,b: 3/4[ ] ad). In all other characters Ctenochiton hymenantherae: s and [mm][ ]s",ad and2/4 measured, this material off Astelia appears to agree with U. pieces of wax test", (all in J.A de Boer's handwriting). hymenantherae. It is uncertain whether this variation is AK: Waitakere Ra, Sharps Bush, Hedycarya arborea un- due to host plant influences or whether this material is of derside leaf, 13 Jul 1997, RCH, #97-080: 1/1[m]2nd. GB: another species. Pohutu, Hedycarya arborea, underside leaf, 4 Nov 1993, RCH, #93-353: 1/1[ ]ad.Hedycarya Pohutu, arborea, Biology. Unknown. undersurface leaves, RCH, 15 Mar 1994, #94-046a-d:4/ 1[ ] ad, 1[ ] 3rd, 1 [m]2nd,ad. As previous, 1 Nov1 1994, Distribution. The relatively few specimens available #95-150a: 1/1[ ]ad. As previous, 3 Nov 1995, #95-106: appear to be widely spread from central North Island to the 1/1 [ ] ad. RI: Pohangina Valley, Totara Reserve, Hedycarya southern end of the South Island (Map 41). arborea, underside leaf, 10 Nov 1994, RC & FL Henderson, #94-097: 1/1[ ] ad. NN: Wairoa Gorge, Melicytus ramiflorus, 13 Dec 1967, J.A. de Boer, No.292: U b n h t n j b t e e so & o gso ew 1/2[ ] ad. MC: Port Hills, Sign of Bellbird, Myrsine s ecies divaricata, 17 Feb 1982, #83-321c: 1/[ ]ad. FD: Secre- sp, Mar 1983, C.F. Butcher, #83-293h: 1/ tary I, ?Myrtus Figs M14, 138 1 [ ](pharate). Unmounted material: test of adult female glassy, white, convex with a crest-like median ridge, highest at anterior Remarks. U. hymenantherae shares the character of large end and tapering in a series of undulations towards anal cone-shaped macropores with U. adelus and U. jubatus, plates. Derm of dorsum of adult female similarly ridged but differs from both in: beneath test in a median row of convex folds. (i) a i g o ge e a a a o e se ae ( - 1 μm as com a e wi 19- μm i e o e s ; Mounted material: body of mature female pear-shaped, (ii) the absence of pregenital disc-pores mediolaterally on narrow at head and widest behind posterior stigmatic abdominal segments II and III (present in the other two clefts, with shallow anal cleft about 1/8th body; with derm species). folds in a median line, largest on head, becoming smaller It is also similar to U. adelus in having few, finely towards anal plates. Length 1.0-2.0 mm; breadth 0.7-2.0 spinose marginal setae, but differs in having: mm. (i) many fewer dorsal macropores; (ii) many more pregenital disc-pores on segment VII (11- Dorsum: dorsal pores distributed in a reticulate 33 as compared with 4-8 on U. adelus); pattern as for genus; dorsal pores of 3 types: (i) (iii muc o ge egs ( oc a e + emu 13 -155 μm microductules, with a narrow straight inner ductule, with a com a e wi 5-1 μm o U. adelus). bi- or trifurcate distal end: most obvious near margin but U. jubatus differs from U. hymenantherae in having: also present medially and submedially in reticulation (i) more numerous, broadly spinose marginal setae; lines; (ii) slightly larger, flat simple pores; and (iii) large, ea i y sc e o ise co e-s a e mac o o es ( ig Μ1 182 d n & nd r n (2000 : C d (In t : pt r : C d most abundant in median reticulation lines but with a few 1 pupa, 2[mm] ad, 1 neonate. in lines extending to abdominal submargin; with 8-9 in posterior medial macropore line. Anal plates: widest at Remarks. U. jubatus shares with U. adelus and U. a e io i a e i g o a e ; e g 115-1 μm hymenantherae large cone-shaped macropores but differs com i e wi s 5-1 7 μm; wi 1-5 mi u e o es o in having more numerous and more broadly spinose u e su ace o eac a e; e g o se ae i e ma gi 1 marginal setae (those of U. adelus and U. hymenantherae 1 -1 μm i e ma gi 1 - 1 μm a ica se ae s e e are fewer and only finely spinose). U. jubatus is similar to s i ose a 1 - μm o g a ou e ma gi se ae se ose U. bullatus in: a 19- μm A oge i a o wi 3- ai s o se ae (i) the shape and number of marginal spines; a o g a e io ma gi a wi 1 ai a e a y; e g o (ii) in having dorsal derm folds, but these are restricted to o ges 33-3 μm a median longitudinal line on U. jubatus, whereas U. Margin: marginal setae sharply spinose, mostly 10- bullatus has 5 longitudinal rows of rounded derm 15 μm o g a e a ia e i si e e icu a io se ae o folds; U. bullatus also differs in having bollard-like i e e ia e ; wi 5-1 o eac si e e wee s igma ic dorsal macropores. c e s; wi a a e a ge mo e co ica s i ose se a os e io o eac s igma ic s i e - 5 μm o g Biology. Nothing known. S igma ic s i es qui e o g 11 -1 μm. Venter: pregenital disc-pores with 5-8 loculi, Distribution. Only collected once from central North distributed as for genus; with none medially; number Island (Map 42). mediolaterally on each Side of abdominal segments: anal cleft/VII, 3-10; VI, 3-4; V, 2-4; IV, 2-3; III, 4-5; and II, Name derivation. From jubatus (L. = maned, crested) 1-2; with none laterad to each metacoxa. Spiracular disc- describing the long median crest of the adult female and pores: with 20-26 in each anterior band and 22-30 in each her test. posterior band. Preantennal pore: with 0-2 near each scape. Ventral microducts present in a single row near margin, and medially on head, thorax, and most abdominal U b n h t n p ll p e e so & o gso ew segments. Ventral tubular ducts as for genus. Ventral s ecies se ae e a a a o e se ae 3- μm o g; wi 1- ai s o a e io a a c e se ae; wi 1 pair of long setae discernable on segment VII on young females but not Figs M17, C95, 139 easily discernible on old females; with 4-8 (total) Unmounted material: young adults positioned length- interantennal setae; with 3-5 small submarginal setae on wise on twig of host plant and lateral margins also curved each side between stigmatic clefts; number of setae partly around twig; patterned chestnut and dark brown in medially on each abdominal segment: VII, 5; VI, 5-6; V, life; shape elongate; test composed of wax plates 7; IV, 10; III, 10; and II, 8; with 2-3 setae near each meso- reinforced with some sort of fibrous material. and metacoxa, and 1 near each procoxa; Submarginal setae o g 7-15 μm i e g a ou equa o e g ma gi a Mounted material: body elongate-oval, slightly narrower s i es u a e muc mo e oa A e ae - at head end. Body length 3.20-3.36 mm, breadth 1.37- segme e 3 segme su equa i e g o es o l.84 mm. a e a a wi ou seu osegme s; o a e g 57- 9 μm; e g o a ica se a 3 - μm C y eo a a Dorsum: dorsal pores distributed in a reticulate s ie 1 3-157 μm o g Wi o s i acu a e i emes pattern as for genus; dorsal pores of 3 types: (i) small dark a e io 3- 7 μm os e io 7-3 μm egs e g s pores which appear to lack a ductule: present in a scattered (me a o acic co a 9 -1 μm oc a e + emu 13 - row between macropores within reticulation lines, with 13 μm i ia 104-1 μm a sus 73- 3 μm c aw 1 -15 groups at junctions of lines and near margin; also with an μm occasional pore on submargin between reticulation junctions on margin; (ii) slightly larger pores, which may Material examined: HOLOTYPE [ ]: NEW ZEALAND: be microductules although inner ductules unclear: in a Ο au u ga oa a Pittosporum turneri, 7 Nov 1982, single or double row along anal cleft, and (iii) very large, C.F. Butcher, NZAC #94-110c: 1/1[ ]ad. sclerotised, stalked, bulbous or mushroom-shaped PARATYPES: as for holotype: NZAC: #94-110a,b: 2/2 [ ] macropores (Fig. M17)—outer bulbous part appears ad; #94-110d–m: 10/6[ ] ad (5 pharate), 2[m][ ]3rd,2nd, 3 fibrous like a shaving-brush under light microscope- n f l nd 4 8 protruding above derm surface, and with a basal previous, underside leaves, 31 July 1995, #97-143a—d: 4/ sclerotised tube-like stem; present in median and 1[ ] 2nd, 3[m]2nd. As previous, leaves, 16 Aug 1996, #96- submedian reticulations rows and also in some lines 153a-b: 2/1[ ] 2nd, 2[m]2nd. As previous, leaves, 17 July extending to inner submargin; with 2-6 in posterior 1997, #97-084a-d: 2[ ] 2nd, 1[m]2nd, 1 pharate 1st-2nd, 4/ medial macropore line. Dorsal simple pores apparently 4 1sts. As previous, leaves, 14 Aug 1997, #97-122, -123, absent. Anal plates: widest in anterior third; length 134- -l24a--d: 6/5 [m]2nd, 1 1st. As previous, except on stems, 1 μm com i e wi s 119-13 μm; wi 5-1 mi u e 22 July 1998, #98-083: 2/1 [ ] ad (infested with fungus), 4 o es o eac u e su ace; e g o se ae i e ma gi 1sts (from female brood chamber). 1 1 -1 μm i e ma gi 1 -1 μm a ica se ae 1 - 1 μm a ou e ma gi se ae s im 1 - 1 μm A oge i a ema ks This species can be immediately separated o wi 3 ai s o a e icke e se ae a o g a e io from other species in the genus by the dorsal macropores, ma gi a a si g e ai a e a y o ges 3 -5 μm which are by far the largest macropores of the genus and of Ma gi marginal setae short, finely spinose, 15-19 an unique mushroom shape. μm o g wi - e wee s igma ic c e s a wi a The test contains some fibrous material, which enables a ge s ou e se a us os e io o eac s igma ic c e it to be partially stained and provides sufficient strength to eac a ou 3 μm o g S igma ic s i es a e i g a allow it to be mounted on a microscope slide, so that the qui e o g e g 9 -115 μm structure of the wax plates can be studied. These mounted e e pregenital disc-pores with 7-8 loculi, tests show occasional holes along the reticulation lines, distributed as for genus but absent medially and laterad to presumably where the macropores extend through the test. metacoxae; number mediolaterally on each side of abdominal segments: anal cleft/VII, 15-18; VI, 6-10; V, io ogy Apparently host specific to Podocarpus totara. 5-7; IV, 2-6; III 0-2; and II -1 Spiracular disc-pores: Immature specimens were collected through the autumn with 20-24 in each anterior band and 23-37 in each and winter, with adult females only being collected in late posterior band, each band 2-4 pores wide near margin. winter (July). As the adult is very cryptic and difficult to Ventral microducts very small, numerous in broad bands find, it is not known whether this signifies a univoltine life- across median thorax and abdomen, in a submarginal line cycle. The preadult and adult females were found on the and also present sparsely within submarginal band of stems of the host plant whereas the more immature stages tubular ducts. Tubular ducts as for genus. Ventral setae: were on the leaves. e a a a o e se ae 53-57 μm o g; wi -3 ai s o a e io a a c e se ae; o ges ege i a se ae a ou 77 is i u io Only known from the one site in Auckland μm; um e s o se ae me ia y o eac a omi a area (Map 43). segme II 9-10; VI, 76-10; V, 13-14; IV, 7-9; III 4- 10; and II -1 ; with 4-5 setae near each metacoxa, 5-7 ame e i a io The specific name pellaspis from near each mesocoxa, and 3 near each procoxa; with 5-8 pellis ( = skin) and asilis (f. = a snake), describing the (total) interantennal setae. Antennae 6-segmented, with 2 snakeskin appearance of the live female, her cryptic pseudosegments in long 3rd segment; total length 292- pattern of browns resembling the twig on which she lies. 3 μm; a ica se a a ou 1 μm o g C y eo a a s ie 1 -17 μm o g Wi o s i acu a e i emes a e io 3 -3 μm os e io - μm egs e g s (me a o acic co a 1 7 μm; oc a e + emu 1 -1 μm; i ia 9 -1 7 μm; a sus - 5 μm; c aw 15 μm o g Ma e ia e ami e O O Y E [ ] NEW ZEALAND: AK Riverhead Forest, Barlow Road reserve, Podocarpus totara stem, 31 July 1997, RC Henderson, #97-114c, NZAC: 1/1 [ ]ad. PARATYPES: as for holotype, NZAC #97-114a-b, d—f: 5/2[ ] ad, 1 [m][ ]2nd,tests. 2 Other material: NEW ZEALAND: AK Riverhead Forest, Barlow Road reserve, Podocarpus totara leaves and stems, 4 May 1995, RCH, #95-040a,c: 2/1 [ ] 2nd, 3[m]2nd. As 84 d n & nd r n (2000 : C d (In t : pt r : C d O E S ECIES A E I E S ECIES Ct n h t n l n t Maske ome u ium The key to the genera found in New Zealand is on page 23. All the adventive or exotic species in New Zealand are Ct n h t n l n t Maskell, 1879: 212; -Maskell, cosmopolitan. They have been redescribed many times 1887: 68; –Cockerell, 1893: 548; –Cockerell, 1896: 330;– and good descriptions are available in a number of recent Fernald, 1903: 160; –Hutton, 1904: 226; –Myers, 1922: publications. Because of this, the following descriptions 199; –Wilke, 1927: 207; –Pape, 1939: 346; –Lindinger, are intended to ensure proper identification within the 1943: 218; –Wise, 1977: 104. framework of the New Zealand fauna and are not intended to be exhaustive or to separate these species from other Maskell described C. l n t in 1879. A single slide similar species elsewhere in the world — nor are the exists, labelled "Ct n h t n l n t , from G n t , references exhaustive. For a fuller bibliography see Ben- two females, April 1878, W.W.M.". (stained and Dov (1993). remounted in Canada Balsam, 2nd Feb. 1995, R.C. The fourteen species recorded for New Zealand and Henderson). This slide is the only material which can three additional species erroneously recorded are treated definitely be assigned to the original collection data and here as belonging to six genera. Some of these species has two specimens on it, both 2nd-instar males! (NOT have been placed in other genera at some time and, where females). Unfortunately, they belong to different species! this is the case, this is discussed in the Remarks section No adult females can be located, although the colour plate under each description. These seventeen non-indigenous in Maskell's paper of 1887 (Plate VII, Fig. 4) clearly species belong to two subfamilies (as defined by Hodgson, shows a female test rather than a male test. The original 1994a), namely the Ceroplastinae (5 species) and the material was collected off G n t sp. There are no Coccinae (Tribes: Coccini, 2 species; Pulvinariini, 5 records of any female specimens of any species being species and Saissetiini 5 species). collected off G n t in New Zealand, although undetermined males have from time to time been collected from this plant. Second-instar males often apparently settle on plants which are not the host plant of the adult Su ami y CE O AS I AE females and so the host plant of this stage may be no indicator of the corresponding female. At the present time, iag osis: es : adult females covered in thick coating of neither of the 2nd-instar males on the slide can be wax at maturity; anal plates carried on a sclerotised caudal definitely placed with any known female but, even if both process in order to reach above surface of wax; form and specimens could be, it would leave open the question as to colour of wax often distinctive for a given species. which is actually C. l n t . We therefore consider that Shape: adult female— denuded of wax— usually the best action is to designate C. l n t a n n highly convex, with a distinct dorsal extension to form d b . caudal process, which may be short or longer than rest of In the later papers, Maskell also lists ndr b p. body and which is at least partly heavily sclerotised. and E r n sp. (both Orchidaceae) as hosts. None of this Stigmatic clefts usually distinct; anal cleft extends up material exists. Presumably based on these latter hosts, underside of caudal process to anogenital area. Eggs and various authors (Wilke, 1927; Pape, 1939; Lindinger, 1st-instar nymphs held beneath concave venter. 1943) have reported this species as a pest of orchids o sum: usually highly sclerotised at maturity, and ( ndr b p. and Ep d ndr sp.) in Europe. It is not may have a mid-dorsal, an anterior, and 2 or 3 pairs of known on what basis these identifications were made but it lateral lobes or clear areas, each lacking dorsal pores or is considered here unlikely that they refer to C. l n t setae. Dorsal setae usually sparse, short (often not much — or indeed any of the New Zealand species. longer than width of basal socket), and blunt. Dorsal pores highly characteristic and heavily sclerotised C r pl t type pores, usually with 1-3 satellite loculi and with a long inner filament which branches terminally into numerous fine filaments; these pores may be found throughout or may be absent from some or all dorsal lobes. Preopercular pores generally present in a group imbedded in a sclerotised caudal process anterior to anal plates. Dorsal tubular ducts, dorsal tubercles, and pocket-like n f l nd 4 8 sclerotisations absent. Anal plates together quadrate, the genus C r pl t could only be considered anterior margins usually almost at right angles to long axis monophyletic if all species were included. This is of body; posterior margins convex; inner margins parallel; followed here. each plate with a group of fine apical and subapical setae. Anogenital fold with setae present along anterior and Key o a u ema e C r pl t s ecies i ew ea a lateral margins. Ma gi marginal setae setose and rather sparse. Antennae 7-segmented; tibio-tarsal articulatory sclerosis Stigmatic spines usually abundant, variable in shape, 1 present; claw digitules similar; dorsal pores mainly sometimes of more than one type, th r extending along trilocular n n margin r radially onto dorsum. Eyespots present just dorsad to margin. —Antennae 6-segmented; tibio-tarsal articulatory sclero- e e membranous, usually thrown into large sis present or absent; claw digitules similar or mediolateral folds on either side of genital opening. dissimilar; dorsal pores bi- or trilocular 2 Pregenital disc-pores usually with 10 loculi. Spiracular disc-pores usually with 5 loculi, present in broad bands Stigmatic spines in a compact, oval group, extending up between margin and each spiracle. Ventral microducts onto dorsum; caudal process very broad, particularly distinctive, with a sclerotised, cruciform pore and at its base, where it is more than ½ width of body on generally most abundant submarginally. Ventral tubular mature adults; ventral tubular ducts only present ducts usually present. Ventral setae sparse. Spiracles posteriorly on abdomen d tr t r rather large. Legs usually well developed but may be —Stigmatic spines more or less restricted to along reduced; each with or without a tibio-tarsal articulation margins of stigmatic clefts, extending some way and articulatory sclerosis; each claw with or without a laterad to mid-point in each cleft but not extending as denticle; claw digitules usually similar, with broadly an oval group on to dorsum; caudal process short on expanded apices, but both may be fine when legs reduced. young specimens but, when long, never very broad at Antennae normally well developed, with 6 to 8 segments. base; ventral tubular ducts present anteriorly between Labium normal. antennae as well as posteriorly on abdomen ceriferus ema ks Three species of C r pl t are known from New Zealand: C. r f r (Fabricius), C. d tr t r Newstead, and C. n n l Guercio. Two other species C r pl t r f r ( a icius I ia wa sca e (C. r b n Maskell and C. r (Linnaeus)) are mentioned in the literature but, as there are no voucher Fig. 140 specimens and subsequent authors treated them as misidentifications, these records are here treated as C r f r Fabricius, 1798: 546. doubtful. C r pl t species are all immediately C r pl t r f r (Fabricius); –Walker, 1852: 1087. identifiable by the thick, soft, waxy test which covers the U mou e ma e ia body covered in thick white or dorsum. pinkish-white wax, often irregular in outline, normally with an anteriorly projecting horn, particularly on young specimens. Length 3.0-12.0 mm, width 3.0-10.0 mm. Ge us CE O AS ES G ay Mou e ma e ia more or less oval, derm becoming y e s ecies C (C r pl t j n r n Gray, heavily sclerotised on old females. Caudal process short 1828: 7. on young specimens but becoming more elongate with maturity, up to 1/3rd length of body. C (C r pl t Gray, 1828: 7. C r pl t Gray; –Signoret, 1872: 35; –Wise, 1977: 103 o sum usually with 11 clear areas devoid of pores [checklist]. and setae in submedian band around body, but none The genus C r pl t very large and the boundaries are medially on dorsum. Dorsal setae rather variable but controversial so that some members have also been placed cy i ica eac a ou μm o g C r pl t t p pores in the genera G rd Targioni Tozzetti and W x ll mainly triangular and trilocular, evenly distributed, with De Lotto in the past. However, a phylogenetic study of fewer oval trilocular, quadrilocular, and bilocular pores. their relationships by Qin & Gullan (1995) suggested that Filamentous ducts present around margin. 86 d n & nd r n (2000 : C d (In t : pt r : C d . 40. C r pl t r f r ( a icius , a u ema e — ( om Gim e t l., 4 . n f Ζ l nd 4 8 Ig0,4 4 Ι. 0 i ν ν dd ig. 4 . C r pl t d tr t r ews ea , a u ema e — ( om e o o, 6 . 88 d n & nd r n (2000 : C d (In t : pt r : C d . 42. C r pl t r b n Maske , a u ema e — ( om Gim e t , 4 . n f l nd 4 8 . 4 . C r pl t r ( i aeus , a u ema e — ( om e i a i & Cam o ese, 4 . 0 d n & nd r n (2000 : C d (In t : pt r : C d . 44. C r pl t n n l G r , d lt f l — ( r G p l t l., 4 . n f l nd 4 Ma gi marginal setae rather few and setose, with C r pl t d tr t r ews ea so wa sca e about 2-4 laterally between stigmatic areas, each about 20 μm o g S igma ic c e s s a ow eac wi - u e - Figs C37, 141 s a e s igma ic s i es e e i g a o g ma gi o c e i Ceroplastes destructor Newstead, 1917: 26; –Cottier, i egu a ows 1956: 273, 329 [key; status; host range]; –Ben-Dov, e e pregenital disc-pores with mainly 10 loculi: 1993:30 [world catalogue]. abundant around genital area and on preceding segment, Gascardia destructor (Newstead); –De Lotto, 1965: 200; becoming infrequent on more anterior segments but also –Wise, 1977: 105 [checklist]. present mesad to each coxa. Ventral tubular ducts with a narrow inner ductule: restricted to medially on posterior U mou e ma e ia test of thick, white, soft wax, `wet' four abdominal segments and between antennae. to touch, strongly convex and irregular in shape. Length Antennae 6-segmented. Legs well developed, with a about 3.0-5.5 mm, breadth 1.5-3.0 mm. separate tibia and tarsus but no articulation; claw digitules unequal and claw with a denticle. Mou e ma e ia broadly oval with a caudal process about 1/3rd body length, very broad at base. Length up to Not recorded on indigenous native Ma e ia e ami e I 6 mm. plants. ,and II On exotic plants: Citrus reticulata, Citrus sinensis o sum with 7 clear areas devoid of pores and setae Citrus x `tangelo'. in a submedian band around body; mid-dorsal area with G (all records of primary association, from I few pores and setae. Dorsal setae short and cylindrical to database). slightly clavate. Ceroplastes-type pores fairly evenly distributed, oval and triangular trilocular pores most ema ks Recent descriptions include: De Lotto (1971a), common but with a few bilocular pores. Williams & Kosztarab (1972), Gimpel et al. (1974), Ma gi marginal setae possibly restricted to a few Hamon & Williams (1984), Avasthi & Shafee (1986), and short, flagellate setae on either side of stigmatic clefts. Williams & Watson (1990). Useful identification Stigmatic clefts broad and shallow, each with about 10 characters are the 6-segmented antennae, well developed short, conical spines along margin and further spines legs lacking an articulatory sclerosis, and presence of extending onto dorsum as a conspicuous round to oval ventral tubular ducts between the antennae. group of short, broadly conical spines, each group with 1- First reported from New Zealand in March 1992 2 much larger spines furthest from margin; total number of (source, PPIN database). stigmatic spines per cleft 37-77. This species has a wide distribution throughout e e pregenital disc-pores with mainly 10-12 southern Asia (China, Japan, India, Sri Lanka, Thailand, loculi: restricted to immediately around the genital area. Vietnam, Indonesia, Malaysia, Papua New Guinea, and Ventral tubular ducts restricted to the mediolateral folds the Philippines (Ben-Dov, 1993)). It has been known in laterad to vulva region. Antennae 6-segmented and quite Australia since the last century (as Ceroplastes australiae long. Legs well developed but comparatively rather small Walker, 1852) but it has not yet become a pest (Qin & for body size; lacking a tibio-tarsal articulatory sclerosis; Gullan, 1994). It has also been recorded from the Cook Is, claw digitules dissimilar; claw without a denticle. Fiji, New Caledonia, Tonga, and Vanuatu by Williams & Watson (1990). It was introduced into the USA in 1936 Ma e ia e ami e I Not recorded on indigenous (Gimpel et al., 1974) where it is now widespread and native plants. causes significant damage to ornamentals. It has a large II. On exotic plants: Actinidia deliciosa, Citrus spp. host range—Ben-Dov (1993) lists 39 plant families. io ogy In cool areas, C. ceriferus overwinters as the ND, AK, WO, BP, G adult female. In the spring, it lays a thousand or more eggs which hatch in 2-3 weeks (Williams & Kosztarab, 1972). ema ks This species has been included in the genus The nymphs settle on the twigs or stems, with the female Gascardia by many authors. Recent descriptions include: going through 4 and the male 5 instars. Males are De Lotto (1965), Williams & Watson (1990), Qin & generally considered to be rare (Gimpel et al., 1974). Gullan (1994), and, describing all stages, Wakgari & Giliomee (1998). It is best identified by the very large, is i u io a s a us i ew ea a Known only broad caudal process, the well developed legs lacking a from Gisborne at present; of low economic importance. tibio-tarsal articulatory sclerosis, and the absence of 2 d n & nd r n (2000 : C d (In t : pt r : C d ventral tubular ducts between the antennae. as misidentification]; –Ben-Dov, 1993: 49 [world C. d tr t r was first reported from New Zealand by catalogue]. Greig (1940). It is currently an important pest on citrus in Northland (Smith t l., 1980; Lo & Blank, 1992; Lo t l., C. r b n was reported from New Zealand (Muggeridge, 1992; Olsen t l, 1993; Lo, 1995; Lo t l., 1996; Blank 1933) but there are no voucher specimens, and the record t l., 1997), and Gisborne (D. Steven, pers.comm.). was considered to be a misidentification by Cottier (1939) This species was first described from tropical Africa and Wise (1977). It is treated here as doubtful. It is a rather where it is widespread; its biology was studied in South common and widespread species in the Pacific (Williams Africa by Cilliers (1967). Away from the Ethiopian & Watson, 1990). lt is highly polyphagous and Ben-Dov region, it is known from Australia and from parts of the (1993) lists 72 plant families as hosts. It is considered to Pacific Region — Norfolk Island, Solomon Is., and Papua be a major pest of citrus in Australia (Sabine, 1969; Qin & New Guinea (Williams & Watson, 1990; Ben-Dov, 1993). Gullan, 1994), Hawaii and Japan (Ebeling, 1959), and the The distribution records for India, Florida, Mexico, and Solomon Is (Williams & Watson, 1990). The test is easily Colombia are believed now to be doubtful (T.-K. Qin, identified by the reddish-pink colour of its wax, while pers. comm. 1999). mounted specimens have poorly developed legs—shorter C. d tr t r has a wide host range. Ben-Dov lists 21 than the antennae—and no ventral tubular ducts. plant families, while Brimblecombe (1956) lists 79 plant Recent descriptions include: Gimpel t l. (1974), species from Australia, and Snowball (1969) 106 plant Kawai (1980), Hamon & Williams (1984), Williams & species from New South Wales alone. However, Cottier Watson (1990), and Qin & Gullan (1994). (1956) stated that it was only known on citrus in New Zealand. C r pl t r ( nn f x l Biology. In Queensland, where it was a major pest of citrus before the 1970's (probably due to overuse of Fig. 143 certain pesticides (Smith, 1970)), it has two generations a year (Smith & Ironside, 1974) but only one in the south C r Linnaeus, 1758: 456. (Qin & Gullan, 1994). In New Zealand, the biology has C r pl t r (Linnaeus); –Signoret, 1872: 35; – been studied by Lo t l. (1992), Olsen t l. (1993), and Muggeridge, 1933: 226 [doubtful record]; –Cottier, Lo (1995). In Northland, New Zealand, it has been shown 1939b: 422 [identification correction]; –Wise, 1977: 103 to have only one generation (Lo t l., 1996). Males are [checklist, as misidentification]; –Ben-Dov, 1993: 51 unknown. The crawlers tend to settle on the leaves but [world catalogue]. then migrate to the twigs (late 2nd or early 3rd instar) for This is another C r pl t species which was reported the rest of their lives. Overwintering is by the 3rd-instar from New Zealand by Muggeridge (1933) but there are no nymphs and adults. voucher specimens and the record was considered to be a The possibility of introducing biological control misidentification by Cottier (1939b) and Wise (1977). agents into Australia from South Africa was first studied Although it has been recorded in some tropical areas (Ben- by Snowball (1969); since then several parasitoids and Dov, 1993), C. r is typical of areas with a predators have been introduced and it is currently under Mediterranean climate where it is a pest of figs; it is also a good biological control (Sands t l., 1986; Qin & Gullan, minor pest of citrus in Israel. 1994). It is highly polyphagous and Ben-Dov (1993) lists 30 plant families. It infests the stems, branches, and leaves of Distribution and status in New Zealand. C. d tr t r its host. lts biology in Italy is described briefly by an important pest of citrus in Northland and Gisborne. Pellizzari & Camporese (1994). Its colour and the arrangement of the white stigmatic wax bands in the red wax test make this species fairly easy to identify. Recent descriptions include: Hodgson (1969) and C r pl t r b n Maske e wa sca e Pellizzari & Camporese (1994). Mounted material is best separated from the other C r pl t species considered Fig. 142 here by the presence of 6-segmented antennae, well C r pl t r b n Maskell, 1893a: 214; –Muggeridge, developed legs with a tibio-tarsal articulatory sclerosis, 1933: 226 [doubtful record]; –Cottier, 1939b: 422 and ventral tubular ducts restricted to area between the [identification correction]; –Wise, 1977: 103 [checklist, antennae. n f l nd 4 C r pl t n n e Gue cio a wa sca e o II. On exotic plants: C tr spp., j ll n , C i ese wa sca e G rd n sp., Il x f l , n r sp., h nb r sp. Figs C38, C39, 144 Τ ND, AK, C , WO, BP, GB, HB. C r pl t n n Del Guercio, 1900: 232; —Cottier, 1939b: 422 [description; status; hosts]; —Cottier, 1956: Remarks. Recent descriptions include: De Lotto (1971a), 273, 327 [key; status; hosts]; -Hoy, 1961: 58 [mention; Williams & Kosztarab (1972), Gimpel t l. (1974), distribution]; —Wise, 1977: 104 [checklist]; —Penman, Hamon & Williams (1984), Tremblay (1988), Gill (1988), 1984: 64 [biology; distribution; hosts]; —Scott, 1984: 28 Williams & Watson (1990), Pellizzari & Camporese [hosts]; —Ben-Dov, 1993: 54 [world catalogue]. (1994), and Qin & Gullan (1994). This species has 7- segmented antennae and well-developed legs with a tibio- Unmounted material: young nymphs have small white tarsal articulatory sclerosis. wax plates in a star-like rosette and are normally found on It was first reported in New Zealand in 1932 (Cottier, leaves; mature females with thick, off-white wax test, 1939b) but then declined (Snowball, 1970), although Hoy coloured pinkish brown in patches, and with white `dry' (1961) described it as widespread and recorded it off both wax in the lateral and dorsal depressions; usually on leaves pt p r p r and K nz [as pt p r ] or stems; oval in dorsal view, hemispherical when viewed r d . It is currently a widespread species on citrus in laterally, without a horn; old females lose pinkish-brown Northland but considered to be less destructive than C. colours, becoming grey-white with sooty mould fungus. d tr t r (Lo t l., 1996). Females often clumped together on stems, when shape of C. n n is widespread in the Palaearctic Region, individual specimens becomes distorted. where it is a sporadic pest in Italy and Spain, and is also found in some southern and eastern states in the USA and Mounted material: broadly oval, dorsum only lightly in Mexico, Brazil, and Chile (Ben-Dov, 1993). In the sclerotised; caudal process short. Length 3.5 mm, width Pacific, it has been found on Norfolk Is (Williams & 3.0 mm. Watson, 1990) and has been causing some concern in Australia, where it was first reported in 1966 (Snowball, Dorsum: with 7 clear areas devoid of pores and setae 1970) and where it is now quite widespread. Qin t l. around submargin and with a distinct clear area mid- ( 4 , in an effort to determine a suitable geographic area dorsally. Dorsal setae short and cylindrical to clavate. from which to import biocontrol agents for its control in Oval, trilocular, C r pl t t p pores most numerous, Australia, looked at the relationships of all the but also with triangular trilocular and bilocular pores. Ceroplastinae and concluded that C. n n was probably Margin: marginal setose setae, rather few, with 2-4 native to Central or South America. e wee a e a s igma ic c e s eac u o a ou μm C. n n has a very large host range. Ben-Dov long. Stigmatic clefts shallow, each with about 30 (1993) and Qin & Gullan (1994) list about 50 plant hemispherical, bullet-shaped spines in 2-4 elongate rows families. Cottier (1939b, 1956) lists the following hosts along margin. from New Zealand: citrus, clematis, Escallonia sp., t Venter: pregenital disc-pores with mainly 10 loculi: sp., j ll n , b sp., matipo ( tt p r abundant around vulva region and in two preceding sp.), Ol r sp., puriri ( t x l n , tree tomato (= segments. Ventral tubular ducts sparse posteriorly on tamarillo, C ph ndr b t , and r n sp.; and abdomen and between antennae on head. Long he noted that it had been sufficiently serious on species of filamentous ducts present in a band around margin. Ol r and b to kill them. Since then, the incidence of Antennae 7-segmented. Legs well developed, with a damage to indigenous plants has become minor. distinct tibio-tarsal articulatory sclerosis; claw digitules both broad; claw with a small denticle. Biology. One generation a year in Italy (Frediani, 1960; Pellizzari & Camporese, 1994), Virginia (Williams & Kosztarab, 1972), and New South Wales, Australia Material examined. I. On indigenous native plants: (Snowball, 1970). Young nymphs feed on the leaves and Av nn r n subsp. tr l , C pr rh n then, in the 3rd instar, migrate to the stems to complete d , C pr r b t , C pr sp., Hebe tr t , development (Hamon & Williams, 1984; Snowball, h r p p ln , M l p pl x, t r d 1970). The biology in New South Wales is described by l nt , S l n v l r , t x l n . Snowball (1970) who found that there were 4 female 4 d n & nd r n (2000 : C d (In t : pt r : C d instars and six male (with an extra 3rd-nymphal instar). sclerotisations only present on Saissetiini. Dorsal tubular Qin & Gullan (1994) illustrate a 3rd- instar male with anal ducts present or absent; when present, usually small. Anal plates and mouthparts (appearing more like the 2nd- instar plates together quadrate, each plate usually with outer male than a normal prepupa) and also with small ventral margins subequal in length or with posterior margin wingbuds. Qin & Gullan (1994) did not illustrate a slightly longer; typically with 3 or 4 setae near apex, prepupa or pupa and have indicated (pers.comm.) that they occasionally with setae or spinose setae along inner did not find any prepupae or pupae in the population that margins; discal setae rarely present except on Saissetiini. they studied. Confirmation of the number of male instars is Anogenital fold with pairs of setae present along both required. In New Zealand, the eggs hatch in early March; anterior and lateral margins. Supporting bars to anal plates development is slow during the winter, with the females usually present. Anal ring with 6-10 setae. becoming mature in December (Cottier, 1939b; Penman, Ma gi marginal setae either spinose or setose, the 1984). The young stages are mainly found on the upper latter frequently fimbriate on Coccini and Saissetiini; leaf surface while the adults are on the stems and small usually present in a single marginal line and not extending branches (Cottier, 1956; Penman, 1984). Its biology in up margins of anal cleft. Stigmatic spines: 3 in each cleft, New Zealand has recently been studied by Lo (1995) and clearly differentiated from marginal setae. Eyespots o t l. (1996) and found to be univoltine, with the situated near margin. development of the early nymphal stages much delayed in e e derm membranous. Pregenital disc-pores the spring and early summer in comparison to C. each with 5-10 loculi; on Saissetiini, usually present d tr t r. medially across most abdominal and thoracic segments; on Pulvinariini, typically restricted to abdominal segments is i u io a s a us i ew ea a C. n n is and metathorax; on Coccini, typically on only 1 or 2 an important pest of citrus in Northland and Gisborne, and pregenital segments. Spiracular disc-pores each with 5 of j in Auckland. It has been recorded from Great loculi; present in bands between margin and spiracles. Island (Three Kings Islands) and from most regions of the Ventral microducts present, usually throughout. Ventral northern half of the North Island. tubular ducts highly variable; typically absent on Coccini or, if present, restricted to medially on thorax (e.g., on C on the Saissetiini, present in a broad submarginal band of generally 1 or 2 types (3 on some Saissetiini); on Su ami y COCCI AE the Pulvinariini throughout and generally of 3 or 4 types. Ventral setae normally sparse; with long pregenital setae y e ge us C Linnaeus, 1758: 455. present on 3 pregenital segments. Antennae each with 5- 9 segments. Labium and mouthparts normal. Spiracular iag osis New Zealand species in 3 tribes. Test covering peritremes normal in size. Legs usually well developed; dorsum either absent, extremely sparse or of thin wax; each leg with separate tibia and tarsus, with or without an woolly ovisac produced ventrally on Pulvinariini, in articulatory sclerosis (typically present on Pulvinariini); which eggs and 1st-instar nymphs are protected; in other claw digitules usually both broad; each claw with or tribes eggs and/or 1st-instar nymphs are held under without a denticle. concave venter. Body oval and usually rather flat, less commonly highly convex when mature (e.g., many Saissetiini); stigmatic clefts rather shallow or absent; anal cleft normal; occasionally with an anal sclerotisation around anterior margin of anal cleft. i e Cocci i a é o sum derm quite thin (thick with dermal y e ge us C Linnaeus, 1758: 455. areolations on Saissetiini), sclerotised at maturity. Dorsal setae typically spinose. Dorsal pores variable, but Members of this tribe in New Zealand are characterised generally including a dorsal microductule. Preopercular by: (i) absence of ventral tubular ducts or their restriction pores generally present: on Pulvinariini usually in a single to medially on thorax; (ii) lack of pocket-like band extending anteriorly from anal plates; spreading sclerotisations; (iii) preopercular pores in an elongate around sides of anal plates on Saissetiini; each pore band anterior to anal plates; (iv) pregenital disc-pores on usually small, but large, convex and heavily sclerotised on 1-3 pregenital segments only. Saissetiini. Dorsal tubercles present or absent; when present, convex and submarginal. Pocket-like n f l nd 4 Ge us COCCUS i aeus series of brown spots which take on a distinct pattern; mature adults finally turning dark brown, slightly convex. C Linnaeus, 1758: 455. Type species: C h p r d Linnaeus, 1758, Mounted material: elongate-oval, up to 6.0 mm long and designated in Opinion 1303 under the plenary powers of 4.0 mm wide. the International Commission on Zoological Nomenclature (name number 2244) (1985). Dorsum: derm membranous, becoming mildly sclerotised with small pale areolations on older specimens. Remarks. Two species currently placed in the genus Dorsal setae small, cylindrical with blunt apices. Dorsal C are known from New Zealand, namely C pores frequent throughout, with a minute microductule in h p r d L. and C l n l (Douglas). The each areolation. Preopercular pores flat, in a diffuse group former is of some importance on horticultural crops. in front of anal plates. Dorsal tubular ducts present or absent; when present, scarce submarginally, sometimes singly but with up to 6 or more per side. Dorsal tubercles Key o a u ema e C s ecies i ew ea a almost invariably present submarginally; each tubercle Antennae 8-segmented; ventral tubular ducts absent; small, simple, and convex; with 0-6 on each side. Anal dorsal setae short, curved, and sharply pointed plates rather pointed; posterior margins usually with a l n l small indentation about 1/2-1/3rd way along length. —Antennae 7-segmented; ventral tubular ducts present Anogenital fold with 2 pairs of long setae along anterior medially on pro- and mesothorax; dorsal setae blunt margin and 2 pairs laterally. and cylindrical h p r d Margin: marginal setae finely spinose with flattened fimbriate apices but setae can appear pointed; with 8-14 setae on each side between stigmatic clefts. Stigmatic clefts usually slightly indented, with 3 stigmatic spines; C h p r d i aeus so ow sca e median spine generally slightly curved, with a blunt apex and 2-3 x length of lateral spines, which are usually Figs C40, C41, 145 straight with a blunt apex. Venter: pregenital disc-pores with mainly 10 loculi: C h p r d Linnaeus, 1758: 455; –Fernald, 1903: sparse immediately around genital opening and with a few 168 [world catalogue]; –Thomson, 1922: 334 [pest status; mediolaterally on preceding 1 or 2 segments. Spiracular hosts]; –Myers, 1922: 199 [checklist]; –Miller, 1935 disc-pores in a narrow band. Ventral microducts small and [status; biology]; –Cottier, 1956: 272, 331 [key; status; rather sparse throughout. Ventral tubular ducts present in hosts]; –Wise, 1977: 104 [checklist]; –Penman, 1984: 61 a rather distinct pattern, with a small group of 0-3 just [biology]; –Somerfield, 1984: 87 [hosts]; –Ben-Dov, mesad to each procoxa, a larger group near each mesocoxa 1993: 73 [world catalogue]. and usually extending thinly across median area of n h p r d (Linnaeus); –Maskell, 1879: 205 mesothorax, and with 0-2 also present lateral to anogenital [description; biology; pest status]; –Maskell, 1887: 80 fold. Antennae 7-segmented. Legs well developed; each [description; records]; –Maskell, 1893a: 218 [synonymy with a distinct tibio-tarsal articulation and a small discussion]; –Maskell, 1893b: 103 [synonymy; status]; – articulatory sclerosis (occasionally absent on some legs); Maskell, 1895a: 15 [checklist]; –Hutton, 1904: 353 claw digitules both broad though one occasionally slightly [checklist]. thinner than other; claws without a denticle. n l t Signoret, 1873b: 400; –Maskell, 1879: 207 [brief description]; –Maskell, 1887: 81 Ma e ia e ami e I O i ige ous native a s [description; records]; –Maskell, 1895a: 16 [checklist]; – l h d t , l hn fr r , r h l tt Hutton, 1904: 353 [checklist]. b ll d d , r h l tt r p nd , C r h l s C l t (L.); –Fernald, 1903: 172 [world Ch rd p rt t v n n , C pr t f l , catalogue]; –Thomson, 1922: 334 [mention]; –Myers, C pr s C r s C th s M p r 1922: 199 [checklist]; –Wise, 1977: 104 [checklist]. l t , Ol r n l r f l , Ol r tr v r , Unmounted material: body elongate-oval to nearly l s tt p r s r t ph l d , d round, sometimes asymmetrical if against leaf vein; rather p n x r f l , d p n x l n , d p n x flat, with a series of faint lateral ridges; markings highly s S n s S l n v l r , nth variable, young adults generally pale green, developing p , t x l n . 6 d n & nd r n (2000 : C d (In t : pt r : C d ig. 4 . C h p r d i aeus, a u ema e — (A a e om o gso , 4a . n f l nd 4 ίί ίg. 46. C l n l ( oug as , a u ema e — ( om Gi , 88 . 19 d n & nd r n (2000 : C d (In t : pt r : C d II. On exotic plants: Ab t l n sp., A t n d d l , is i u io a s a us i ew ea a C. h p r d Alt rn nth r ph l x r d , v rd sp., "broom", has been recorded from the Kermadec Islands and C tr spp., C rb t x , C b d sp., ndr Chatham Islands as well as throughout the North and b sp., nth sp., t sp., sp., G rd n South Islands where it is a serious pest of ornamentals, sp., d r sp., b t l , Il x sp., Laurel, both indoors and outdoors. It has also been recorded from l sp., M l x d t , M d t v , r at least 22 indigenous plant species. r n , h l n p sp., hl x sp., n r d t , n r tr f l t , r t nth r n t , r n r n , r n v , r n l r r , r n p r var n p r , r n l n , b n r , C l n l ( oug as o g ow sca e r n sp, b x [as loganberry], S ll h t r ph ll , t v n f r . Figs C42, 146 KE / AK WO G K WA WI W / n l n l Douglas, 1887: 97; -Maskell, 1891: S M W MC SC O CO S / C 16 [synonymy]; -Maskell, 1893a: 221 [Sandwich Is]; - Maskell, 1897: 310 [first record]. ema ks Recent descriptions of C. h p r d can be n h r l Maskell, 1890c: 137; -Maskell, found in: Williams & Kosztarab (1972), Gill t l. (1977), 1891: 16 [synonymy]; —Fernald, 1903: 171 [world Kawai (1980), Hamon & Williams (1984), Gill (1988), catalogue]. Williams & Watson (1990), and Hodgson (1994a). The 7- C l n t (Signoret); —Fernald, 1903: 168 [world segmented antennae and the distribution of the ventral catalogue]; —Wise, 1977: 104 [checklist]. tubular ducts are particularly good identification C l n l (Douglas); —Fernald, 1903: 171 [world characters, along with such anal plate characters as lateral catalogue]; —Thomson, 1922: 334 [status]; —Myers, 1922: margin indentation, 4 pairs of small setae near apex, and 199 [checklist]; -Wise, 1977: 104 [checklist]; —Ben-Dov, noticeably sclerotised supporting bars. 1993: 80 [world catalogue]. C. h p r d is one of the most cosmopolitan and polyphagous insects (Ben-Dov (1993) lists 91 plant U mou e ma e ia elongate oval, moderately convex families), but Gill (1988) states that it has been found on and smooth; yellowish to greyish-brown. almost every kind of plant except grasses. It is an important pest on citrus and ornamentals, including those Mou e ma e ia elongate, up to 6 mm long and 3 mm under glass. C. h p r d was first reported in New wide. Zealand by Maskell in 1879, when he considered that it o sum derm membranous to slightly sclerotised, was becoming a veritable pest on introduced garden trees. with conspicuous areolations. Dorsal setae often curved, Cottier (1956) considered that it had a wide host range but sharply pointed. Dorsal microductules present in each was of minor importance, while Penman (1984) stated that areolation. Dorsal tubular ducts absent. Dorsal tubercles it was often a problem on citrus in the northern North present, totalling up to about 19. Preopercular pores in an lsland. An encyrtid parasitoid, M r t r fl v elongate group in front of anal plates. Anogenital fold (Howard), was introduced to N.Z. for its control in 1921 with 4 pairs of setae along anterior margin and 4 pairs (Hill, 1989). laterally. io ogy C. h p r d has 3-5 generations a year Ma gi marginal setae setose, rarely with frayed outdoors in California and more under protective apices, with about 12 between lateral stigmatic clefts. cultivation (Gill, 1988); it is mainly parthenogenetic and Stigmatic clefts shallow, with three stigmatic spines, ovoviviparous. It attacks leaves and twigs, and usually is median spine 2 to 3 times longer than lateral spines. associated with large amounts of honeydew, resulting in a Eyespot set slightly onto dorsum in a clear area of derm. thick covering of sooty mould, which is often more e e pregenital disc-pores with mainly 7 loculi: important than any physical damage done by the insect. abundant around vulva and occasionally on 1 or 2 Where toxic chemicals are infrequently used, it is usually preceding segments. Ventral microducts sparse kept under control by natural enemies. Penman (1984) throughout, most common near labium. Ventral tubular notes that generally within an orchard of subtropical fruit ducts absent. Antennae 8-segmented. Legs well trees, large numbers of C. h p r d may be found in developed, with a tibio-tarsal articulatory sclerosis. very small areas on a single tree or only on isolated trees. There are 3-4 (Miller 1935) or possibly up to 5 (Penman Ma e ia e ami e I On indigenous native plants: 1984) generations per year in New Zealand. C r h l tr l , C r h l sp., n f l nd 4 Ch rd p rt t v n n , M l p t rn t . Key o a u ema e Saisse ii i i ew ea a II. On exotic plants: C tr sp., p r sp., M n l 1 Pregenital disc-pores present medially on thorax and t ll t , t v n f r . head as well as on abdomen; dorsal setae of distinctly ΑK, M . two sizes, largest setae forming a mid-dorsal band anterior to anal plates; ventral tubular ducts of two Remarks. Recent descriptions—often as C. l n t types, with a smaller type completing the submarginal (Signoret)—include: Ben-Dov (1977), Gill t l. (1977), band between antennae . rth n l n spp 2 Kawai (1980), Hamon & Williams (1984), Gill (1988), —Pregenital disc-pores not present anteriorly to and Williams & Watson (1990). The presence of 8- metathorax; dorsal setae of one size, evenly segmented antennae, slender, pointed, often curved dorsal distributed over dorsum; larger ventral tubular ducts setae, and absence of dorsal and ventral tubular ducts forming a complete submarginal band between quickly separate this species from C. h p r d . antennae 3 A cosmopolitan species found in most countries with a warm climate or under protected cultivation. It has a large 2 Submarginal band of ventral tubular ducts with three host range— Ben-Dov (1993) lists 45 families. It was first types of duct, one of which has inner ductule as wide reported in New Zealand by Maskell in 1897 when he as or wider than outer ductule; ventral microducts thought it likely that it would become established in the abundant between band of ventral tubular ducts and warmer regions; the record by Dale t l. (1976) was not margin; legs with a tibio-tarsal articulatory sclerosis therefore the first. Parthenolecanium persicae Biology. This species is parthenogenetic (Ben-Dov, 1977, —Submarginal band of ventral tubular ducts with only 1993). It is generally found on the leaves, branches and two types of duct, neither of which has inner ductule as twigs. It is rarely of economic importance. broad as outer ductule; ventral microducts very scarce or absent between submarginal band of tubular ducts Distribution and status in New Zealand. C. l n l is and margin; legs without a tibio-tarsal articulatory seldom collected in New Zealand; most records are from sclerosis Parthenolecanium corni the Auckland area. 3 Anal plates without setae in discal position; dorsal setae spinose with clavate apices; legs without a tibio-tarsal articulatory sclerosis; dorsum with numerous polygonal cell-like areas Parasaissetia nigra —Anal plates each with a discal seta; dorsal setae spinose with pointed apices; legs each with a tibio-tarsal i e Saisse ii i o gso articulatory sclerosis; dorsum never with polygonal Type genus: S t Déplanche, 1859. cell-like areas but with distinct dermal areolations ... S t spp.... 4 Members of this tribe in New Zealand differ from others in the Coccinae in having: (i) a broad submarginal band of ventral tubular ducts of 1 or 2 types; (ii) dorsal tubercles 4 Submarginal band of ventral tubular ducts of one type and often pocket-like sclerotisations; (iii) pregenital disc- only, none with inner ductule as wide as outer ductule pores, usually each with 10 loculi, present medially on Saissetia oleae thorax as well as abdomen; (iv) preopercular pores in a —Submarginal band of ventral tubular ducts with three group around or anterior to anal plates; and (v) dorsal types of duct, one of which has inner ductule as wide tubular ducts generally absent (present on as outer ductule Saissetia coffeae rth n l n rn . Remarks. In New Zealand this tribe contains three genera and five species: r t nί r (Nietner), rth n l n rn (Bouché), rth n l n p r (Fabricius), S t ff (Bernard), and S t l (Olivier). 200 d n & nd r n (2000 : C d (In t : pt r : C d ig. 4 . r t n r ( ie e , a u ema e — (A a e om o gso , 4a . n f l nd 4 20 ig. 48. rth n l n rn ( ouc é , a u ema e — (A a e om o gso , 4a . 202 d n & nd r n (2000 : C d (In t : pt r : C d ig. 4 . rth n l n p r ( a icius , a u ema e ( om Gi , 88 . n f l nd 4 20 . 0. S t ff (Wa ke , a u ema e — (A a e om o gso , 4a . 204 d n & nd r n (2000 : C d (lIn t : pt r : C d . . S t l (O i ie , a u ema e — ( om Gi , 88 . n f l nd 4 5 Ge us A ASAlSSE lA aka as i typically cylindrical, with a clavate apex; frequent throughout. Dorsal microductules each in a polygonal r t Takahashi, 1955: 26. areolation. Preopercular pores highly convex, round, Type species: n n r Nietner, 1861. sclerotised, with a granular surface: present in a small group of 6-23 pores immediately anterior to anal plates. Dorsal tubular ducts absent. Dorsal tubercles fairly small r t n r ( ie e ig a sca e and convex, with 1-26 in a submarginal ring. Pocket-like sclerotisations generally present, total 0-17. Anogenital Figs C67, 147 fold with 6-8 setae along anterior margin and 3 or 4 pairs setae laterally. Anal ring with 8 setae. n n r Nietner, 1861: 9; –Maskell, 1894a: 166 Margin: marginal setae each often apically flattened, [synonymy]; –Maskell, 1894b: 73 [mention]; –Hutton, broadened, and frayed; with 11-23 setae on each side 1904: 353 [checklist]; –Green, 1929: 376 [record]. between stigmatic clefts. Stigmatic clefts shallow, each n d pr Targioni Tozzetti, 1867: 29; – with 3 stigmatic spines; median spines 2 x to 4 x length of Maskell, 1879: 206 [description]; –Maskell, 1887: 79 lateral spines, all rather blunt. [description; records]; –Maskell, 1893a: 220 [description; Venter: preopercular pores with mainly 10 loculi; status]; –Maskell, 1894a: 166 [synonymy]; –Maskell, abundant around genital opening, becoming progressively 1894b: 73 [mention]. less frequent on preceding abdominal segments. Ventral n n r νa d pr Targioni Tozzetti; – tubular ducts of one type: in a submarginal band. Maskell, 1895a: 16 [checklist]. Antennae 8-segmented but occasionally with only 7 n (S t n r Nietner; –Cockerell & segments. Legs well developed; each with a separate tibia Parrott, 1899: 163; –Green, 1929: 376 [record]. and tarsus but no articulatory sclerosis; claw digitules both S t n r (Nietner); –King, 1902: 296; –Fernald, broad and marginally shorter than tarsal digitules; claws 1903: 204 [world catalogue]; –Thomson, 1922: 335 without a denticle. [mention]. S t d pr (Targioni Tozzetti); –Fernald, 1903: Material examined. I. Not recorded on indigenous native 201 [world catalogue]; –Myers, 1922: 200 [checklist]; – plants. Wise, 1977: 106 [checklist]. r t n r (Nietner); –Takahashi, 1955: 26; – II. On exotic plants: `broom', C tr spp., phn sp., Wise, 1977: 106 [checklist]; –Ben-Dov, 1993: 209 [world j ll n , Il x sp., Ir r n , r n catalogue]. r n . Unmounted material: young females usually elongate ΑΚ ΤΟ WI W / ΝΝ M MC O oval and flat to rounded and convex, with elongate individuals usually on twigs and rounded ones on leaves. Remarks. Recent descriptions include: De Lotto (1967), Dorsum of older, darkened females smooth, without "H" Ezzat & Hussain (1969), Ben-Dov (1978), Kawai (1980), pattern, polygonally reticulated and often with a single Hamon & Williams (1984), Gill (1988), Williams & series of staggered, squarish white wax plates around Watson (1990), and Hodgson (1994a). The reticulated margin and with 5 longitudinal dorsal rows of smaller wax pattern on the dorsum separates this species from all others plates. Shape and colour varies with type of host and considered here. location on host. Young adult females translucent yellow, . n r is a cosmopolitan species but is mainly occasionally with brown or red mottling, becoming shiny restricted to the warmer countries, unless under glass. It and dark brown to purple-black with age. Young females has a huge host range, Ben-Dov (1993) listing 81 host about 2.0 mm long but growing to about 5.5 mm long and families. It is mainly a pest on ornamentals. 4.0 mm wide. It was first reported from New Zealand by Maskell (1879) — as n d pr Targioni Tozzetti— Mounted material: body almost round to elongate oval, occurring in greenhouses. with shallow stigmatic indentations: about 2.0-3.5 mm long and 1.5-3.2 mm wide. Biology. . n r has 1 generation outdoors in California but up to about 6 in greenhouses in Israel (Ben-Dov, Dorsum: on all but newly moulted adults, derm with 1978). Reproduction is parthenogenetic. It is found on the numerous polygonal areas except at extreme margins; leaves, twigs, branches, and fruits. In California, this each polygon with a central areolation. Dorsal setae species overwinters as the 2nd- or 3rd-instar nymphs (Gill, 206 d n & nd r n (2000 : C d (In t : pt r : C d 1988). See Smith (1944) for a study of its biology, etc., in Dorsum: derm becoming evenly and heavily California. sclerotised at maturity; lacking cell-like areolations. Dorsal setae of 2 sizes: rather large, stout, blunt spines, Distribution and status in New Zealand. Sporadic eac 1 -3 μm o g ese i a mo e o ess ou e i e throughout North and South Island — not a serious pest. medially anterior to anal plates extending as far forward as mouthparts; much smaller, rather blunt, spines, each about 5-1 μm o g a e s a se oug ou es o o sum Ge us A E O ECA IUM Š ulc Dorsal pores of 2 types: present throughout. Preopercular pores circular, moderately large, with a rough surface: Parthenolecanium Šulc, 1908: 36. present in a small loose group of 6-26 pores just anterior Type species: Lecanium corni Bouché, 1844, designated to anal plates. Dorsal tubular ducts present though by Opinion 1303 under the plenary powers of the frequency highly variable, ranging from abundant to International Commission on Zoological Nomenclature sparse or even absent. Dorsal tubercles normal, large, and (1985). convex; total 0-18 in a submarginal band. Pocket-like sclerotisations varying from 0-7 pairs; when present, more or less within submarginal band of dorsal tubercles. Anogenital fold with 2 pairs of long setae along anterior margin and 2 pairs of shorter setae laterally. Anal ring with rth n l n rn ( ouc é ow sca e o 8 setae. Margin: marginal setae each bluntly spinose; present Eu o ea ui eca ium more or less in 2 lines, rather unevenly spaced, with 11-20 Figs C68, 148 setae on each side between stigmatic areas. Stigmatic Lecanium corni Bouché, 1844: 298; –Wise 1977: 105 clefts absent. Stigmatic spines all bluntly spinose, in [checklist]. groups of 3 in each stigmatic area, each median spine Lecanium ribis Fitch, 1857: 427; –Maskell, 1891: 16 about 1.5 x longer than laterals and generally slightly [description; status]; –Maskell, 1892: 22 [pest status]; – curved. Maskell, 1895a: 16 [checklist]; –Maskell, 1898: 237 Venter: pregenital disc-pores with mainly 10 loculi; [record]; –Hutton, 1904: 353 [checklist]; -Thomson, fairly abundant around genital opening, becoming 1922: 335 [mention]. progressively less frequent across preceding abdominal Lecanium rosarum Snellen von Vollenhoven, in De Graaf segments; with small groups also mesad to each coxa and et al., 1862: 94; –Maskell, 1892: 22 [pest status]; – laterad to each metacoxa. Ventral microducts abundant in Maskell, 1895a: 17 [checklist]; –Thomson, 1922: 335 a submarginal band and much less frequent medially, [mention]. particularly on abdomen. Ventral tubular ducts of 3 types Eulecanium corni (Bouché); –Fernald, 1903: 185 [world present: (i) small duct: present in small submarginal catalogue]; –Myers, 1922: 199 [checklist]; –Miller, 1935 groups between antennae and on either side of anal cleft; [status; biology]. abundance highly variable between specimens; (ii) Lecanium (Eulecanium) corni (Bouché); –Brittin, 1940a: slightly larger duct, with outer ductule rather longer than 411 [mentions]. type-(i), with a large terminal gland: rather sparse Parthenolecanium corni (Bouché); –Borchsenius, 1957: mediolaterally on head, thorax and abdomen; and (iii) 356; –Ben-Dov, 1993: 214 [world catalogue]. fairly large duct: in a broad submarginal band extending from anterior to each antenna to near anal cleft. Antennae Unmounted material: shape and colouration extremely 7-segmented (rarely 6 or 8). Legs normally developed; variable. Mature females chestnut-brown and leathery, each with a separate tibia and tarsus but no articulatory varying from slightly convex to pyramidal or hemispherical sclerosis; claws rather long and narrow with a minute in profile. Stages on leaves semi-transparent yellow- denticle; claw digitules dissimilar, one broader than the green; stages on twigs mottled yellow and brown, often other. with darker bands on abdomen. Eggs white. Young adult Material examined. I. On indigenous native plants: females and 2nd- and 3rd-instar nymphs on twigs well Aristotelia sp. camouflaged and difficult to detect. II. On exotic plants: Prunus sp., Prunus armeniaca, Mounted material: elongate oval, widest in abdomen; Prunus avium, Quercus palustris, Ribes nigrum, Vitis length up to 4.3 mm and width up to 3.0 mm. vinifera. n f l nd 4 20 BP, WO, HB, WN ΜΒ, CO. Hutton, 1904: 353 [checklist]; –Thomson, 1922:334 [checklist]. Remarks. Recent descriptions include: Williams & n p r (Fabricius); –Comstock, 1883: 134; – Kosztarab (1972), Kawai (1980), Hamon & Williams Green, 1929: 376 [record]; –Wise, 1977: 105 [checklist]. (1984), Danzig (1986), Kosztarab & Kozár (1988), Gill E l n b rb r (Schrank); –Fernald, 1903: 182 (1988), and Hodgson (1994a). The presence of [world catalogue]; –Miller, 1935: 37 [description; status]. multilocular disc-pores medially on all thoracic segments n (E l n p r (Fabricius); –Brittin, and two sizes of dorsal setae identifies this genus, while 1940: 411 [mention]; –1940b: 413 [life history; the absence of a type of ventral tubular duct with a very description immatures]. broad inner ductule separates . rn from . p r . n (E l n p r p n Brittin, Distributed mainly in the Palaearctic, Nearctic, and 1940b: 420; –Wise, 1977: 105 [checklist]; syn. nov. Holarctic; within the Australasian and Pacific regions, E l n p r (Fabricius); –Cottier, 1956: 272, only known from New Zealand and Australia. 333 [key; description; status; biology]. Polyphagous — Ben-Dov (1993) lists 40 plant families. rth n l n p r p n (Brittin); –Ben- Highly polymorphic, appearance varying depending on Dov, 1993: 224 [world catalogue]. the host plant, the part of the plant attacked, and the age of rth n l n p r (Fabricius); –Borchsenius, the scale (see Ebeling, 1938). Often an important pest of 1957: 350; –Ben-Dov, 1993: 221 [world catalogue]: deciduous fruit, vines, and ornamentals. Unmounted material: highly variable; not strongly First reported in New Zealand by Maskell in 1891 (as convex, elongate oval with a medial longitudinal ridge. n r b Fitch). Wise (1977) included . r as a Young adult females usually yellowish with brown synonym of E l n ( rth n l n rn but . markings or mottling, becoming uniformly brown with r is actually a synonym of . p r in addition, age. Colour of eggs unkown. Wise (1977) included . r r under . p r but . r r is a synonym of . rn (Ben-Dov 1993). It is Mounted material: elongate oval, with distinct stigmatic possible that some of the early records (such as those of clefts; length up to 5.0 mm and width to 3.0 mm. Hutton (1904), Thomson (1922), and Myers (1922)) could refer to either species. Dorsum: derm membranous when young, becoming mildly sclerotised when old. Dorsal setae of 2 sizes: rather Biology. Usually has only a single generation in the USA, large, stout, blunt spines: present in a more or less double where it overwinters as a 2nd-instar nymph on the twigs line medially anterior to anal plates extending as far and branches, remaining there to mature in the spring. The forward as mouthparts; much smaller, rather blunt, spines: resultant crawlers disperse to the leaves but return to the rather sparse throughout rest of dorsum. Dorsal pores of 2 twigs and branches in the autumn. On evergreen hosts, the types, present throughout. Preopercular pores circular, entire life cycle can occur on the leaves (Gill, 1988). The moderately large, with a rough surface: present in a small life cycle in southern England is described by Birjandi loose group of about 20-26 pores just anterior to anal (1981) and Habib (1955a, 1955b). plates. Dorsal tubular ducts absent. Dorsal tubercles normal, large, and convex; total of 24-42 around Distribution and status in New Zealand. . rn is a submargin. Pocket-like sclerotisations probably present minor pest of plums, apricots and grapevines in both the (included in illustration by Gill, 1988). Anogenital fold North and South Islands. with 2 pairs of long setae along anterior margin and 2 pairs of shorter setae laterally. Anal ring with 8 setae present. Margin: marginal setae long, slender, curved, and rth n l n p r ( a icius eac sca e pointed, in a single band; with about 8-12 setae on each side between stigmatic areas. Stigmatic clefts distinct, each with 3 stigmatic spines, all sharply spinose and about Fig. 149 as long as marginal setae; each median spine slightly Ch r p r Fabricius, 1776: 304. longer than laterals and generally slightly curved. C b rb r Schrank, 1801: 146. Venter: pregenital disc-pores with mainly 10 loculi; n r Signoret, 1874: 407; –Maskell, 1885: 29 fairly abundant around genital opening, becoming [description; status]; –Maskell, 1887: 82 [description; progressively less frequent across preceding abdominal records]; –Maskell, 1894b: 75 [description]; –Maskell, segments; with large groups also present mesad to each 1895a: 16 [checklist]; –Maskell, 1896: 392 [status]; – coxa. Ventral microducts abundant in a submarginal band 208 d n & nd r n (2000 : C d (In t : pt r : C d and near labium, much less frequent medially, particularly A study of the available material of n on abdomen. Ventral tubular ducts of 3 types: (i) small (E l n p r p n Brittin (1940b) indicates duct: present in small submarginal groups between that this is a synonym of . p r (Fabricius) (sy antennae; (ii) a slightly larger duct, with outer ductule o It was collected off W t r [published as rather longer than type-(i), with a large terminal gland: `W t r ] sp. There is some doubt about the published rather sparse, intermixed with type-(iii); and (iii) large collection locality of Ngatea (WO) that differs from the duct, with an inner ductule as wide as or wider than outer localities given on the slides—variously Ngongotaha (BP) ductule: present in a broad submarginal band extending and Waihi (BP)—but these three sites are all within a from anterior to each antenna to near anal cleft. Ventral relatively small area of the North Island, and the chosen setae: submarginal setae in a double row. Antennae 8- or lectotype specimen agrees closely with Brittin's 9-segmented (rarely 6- or 7-segmented). Legs normally description. developed; each with a tibio-tarsal articulatory sclerosis; claws rather long and narrow with a distinct denticle; claw io ogy Probably almost entirely parthenogenetic, digitules broad and similar. although males are known. Has l or 2 generations a year, overwintering as the 2nd-instar nymph on the twigs or Ma e ia e ami e I Not recorded on indigenous stems. The subsequent crawlers disperse to the leaves and native plants. the next generation can be on the leaves. II. On exotic plants: " n b rb r d , [no host or is i u io a s a us i ew ea a It is locality], 1896, W.M.M.": 1/1 `head & antenna'; . occasionally a minor pest of citrus and grapevines; p r Fab., det. J.G. Saunders, Nov. 20 1909, currently known from the southern half of North Island to Washington, D.C.; 1/1 '2nd-stage female'; 1/1 `larvae'; the northern half of South Island. NZAC. n (E l n p r (Fab.) p n Brittin: LECTOTYPE [ ]here'Eulecaniump r designated: Ge us SAISSE IA é a c e Fabr. variety, No. 248, on W t r , Ngongotaha, 4 Jan 1934, " G. Brittin Collection; NZAC: 1/1 young [ ] S t Déplanche, 1859: 6. ad. n ff Walker, 1852, proposed PARALECTOTYPES: 2/2 old [ ] ad.; collection data as y e s ecies by Ben-Dov, 1989: 116, approved by the International above except locality data: 1/ "Ngongotahi Waihi"; 1/ Commission on Zoological Nomenclature (Opinion 1627, "Waihi". 1991). III. From PPIN database (all primary associations): C tr spp., G W M SC t v n f r , HB. S t ff (Wa ke emis e ica sca e ema ks Recent descriptions include: Williams & Kosztarab (1972), Kawai (1980), Hamon & Williams Figs C88, C89, 150 (1984), Danzig (1986) (as . th , Gill (1988), and Kos a a & Κo á (19 e ese ce o mu i ocu a n ff Walker, 1852: 1079. isc- o es me ia y o a o acic segme s a e wo n h ph r Targioni Tozzetti, 1867: 26; – si es o o sa se ae i e i y is ge us w i e e ese ce Maskell, 1885: 29 [description; pest status]; –Maskell, o e a u u a uc s wi a e y oa i e uc u e 1887: 80 [description; records]; –Maskell, 1895a: 15 se a a es i om . rn . [checklist]; –Maskell, 1895b: 59 [Australian records]; – Cosmopolitan, widespread in the Palaearctic and Cockerell & Parrott, 1899: 164 [synonymy]; –Hutton, Nearctic, but also known from the Neotropics, Oriental, 1904: 353 [checklist; synonymies]; –Thomson, 1922: 335 Australasian regions, and New Zealand. It has a fairly [record; host]. wide host range (Ben-Dov (1993) lists 21 plant families). Ch r f l Boisduval, 1867: 328. It is a minor pest in deciduous orchards. S t f l (Boisduval); –Fernald, 1903: 201 [world . p r was first reported from New Zealand by catalogue]; –Wise, 1977: 106 [checklist]. Maskell (1892) — as n r r Snellen von Ch r h b rn l r Boisduval, 1867: 328; – Vollenhoven — but see comments under . rn above Maskell, 1879: 207 [brief comment]; –Hutton, 1904: 353 regarding the identity of early records. [checklist]. n f l nd 4 9 n h b rn l r (Boisduval); –Maskell, 1887: Spiracular disc-pores in quite broad bands between 81 [description; records]; –Maskell, 1895a: 15 [checklist]. margin and each spiracle, with 40-86 disc-pores in each S t h ph r (Targioni Tozzetti); –Kuwana, band. Ventral microducts present throughout venter. 1902: 63; –Fernald, 1903: 202 [world catalogue]; – Ventral tubular ducts of 3 types present: (i) large ducts Thomson, 1922: 335 [record, host]; –Myers, 1922: 200 with wide outer and inner ductules and a large terminal [checklist]; –Miller, 1935: 36 [status; biology]. gland: present in a distinct submarginal band but absent n (S t h ph r (Targioni Tozzetti); from wide gaps at each band of spiracular disc-pores; also –Green, 1929: 376 [record]. with l-5 present just mesad to each pro- and mesocoxa; S t ff (Walker); –Kirkaldy, 1902: 105; – (ii) small ducts with a short outer ductule and a short Cottier, 1939a: 146 [description; status; hosts]; –Cottier, filamentous inner ductule: present between submarginal 1956: 272, 325 [key; description, status, biology]; –Wise, band of type (i) ducts and margin; and (iii) ducts with a 1977: 106 [checklist]; –Penman, 1984: 63 [biology]; – fairly narrow outer ductule, a much thinner inner ductule Scott, 1984: 28 [pest status]. and a large terminal gland: present medially and mediolaterally on all segments, though sparse on thorax; U mou e ma e ia older adults convex and rounded; particularly abundant just laterad to band of type -(i) ducts. colour shiny tan to light brown with many evenly spaced Antennae generally 8-segmented. Legs well developed, light dots over most of dorsal surface. Nymphs and young with a tibio-tarsal articulation and generally with a small adult females much flatter and light yellow to pink with articulatory sclerosis; claws without a denticle; claw some darker mottling. Characteristic `H' pattern digitules both broad and slightly shorter than tarsal noticeable on nymphs and young adults, but disappears as digitules. adults become convex, so the dorsal surface is completely smooth on mature individuals. Adult females 2-4 mm Ma e ia e ami e I On indigenous native plants: long. Eggs pink. A pl n b lb f r , A pl n p p r t , A pl n sp., Av nn r n subsp. tr l , body oval, lacking obvious stigmatic Mou e ma e ia l hn fr r , C pr ?pr p n , C rd l n clefts; length 1.4-2.0 mm, width 1.2-1.8 mm. tr l , G n t sp., b r rp , M p r o sum derm membranous when young, becoming l t , r h b l ll , t r t t , r t heavily and uniformly sclerotised when mature, with small ph l d , t r d l nt . oval areolations, each with a microductule. Dorsal setae II. On exotic plants: Alt rn nth r ph l x r d , s i ose a qui e s o 3-7 μm o g eque oug ou `Asparagus fern', C tr spp., C rn lt rn f l , o sum o sa o es o y es i c u i g a mi u e C d n bl n , ndr b n bl , t j p n , mic o uc u e ese i eac a eo a io eo e cu a `Ferns', G rd n sp., l l d , l sp., Picea o es a e sma a ia e i si e eac co e wi a p n n cv. Glauca, r n p r νa n p r , u y oi e a e i a we -s ace g ou o - o es r n l n , S ntp l sp., h nb r sp., t a e io o a a a es o sa u u a uc s a se o sa v n f r , W h n t n r b t . u e c es eac sim e a co e wi 1- ai s i a s a se su ma gi a a A oge i a o wi 3 o se ae AK WO G K WI / M C O ese a o g a e io ma gi a 3 o ai s a e a y A a i g wi se ae ema ks Recent descriptions include: Kawai (1980), Ma gi marginal setae of 2 sizes, both with slightly Hamon & Williams (1984), Gill (1988), Tremblay (1988), expanded and more or less fimbriate apices although this Williams & Watson (1990), and Hodgson (1994a). The may be hard to see in some views; with 12-18 setae on presence of a discal seta on each anal plate, pregenital disc- each side between stigmatic areas, of which 5-8 are short; pores medially across all abdominal segments and on the e g o eac se a 1 -7 μm; wi mo e a 3 se ae metathorax, and a submarginal band of tubular ducts a ou e ea e wee e a e io s igma ic c e s identify this as belonging to the genus S t , whilst S. S igma ic c e s eac wi 3 s i es; me ia s i e s ig y ff can be separated from S. l by the presence of e wi a u a e eac a ou 5 o ge a a e a ventral tubular ducts with a broad inner ductule and by s i es having more than 30 marginal setae around the head e e pregenital disc-pores with mainly 10 loculi: between the anterior stigmatic clefts. abundant around genital opening and across all preceding S. ff is tropicopolitan and is also an important abdominal segments; a few occasionally present medially pest under glass. It is highly polyphagous — Ben-Dov on metathorax and with small groups laterad to each coxa. (1993) lists 82 plant families. It is an important pest of 1 d n & nd r n (2000 : C d (In t : pt r : C d ornamentals. Penman (1984) states that, unlike black scale S t l (Bernard); –Cockerell, 1901: 31; – (S t l , honeydew and the accompanyng black Fernald, 1903: 205 [world catalogue]; –Myers, 1922: 200 sooty mould are not characteristic. Cottier (1939a, 1956) [checklist]; –Thomson, 1922: 335 [mention]; –Miller, lists the following as hosts in New Zealand: asparagus, 1935: 35 [description; status; biology]; –Cottier, 1939a: camellia, citrus, cucumber (C t v , currant 145 [description; status; hosts]; –Cottier, 1956: 272, 324 ( b sp.), eggplant (S l n l n n , japonica [key; description; status; biology]; –Wise, 1977: 106 (Ch n l sp.), oleander ( r l nd r , orchids, [checklist]. and palms. Scott (1984) mentions passionfruit ( fl r S t l (Olivier); –De Lotto, 1971b: 149 d l and blackcurrant ( b n r as being [authorship correction];–Penman, 1984:61 [biology; pest commonly attacked. status]; –Ben-Dov, 1993: 313 [world catalogue]. S. ff was first reported from New Zealand by Unmounted material: adults hemispherical, young adults Maskell in 1879—as . h b rn l r — and then as . round and fairly flat, immatures oval. Young adults and h ph r in 1885. nymphs yellow or grey, rough or granular in appearance, older females dark brown or black; all stages with "H" Biology. Probably entirely parthenogenetic as no males pattern on dorsum. Adult females 2-5 mm long. are known. It has up to 8 overlapping generations a year in the tropics but fewer under cooler conditions. There is Mounted material: body round to oval, widest in more than 1 generation a year in New Zealand (Cottier, abdomen, with shallow stigmatic indentations; up to 4.5 1956). Often effectively controlled by parasitoids. It mm long and 4.0 mm wide. occurs along the main veins of the leaves and on the stems (Miller, 1935). Overwinters as immature stages but is Dorsum: derm fairly thick, with numerous pale mature by the spring when it migrates to the leaves or areolations present throughout; unsclerotised on young young twigs to reproduce (Cottier, 1939a; Penman, 1984). individuals apart from a dense anal sclerotisation anterior to anal cleft; derm becomes much thicker and heavily Distribution and status in New Zealand. Cottier (1956) sclerotised throughout on old individuals. Dorsal setae stated that S. ff could be a major pest of citrus and s o g y s i ose; e g o eac se a -13 μm; a e passionfruit; currently it is a pest on ornamentals sparse throughout. Dorsal pores of 2 types including especially ferns and potted plants, and is occasionally heavily sclerotised microductules in each areolation. found in native forest margins. Recorded from the north of Preopercular pores quite large and heavily sclerotised, the North Island to the Otago Lakes in the South Island. probably roundly convex: in a broad group of 3-46 anterior to anal plates. Dorsal tubular ducts absent. Dorsal tubercles each rather small and convex, total 4-16 in a submarginal ring. Pocket-like sclerotisations absent. Anal S t l (O i ie ack sca e o o i e sca e a es wi a a ge isca se a a ou 17-1 μm o g Anogenital fold with 3 or 4 pairs of fairly long setae Figs C90, C91, 151 present along anterior margin and 2 or 3 pairs laterally. C l Olivier, 1791: 95. Anal ring with 8 setae. n l (Bernard); –Walker, 1852: 1070; – Margin: marginal setae quite large, spinose, with Maskell, 1885: 28 [pest status]; –Maskell, 1887: 82 rather parallel sides and either tapering, blunt, or with a [description; records]; –Maskell, 1895a: 16 [checklist]; – slightly frayed apex; tending to be of 2 sizes, smaller setae Hutton, 1904: 353 [checklist]. a ou 1 - μm o g; o ge se ae mo e ume ous a n n Maskell, 1891: 15 [description; 35- μm o g; wi 5-1 se ae o ei e si e e wee e status]; –Maskell, 1895a: 15 [checklist]; –Hutton, 1904: anterior and posterior stigmatic clefts, of which 0-3 may 227 [checklist]; –Borschenius, 1957: 336 [synonymy]. be short; with 15-30 setae around the head between the S t n (Maskell); –Fernald, 1903: 200 [world anterior stigmatic clefts. Median stigmatic spine about 4 x catalogue]; –Myers, 1922: 200 [checklist]; –Wise, 1977: length of lateral spines, slightly curved, tapering to a blunt 106 [checklist]; –Ben-Dov, 1993: 303 [synonymy point; lateral spines each tapering to a fairly sharp point. uncertain]. Venter: pregenital disc-pores with mainly 10 loculi: n (S t n (Maskell); –Cockerell & present in a dense group around genital opening and less Parrott, 1899: 163. frequently across preceding abdominal segments and n (S t l (Bernard); –Green, 1929: 376 metathorax. Ventral microducts present throughout. [record]. Ventral tubular ducts of 2 types: (i) larger duct with a fairly n f l nd 4 11 broad outer ductule, thin inner ductule and moderate-sized tubular ducts with a broad inner ductule, and by having terminal gland: present in a fairly broad submarginal band; less than 30 marginal setae around the head between the and (ii) a slightly smaller duct with a filamentous inner anterior stigmatic clefts. ductule and no terminal gland: only present mediolaterally A study of Maskell's material of n n on last 2 or 3 abdominal segments. Antennae each 7- or 8- Maskell shows that it is identical with S. l and a segmented. Legs well developed; each with separate tibia lectotype is here designated; we therefore uphold the and tarsus but articulatory sclerosis usually absent; claws earlier synonymy by Borschenius (1957). S. l was without a denticle; claw digitules both broad and slightly first reported from New Zealand by Maskell in 1885 — as shorter than fine tarsal digitules. n l . In the past, other species of S t , such as S. Ma e ia e ami e r nd (Cockerell & Parrott) and S. n l t De Lotto, I n n Maskell. LECTOTYPE [ ]: here des- have been misidentified as S. l and thus its true ignated; labelled " n n , adult female, Mar distribution is hard to determine. Nonetheless, it appears 1890, W.M.M."; NZAC: 1/1[ ]ad. Host given by Maskell to be cosmopolitan and has a wide host range-50 plant (1890) as "C n l pt ph ll " [= Oz th n families are listed in Ben-Dov (1993). It is a major pest on l pt ph ll ], and localities as "Wellington, Waiararapa, some crops such as citrus and olives under subtropical and [and] Hawke's Bay." temperate conditions. Cottier (1939a; 1956) lists the PARALECTOTYPE: " n n , female 2nd- following as hosts in New Zealand: apple (M l x stage, Mar 1890, W.M.M."; NZAC: 1/1[ ]nymph (un- d t , apricot ( r n r n , camellia, citrus, cleared, unstained). daphne, grapevine ( t v n f r , guava ( d sp.), Other material: " n n , adult female, 1891, holly (Il x sp.), laurel, oleander ( r l nd r , pear W.M.M."; NZAC: 1/1 [ ] ad. ( r n , pepper tree (S h n sp.), palms, plums ( r n spp.), rose, r p n , and II O i ige ous native a s Av nn r n su s wisteria; he also states that "it is said to infect various tr l , r h l tt r p nd , C pr repens, native trees". C pr s h ndr r p n , Ent l rb r n , Several predators and parasitoids have been b ll pt , b s M p r l t , M r n introduced into New Zealand as potential biological tr l , t l n l t , Ol r p n l t , control agents against S. l (Morales, 1989). The Ol r s Oz th n l pt ph ll , Oz th n s coccinellid predators l [as Or ] h l b (Bois- tt p r s l nth d v r t , l nth duval) and h z b f r t r (Mulsant) are now well s t r t t , d p n x s t r d established although they appear to be relatively l nt , S l n v l r , t x l n . ineffective. Of the several species of hymenopterous parasitoids that have been introduced, the encyrtid III O e o ic a s Ab t l n s A t n d d l , M t ph l n b r (Howard) has become established, Ar l s A p r , Ch t rn t , C tr s although a later introduction of three other species (in C rb t p p , C s C b d s E n n parasitised S. l and S. ff was unsuccessful. fl r , b s dr n s n r p t r n , Other parasitoid species recorded from S. l in New v t r s t h h n n , Ol r p , r Zealand are C ph hr Howard n , r p r f l , s r n s (Aphelinidae), which was introduced for the control of t v n f r . mealybugs, and E x nth ll ph l pp Silvestri ΚΕ Τ / AK C WO G K I WI (Aphelinidae) and M r n l l f rn (Howard) W / M MC SC O (Pteromalidae). ema ks Recent descriptions include: De Lotto (1965, io ogy Probably almost entirely parthenogenetic as 1971c), Kawai (1980), Hamon & Williams (1984), males very rarely reported (however, empty male tests Tremblay (1988), Gill (1988), Williams & Watson (1990), seen in Auckland on one occasion in 1998). Usually has 1 and Hodgson (1994a — as Bernardia l . The or 2 generations a year. Crawlers can settle almost presence of a discal seta on each anal plate, pregenital disc- anywhere but, on deciduous trees, late-instar nymphs pores medially across all abdominal segments and on the emigrate back onto the twigs to become adult. metathorax, and a submarginal band of tubular ducts In New Zealand, S. l overwinters as eggs and identify this as belonging to the genus S t , whilst it nymphs (Miller 1935); the adults take about 3 months to can be separated from S. ff by the absence of ventral develop and there is only one generation a year. According 2 2 d n & nd r n (2000 : C d (In t : pt r : C d to Penman (1984) the females lay large numbers of eggs in Key o a u ema e lv n r s ecies i ew ea a November – December; these hatch mainly from Dorsal tubercles absent 2 December – January and the crawlers settle on leaves. After feeding for 4-6 weeks, the young scales generally —Dorsal tubercles present 3 migrate to woody parts of the host plant and become permanently attached. When common, S. l is Marginal setae spinose, with a rather blunt apex; anal associated with much sooty mould (Cottier, 1939a) and plates without setae in discal position; pregenital disc- this can cause major damage (Penman, 1984). pores with mainly 10 outer loculi mesembryanthemi is i u io a s a us i ew ea a Cottier (1956) —Marginal setae finely spinose with a sharp apex; anal stated that S. l was found wherever citrus was grown; plates each with a seta in discal position; pregenital it is currently one of the more important pests of citrus, disc-pores with mainly 7 or 8 outer loculi. particularly in the Gisborne area. Its range extends from hydrangeae the Kermadec Islands and Three Kings Islands, throughout the North Island and most of the South Island. 3 Marginal setae slightly fimbriate; pregenital disc-pores with mainly 7 outer loculi fl f r —Marginal setae finely pointed; pregenital disc-pores with mainly 10 loculi v t Ge us U I A IA a gio i o e i lv n r Targioni Tozzetti, 1866: 146; Targioni Tozzetti, 1867: 13. Type species: C v t Linnaeus, 1758. i e u i a ii i a gio i o e i Pulvinati Targioni Tozzetti, 1868: 727. Pulvinariini Targioni Tozzetti; Ashmead, 1891: 98. lv n r fl f r (Wes woo co o y came ia lv n r Targioni Tozzetti, 1867: 13. y e ge us sca e iag osis in New Zealand, members of the Pulvinariini are typically characterised by: (i) production of a woolly Figs C83, 152 ovisac by the reproducing female, which protrudes from C fl f r Westwood, 1870: 308. beneath the posterior end of abdomen, often lifting the lv n r l l Signoret, 1873a: 32; –Maskell, insect so that it appears to be standing on its head; and the 1879: 207 [description]; –Maskell, 1887: 83 [description; presence of: (ii) small dorsal tubular ducts; (iii) a tibio- records]; –Maskell, 1895a: 17 [checklist]; –Myers, 1922: tarsal articulatory sclerosis; and (iv) ventral tubular ducts 199 [checklist]; –Wise, 1977: 106 [checklist]. of 3 or 4 types, including (a) a large duct with inner ductule Chl r p lv n r fl f r (Westwood); –Borchsenius, of similar length and width, with a large terminal gland, (b) 1952: 300. a similar duct (intermediate duct) but with a narrow inner lv n r fl f r (Westwood); –Green, 1897: 72; – ductule and large terminal gland, and (c) a short duct with Thomson. 1922: 335 [mention]; –Ben-Dov, 1993:261 a filamentous inner ductule and no glandular end; and (v) [world catalogue]. the absence of pocket-like sclerotisations. U mou e ma e ia body elongate oval, slightly ema ks Four species of lv n r have been recorded convex, usually widest near centre; cream to tan-coloured, from New Zealand and are minor pests: . fl f r mottled with brown and usually with a brown border (Westwood), . h dr n Steinweden, . around body. Adult females on stems and leaves of host, br nth (Vallot), and P. v t ( The record of secreting an elongate, very white, and fluffy ovisac; at . p d in Fernald (1903) is erroneous; this species has maturity; spent female falls off, leaving only the ovisac never been recorded from New Zealand. attached to plant. n f l nd 4 2 ig. 2. lvln r fl f r (Wes woo , a u ema e — (A a e om o gso , 4a . 2 4 d n & nd r n (2000 : C d (In t : pt r : C d . . lv n r h dr n S ei we e , a u ema e — ( om Gi , 88 . n f l nd 4 2 . 4. lv n r br nth ( a o , a u ema e (A a e om o gso , 4a . 2 6 d n & nd r n (2000 : C d (In t : pt r : C d ig. . lv n r v t ( i aeus , a u ema e (A a e om o gso , 4a . n f l nd 4 2 Mou e ma e ia body elongate oval, widest in ema ks Recent descriptions (often as Chl r p lv n r abdomen. Length 1.5-4.0 mm, width 1.0-2.5 mm. fl f r (Westwood)) include: Williams & Kosztarab (1972), Kawai (1980), Hamon & Williams (1984), Gill derm membranous, with pale areolations on o sum (1988), Kosztarab & Kozár (1988), Qin & Gullan (1992), older females. Dorsal setae rather short and spiniform, 4- and Hodgson (1994a). This species has dorsal tubercles, μm o g eque o e e i e o sum o sa o es o mildly fimbriate marginal setae, pregenital disc-pores with y es i c u i g a sma mic o uc u e oca e mai y i mainly seven loculi, and no ventral tubular ducts a eo a io s eo e cu a o es a e sma a a submarginally anterior to the antennae. a e y sc e o ise i a sma g ou o 7- 5 i o o a a . fl f r has an almost world-wide distribution a es o sa u u a uc s eque oug ou o sa and is particularly widespread in the Holarctic Region. In u e c es a e sma wi o a o -1 su ma gi a y the Australasian and Pacific regions, it is only known from ( a e y a se A a a es oge e qua a e A oge i a Australia (Qin & Gullan, 1992) and New Zealand, and o wi 1 o ai s o se ae o a e io ma gi a 1 o only from Vietnam in the Orient (Ben-Dov, 1993). Ben- ai s a e a y A a i g wi 3 ai s o a ge se ae a Dov (1993) lists 27 plant families as hosts. - ai s o sma e se ae . fl f r was first reported from New Zealand by Ma gi marginal setae rather long, slightly shorter Maskell (1879) — as lv n r ll l , a than median stigmatic spine, typically with a slightly misspelling of . l l Signoret. spatulate apex, but may be pointed, frayed, or divided; Often placed in the genus Chl r p lv n r Borch- each seta generally slightly curved, with well-developed senius, 1952. basal sockets; with 15-27 setae on either side between s igma ic c e s; e g 3-93 μm (usua y a ou 5 μm io ogy Has a single generation in USA (Williams & S igma ic c e s wi 3 a e s ou s igma ic s i es eac Kosztarab, 1972) and Japan, although 2 generations are a e i g o a u a e ; me ia s i e o 3 e g o known off E r in Tokyo (Takahashi, 1955). See El- a e a s i es; eac me ia s i e 55-7 μm a e a s 17- Minshawy & Moursi (1976) for some details of its μm biology. e e pregenital disc-pores each with mainly 7 loculi: present around genital opening and on mediolateral is i u io a s a us i ew ea a Currently only areas of preceding 3 to 4 segments; small groups also known from tea (C ll n n and camellias in New present laterad to each meta- and (occasionally) Zealand. mesocoxa. Spiracular disc-pore bands narrow, with 30- 71 pores in each band. Ventral microducts present in a broad submarginal band and sparsely throughout elsewhere. Ventral tubular ducts: (i) large ducts: present lv n r h dr n co o yS ei we e medially on head, thorax, and first 1 or 2 abdominal y a gea sca e segments, extending laterally to spiracles; (ii) intermediate ducts: present medially on posterior abdominal segments; Figs C84, 153 and (iii) short ducts: present in a broad submarginal band, lv n r h dr n Steinweden, 1946: 7; –Ben-Dov, abundant posteriorly on abdomen, becoming rather thinly 1993: 265 [world catalogue]. distributed between bands of spiracular disc-pore, very yellowish, mottled with brown, scarce anterior to prothorax and absent anterior to U mou e ma e ia rather flat, developing transverse ridges with age; ovisac antennae. Antennae each with 8 segments, 2nd and 3rd up to about 10 mm long, broad, white, and ribbed. Length segments subequal. Legs well developed; each with a 2.00-5.00 mm, width 1.5-2.0 mm. tibio-tarsal articulation and an articulatory sclerosis; claws without a denticle; both claw digitules broad and slightly ovoid. Up to 4.00 mm long and 3.5 shorter than thin tarsal digitules. Mou e ma e ia mm wide. Ma e ia e ami e I Not recorded on indigenous o sum derm membranous. Dorsal setae short and native plants. tapering to a sharp point; present throughout. Dorsal pores: perhaps only minute microductules: frequent II On exotic plants: C ll spp. throughout. Preopercular pores small, with a granulate surface: in an elongate group anterior to anal plates. BP, TK / NN. Dorsal tubular ducts small; frequent throughout. Dorsal 1 d n & nd r n (2000 : C d (In t : pt r : C d tubercles absent. Anal plates together quadrate, with 3 lists 7 plant families; of these, the Hydrangeaceae are the fine setae near apex and another in the discal position. main hosts (Gill, 1988). As it appears to be able to survive Anogenital fold with 2-3 pairs of setae along anterior quite cold winters, it could occur almost anywhere in New margin and 3 pairs on lateral margin. Anal ring with 4 Zealand. pairs of setae, I pair rather short. Ma gi marginal setae slender, mostly with an acute io ogy Has 1 generation a year in California (Gill, apex but some fimbriate, slightly curved; rather sparse, 1988), where it overwinters as the 3rd-instar female on the with about 10-20 setae on each side between stigmatic twigs, matures in the spring, and lays eggs. The crawlers clefts. Stigmatic clefts shallow, with 3 stigmatic setae; then return to the leaves for initial development. In Europe each median spine 1.5-3.0 x longer than lateral spines, it is parthenogenetic (Canard, 1965) and the female curved. migrates back to the leaves to produce its ovisac. e e pregenital disc-pores each with mainly 7 loculi, central loculus slit-like: present in a dense group is i u io a s a us i ew ea a Of no around genital opening, becoming much less frequent economic importance. Records from between central across anterior abdominal segments, with a group laterad North Island and the northern end of the South Island. to each metacoxa and sometimes each mesocoxa. Spiracular disc-pores in a band extending medially and merging with groups of pregenital disc-pores laterad to each coxa; with 55-70 disc-pores in each band. Ventral lv n r br nth ( a o ice a microducts, abundant in a broad submarginal band; sca e apparently absent elsewhere. Ventral tubular ducts: (i) Figs C85, 154 large ducts: abundant medially and submedially on head and thorax (sometimes abdominal segments II & III); (ii) Coccus mesembryanthemi Vallot, 1829: 30. intermediate ducts: present medially and mediolaterally on Pulvinariella mesembryanthemi (Vallot); –Borchsenius, abdomen and laterad to type (i on head and thorax; and 1953: 287. (iii) short ducts: present in a complete, broad, submarginal Pulvinaria mesembryanthemi (Vallot); –Ben-Dov, 1993: band. Antennae 8-segmented. Legs well developed; each 270 [world catalogue]. with a tibio-tarsal articulation and an articulatory U mou e ma e ia adult females oval to circular, sclerosis; claws without a denticle; both claw digitules moderately convex; bright green in colour; ovisac convex broad and slightly shorter than thin tarsal digitules. and white, almost as long as body; females also secrete a sparse dorsal covering of white mealy wax. Ma e ia e ami e I Not recorded on indigenous native plants. Mou e ma e ia body oval to almost round, widest in abdomen, rather flat; stigmatic clefts shallow; a shallow II O exotic plants: Hydrangea sp., Prunus avium, cleft often present near each eyespot. Length 1.6-2.3 mm, Prunus serrulata. width 0.9-1.6 mm. WO G W / MC o sum derm membranous. Dorsal setae small (5-8 ema ks Recent descriptions (sometimes as Eupulvinaria μm o g s i ose wi a u a e eque oug ou hydrangeae (Steinweden)) include: Canard (1965), Dorsal pores of 2 types: a minute microductule and a larger Williams & Kosztarab (1972), Pellizzari Scaltriti (1976), simple pore: fairly frequent throughout. Preopercular Kawai (1980), Hamon & Williams (1984), Gill (1988), pores rather small, round to slightly oval, flat to slightly and Qin & Gullan (1992). As mounted specimens, it can convex, with a granular surface: in a small narrow group of easily be identified by the discal seta on each anal plate and 11-51 pores anterior to anal plates. Dorsal tubular ducts the absence of dorsal tubercles. small; frequent throughout. Dorsal tubercles absent. Anal First recorded in New Zealand in 1977 (Archibald et plates together quadrate. Anogenital fold with 2 pairs of al. 1979). Within the last 20 years there have been a few setae present along anterior margin and 2 or 3 pairs records of this species in New Zealand— it remains scarce laterally. Anal ring with 3 pairs of large and 0-1 pair of and of no economic importance. The only other records smaller setae present. from the Southern Hemisphere are from New South Wales Ma gi marginal setae stoutly spinose, slightly (Qin & Gullan, 1992), otherwise it is only known from the curved, blunt apically, and with rather parallel sides; rather Nearctic and Palaearctic regions. Ben-Dov (1993) only sparse, in more or less 2 lines, more ventral line with n f l nd 4 2 sma e se ae; a ge se ae -3 μm o g s o e se ae as occasionally Chenopodiaceae (Ben-Dov, 1993)). Its life s o as 1 μm; wi 5-1 a ge a 1-3 sma e se ae o cycle has been studied by Washburn & Frankie (1981, eac si e e wee s igma ic c e s S igma ic c e s wi 3 1985) and Washburn t l. (1985); it usually has 2 s igma ic se ae; eac me ia s i e 37- 5 μm o g generations a year in northern California but multiple s ig y cu e occasio a y s ig y swo e a ica y; generations in the south (Gill, 1988). a e a s i es ge e a y s o eac 1 - μm o g e e pregenital disc-pores with mainly 10 loculi: is i u io a s a us i ew ea a Recorded from present in a dense group around genital opening, Great Island (Three Kings Islands), the Auckland region becoming much less frequent across anterior abdominal and from Thames, Napier, and Christchurch. segments and metathorax; a group also present laterad to each coxa. Spiracular disc-pores in a band extending medially and merging with groups of pregenital disc-pores lv n r p d Maske g ee s ie sca e laterad to each coxa; with 32-65 disc-pores in each band. Ventral microducts, abundant near margin and labium but lv n r p d Maskell, 1893a: 223; -Fernald, 1903: less frequent elsewhere. Simple pores in a marginal band 137 [world catalogue]; -Wise, 1977: 106 [checklist]; - among marginal setae, with 3-5 pores on each side Ben-Dov, 1993: 278 [world catalogue]. between stigmatic clefts. Ventral tubular ducts: (i) large . p d has never been recorded from New Zealand. ducts: abundant medially and submedially on head, Maskell described it in 1893 in the r n t n nd thorax, and 2nd abdominal segment; (ii) intermediate r d n f th l nd In t t t from Hawaii ducts: abundant submedially throughout abdomen, (the Sandwich Is). Without apparently sighting the paper, merging with type-(i) anteriorly; occasionally present Fernald (1903) included this reference in her World medially on abdominal segments III- I; and (iii) short Catalogue as being recorded in New Zealand and this error ducts: present in a complete, broad, submarginal band, and was repeated by Wise (1977); Ben-Dov (1993) gives the also submedially on abdomen and medially on posterior correct distribution status for . p d in his World abdominal segments. Antennae 8-segmented. Legs rather Catalogue. large; each with a distinct tibio-tarsal articulation and articulatory sclerosis; each claw possibly with a minute denticle; claw digitules both broad and subequal in length with tarsal digitules. lv n r v t ( i aeus co o y i e sca e Ma e ia e ami e I On native (non-indigenous) Figs C86, C87, 155 plants: ph tr l . C v t Linnaeus, 1758: 456. II On exotic plants: C rp br t sp., pr nth sp., lv n r v t (L.); -Signoret, l873a: 45; -Ben-Dov, M br nth r t ll n , M br nth 1993: 288 [world catalogue]. sp. U mou e ma e ia live adult female highly variable, ΤΗ /AK C ΠΒ / MC depending on age, host-plant species, and position on host; teneral female pale beige, reddish-brown, dark- ema ks Recent descriptions (often as lv n r ll brown or blotched dark-grey, occasionally with raised br nth (Vallot)) include: De Lotto (1967), pale-yellow longitudinal mid-dorsal band; body becoming Hodgson (1967a, 1968, 1994a), Gill (1988), and Qin & greatly convex, heavily sclerotised, uniform dark-brown Gullan (1992). Can be identified by lack of dorsal and wrinkled at maturity; ovisac large, white, strongly tubercles, bluntly spinose marginal setae, pregenital disc- convex, up to 10.0 mm long. Eggs pink. pores with mainly 10 loculi and ventral tubular ducts present anterior to antennae. Mou e ma e ia body elongate oval, widest in . br nth is a potential pest of Aizoaceae abdomen, with shallow stigmatic areas. Length 1.5-8.5 wherever they are grown. Considered to have originated mm and width 0.9-6.9 mm. from South Africa. It has been known in New Zealand since 1987. o sum derm membranous on young specimens, becoming heavily sclerotised when mature, with sparse io ogy . br nth has a very restricted host- pale areolations. Dorsal setae finely spinose with a sharp range, being known only from Aizoaceae (and oi some imes s ig y cu e eac -15 μm o g; 220 d n & nd r n (2000 : C d (In t : pt r : C d frequency variable. Dorsal pores of 2 types: a minute dorsal tubercles, pregenital disc-pores with 10 loculi, microductule in areolations and a slightly larger simple ventral tubular ducts absent from anteriorly on head pore. Preopercular pores round, flat to slightly convex, between antennae, marginal setae finely spinose, and a with a granulate surface, rather variable in size: in an denticle on the claw. elongate group of 7-170 pores. Dorsal tubular ducts small . v t is a highly polymorphic species and the many and sparsely distributed on some specimens, rather forms of it have been studied in Europe by Malumphy frequent on others. Dorsal tubercles in a submarginal ring (1991) and Lagowska (1996), who considered that many totalling 0-14. Anogenital fold with 2 pairs of long setae of the lv n r species from Europe were synonyms of present along anterior margin and 3 or 4 pairs laterally. . v t . Based on their interpretation of . v t , this Anal ring with 6 large and 2 shorter setae. species has a fairly narrow host range of only about 14 Margin: marginal setae spinose, fine, often curved, plant families (Ben-Dov, 1993). eac 1 μm o g is i u e i mo e o ess ows It appears to have arrived in New Zealand only fairly with 8-19 setae on each side between stigmatic clefts. recently. Elsewhere, it is mainly found in the Palaearctic Stigmatic clefts shallow, each with 3 stigmatic spines; Region, where it is an important pest of grapevine (in median spine longest, often slightly curved, bluntly Europe); also present in the Nearctic Region, where it is pointed; length of median spine about twice length of important on peaches (in Canada). In the Southern lateral spines. Hemisphere, only known from Brazil and New Zealand Venter: pregenital disc-pores with mainly 10 loculi: (Ben-Dov, 1993). In New Zealand, sporadic outbreaks abundant around genital opening, becoming progressively have occured on apricots in Central Otago orchards, with less frequent across preceding abdominal segments and records clustered around 1951-52, 1984-85, and (at metathorax; also present laterad to each meso- and present) in 1998-1999. Occasionally also on grapevines, metacoxa. Spiracular disc-pores in broad bands, with to which it can transmit grapevine leafroll closterovirus about 60-110 disc-pores in each band. Ventral disease (Belli t l., 1994). microducts abundant near margin, less frequent elsewhere. Ventral tubular ducts abundant throughout Biology. A univoltine species throughout most of its except medially on head and anterior to antennae; 3 types range. Many details regarding the biology of this species present: (i) large ducts: only duct present medially on (in Poland) can be found in Lagowska (1996). head, thorax, and more anterior abdominal segments; extending laterally to band of type (ii) and (iii) ducts, Distribution and status in New Zealand. Of little where they become infrequent or absent; (ii) intermediate importance economically; verified records from the South ducts: frequent medially on more posterior abdominal Island only — range possibly more extensive. segments and laterally interspersed with type (i) ducts; and (iii) short ducts: frequent in a broad submarginal band extending from anal cleft to near antennae. Antennae 8- segmented. Spiracles quite large. Legs well developed; each with a distinct tibio-tarsal articulation and articulatory sclerosis; claws with a small denticle; claw digitules both broad and marginally shorter than tarsal digitules. Material examined. I. Not recorded on indigenous native plants. II. On exotic plants: p l ?n r hybrid, Prunus r n , Prunus p r , Prunus sp., r n , b n r , t v n f r . MC SC O CO S Remarks. Recent descriptions include: Danzig (1986), Kosztarab & Kozár (1988), Gill (1988), Hodgson (1994a), and Lagowska (1996). This species can be identified by the following combination of characters: presence of n f l nd 4 22 E E E CES 1993: A Systematic Catalogue of the Soft Scale In- sects of the World (Homoptera: Coccoidea: Coccidae). l r nd n ndb n . . Gainsville, Sandhill Archangelskaya, A.D. 1937: The Coccidae of Middle Asia. Crane Press. 536 pp. Izdatelstvo Komiteta Nauk UZSSR, Tashkent. 158 Birjandi, A.K. 1981: Biology and ecology of pp. rth n l n species. Ent l t M nthl Archibald, R.D.; Cox, J.M.; Deitz, L.L. 1979: New records M z n : 1400-1403. of plant pests in New Zealand. l nd rn l Blank, R.H.; Olson, M.H.; Gill, G.S.C.; Dow, B.W. 1997: f A r lt r l r h 22: 201-207. Timing of insecticide applications for control of soft Ashmead, W.H. 1891: A generic synopsis of the Coccidae. wax scale (Homoptera: Coccidae) on citrus. New Zea- Family X — Coccidae. r n t n f th A r n land rn l f Cr p nd rt lt r l S n 2 : Ent l l S t 8: 92-102. 311-317, Atkinson, E.T. 1886: Insect pests belonging to the Blumberg, D. 1997: Encapsulation of parasitoids. Pp. 375- homopterous family Coccidae. rn l f th A t 387 n Ben-Dov, Y; Hodgson, C.J. (eds), Soft Scale S t f n l, t r l t r 55: 267-298. Insects, Their Biology Natural Enemies and Control. Avasthi, R.K.; Shafee, S.A. 1986: Species of Ceroplastinae World Crop Pests Series, 7A. Amsterdam, Elsevier (Homoptera: Coccidae) from India. rn l f th Press. xxiv + 452 pp. b t r l t r S t 8 : 327-338. Bodenheimer, F.S. 1935: Studies on the zoogeography and Balachowsky, A. 1932: Etude biologique des coccides du ecology of Palaearctic Coccidae, I-III. E ( 4 Bassin Occidental de la Mediterranée. Paris, P. 0: 237-271. 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Chapter X, pp. 868-908, n Schlechter, F. of the New Zealand sub-region. The smaller orders. R. R.; Miethe, E. (eds), Die Orchideen. Berlin, Paul Parey. A l nd In t t t nd M ll t n . 176 pp. n f l nd 4 2 A e i A. A a e ica is i g o os a s Av nn r n su s . tr l (Wa . . i c u i g ami y, wi ei associa e so sca e E e e [sy . Av nn r n f r G. s ecies i ew ea a . a ames a au o s o s .], A ice iaceae, as i G i i s ( 4 a a so s t l. ( C r pl t n n , A ( e e e ces i c u e u o ecem e , wi S t ff , A ei wiese & Wa ( a Mi c e t l. ( . S t l , A A, a e i e, cu i a e a o a u a ise E, Av nn s ., A ice iaceae, e emic , a i e, u o e emic [ ], u is e C r pl t n n , A eco , o e i ie . l h d t (A. Cu . e . & ook . e Ab t l n s ., Ma aceae, A Ki k, au aceae, Ε C h p r d , A Aph n h t n p b n , Ε S t l , A C h p r d , A A t n d d l (A. C e . C. . ai g & A. . l h t n l rp , Ε e guso , Ac i i iaceae, A l h d t r Α.Ε. W ig , au aceae, Ε C r pl t d tr t r, A Cr t ll t t rn t , Ε C h p r d , A l h t n l rp , Ε S t l , A l h t n fl v , Ε Al tr n x l Gae ., Sa i aceae, Ε S t ff , A Ep l d h t n p p r , Ε l hn fr r (A. Cu . ue ss., au aceae, Ε Al p fl v d ( ook. . ieg em, o a aceae, Ε C h p r d , A Ct n h t n p r v r d , Ε r p z l b t , Ε Alt rn nth r ph l x r d (Ma . G ise ., n n t , Ε Ama a aceae, A S t ff , A A C h p r d , v rd s ., u iaceae, A A S t ff , C h p r d , A A t l b n A. Cu ., As e iaceae, Ε r h l tt b ll d d ( ook. . . o ., U b n h t n h n nth r ?, Ε As e aceae, Ε Ar l s ., A a iaceae, A C h p r d , A S t l , A r h l tt r p nd . . o s & G. o s ., Ar t t l fr t ook. ., E aeoca aceae, Ε As e aceae, Ε Aph n h t n n n p , Ε C h p r d , A Ar t t l rr t ( . . o s . & G. o s . W. . . Cr t ll t t f , Ε O i e , E aeoca aceae, Ε S t l , A Ep l d h t n p p r ,Ε r h l tt s ., As e aceae, Ε Ar t t l s ., E aeoca aceae, Ε l h t n fl v , Ε rth n l n rn , A oom , a aceae, A A p r s ., As a agaceae, A C h p r d , A S t ff , A r t n r , A S t l , A as a agus e [ A p r t ], C ll s ., eaceae, A As a agaceae, A lv n r fl f r , A S t ff , A S t ff , [ ], A A pl n b lb f r G. o s ., As e iaceae, Ε S t l , [ ], A S t ff , A C r h l tr l . . i i . [sy . C. A pl n p p r t ow sey & Ρ. ackso , l r G. Sim so & C. nn n h As e iaceae, Ε aou sy . Ch rd p rt t v n n S t ff , A C eesema ], a aceae, Ε A pl n s ., As e iaceae, Ε C h p r d , A S t ff , A C l n l , A A t l b n A. Cu ., As e iaceae, Ε C r h l s ., a aceae, Ε U b n h t n h n nth r ?, Ε C l n l , A 2 2 d n & nd r n (2000 : C d (In t : pt r : C d C rp br t s ., Ai oaceae, A C pr rh n d A. Cu ., u iaceae, E lv n r br nth , A C r pl t n n , A C n l pt ph ll [i c u i g C n v v ll r ] K l r d pr , E see Oz th n l pt ph ll , E C pr r b t aou , u iaceae, E Ch n l s ., osaceae, A C r pl t n n , A S t ff , [ ], A Ep l d h t n p p r , E Ch p r l n n (Mu ay a ., C pr p th l t A. Cu ., u iaceae, E Cu essaceae, A K l r p rf r t , E Ep l d h t n p p r , E C pr s ., u iaceae, E Ch n hl fl v n o o , G ami eae, E Aph n h t n n n p ,E Aph n h t n h n hl , E Aph n h t n pr n , E Ch t rn t Η K., u aceae, A C r pl t n n , A S t l , A C h p r d , A Ch rd p rt t v n n see C r h l Cr t ll t t f , E tr l Ct n h t n h l n, E C tr s ., u aceae, A Ct n h t n v r d , A C r pl t r f r , A K l r d pr , E C r pl t d tr t r, A K l r p rf r t , E C r pl t n n , A l h t n l rp , E C h p r d , A S t l , A C l n l , A C rd l n tr l (G. o s . E ., As e iaceae, E r t nί r , A S t ff , A rth n l n p r , A C rd l n b n ook. ., As e iaceae, E S t ff , A Aph n h t n p b n , E S t l , A C rn lt rn f l . ., Co aceae, A Cl t nn n h u c ., a u cu aceae, E S t ff , A Ct n h t n h l n, E C r l t n t r aou , Esca o iaceae, E Cl t s ., a u cu aceae, A Aph n h t n n n p , E S t ff , [ ], A C r s ., Esca o iaceae, E C pr t f l ook. ., u iaceae, E C h p r d , A C h p r d ,A C r n rp l v t . . o s . & G. o s ., C pr rb r Ki k, u iaceae, E Co y oca aceae, E K l r d pr , E Aph n h t n bt l , E C pr l n ook. ., u iaceae, E Ct n h t n h l n, E K l r d pr , E Ep l d h t n p p r , E K l r p rf r t , E C t v ., Cucu i aceae, A C pr f t d . . o s . & G. o s ., S t ff , [ ], A u iaceae, E C rb t x uc es e i am ., E K l r d pr , Cucu i aceae, A K l r p rf r t , E C h p r d , A C pr l d . . o s . & G. o s ., C rb t p p ., Cucu i aceae, A u iaceae, E S t l , A K l r p rf r t , E C th s ., Cya aceae, E C h p r d , A C pr r rp C eesema , u iaceae, E K l r d pr , E Ep l d h t n p p n , E C s ., A ecaceae, A C pr pr p n A. Cu ., u iaceae, E S t l , A Aph n h t n n n p , E C d n bl n M ll., osaceae, A S t ff , A S t ff , A C pr r p n ook ., u iaceae, E C b d s ., O c i aceae, A K l r p rf r t , E C h p r d , A S t l , A S t l , A n f l nd 4 2 C ph ndr b t (Ca . Se e , Ε In l p t ll , So a aceae, Α l h t n l rp , Ε C r pl t n n , [ ], Α El n t s ., My si aceae, Ε Aph n h t n bt l , Ε Ct n h t n h l n, Ε r rp d r d d (A. ic . e au ., Ent l rb r n . ., i iaceae, Ε o oca aceae, Ε S t l , Α r p z d r d , Ε E ll n s ., Esca o iaceae, Α r d pr n am ., o oca aceae, C r pl t n n , [Ρ], Α Ε E n n fl r ., My aceae, Α r p z l b t , Ε S t l , Α r p z d r d , Ε phn s ., yme aeaceae, Α t j p n ( u . ec e. & a c ., r t n r , Α A a iaceae, Α S t l , [ ], Α Α C h p r d , ndr b s ., O c i aceae, Α S t ff , Α C h p r d , A S t l , Α S t ff , Α j ll n ( e g e g, My aceae, Α nth s ., Ca yo y aceae, Α C r pl t n n , Α C h p r d , Α r t n r , Α h ndr r p n . . o s . & G. o s ., e , Ε Α Co o u aceae, C h p r d , Α S t l , Α n n t , Ε r t t aou , am aceae, Ε S t ff , Α K l r p rf r t , Ε s . [as u e a ], Mo aceae, Α ph tr l (W. . Ai o Ν.Ε. ., Ai oaceae, C h p r d , Α lv n r br nth , Α G rd n s ., u iaceae, Α d n v acq., Sa i aceae, C r pl t n n , Α Ct n h t n p r v r d , Ε C h p r d , Α r ph ll n l r Ch n, E ac i aceae, S t ff , Α Ε G lth r s ., E icaceae, Ε Aph n h t n r , Ε l h t n d d , Ε r ph ll b l t ook. ., E ac i aceae, G nl t s ., oga iaceae, Ε Ε S t ff , Α Aph n h t n r , Ε Gr ln l tt r l aou , G ise i iaceae, Ε r ph ll tr v r ook. ., E ac i aceae, Ε Ct n h t n p r v r d , Ε Aph n h t n d r , Ε K l r p rf r t , Ε r ph ll s ., E ac i aceae, Ε Gr l n l d G. o s ., G ise i iaceae, Ε Aph n h t n d r , Ε Aph n h t n p b n , Ε x l p t b l (G. o s . ook. ., Ct n h t n p r v r d , Ε Me iaceae, Ε Ep l d h t n p p r , Ε b br h ph n Summe ., Sc o u a iaceae, Ε El rp d nt t ( . . o s . & G. o s . K l r p r d pr , Ε M. a ., E aeoca aceae, Ε b ll pt (G. o s . e e , Sc o u a iaceae, l h t n fl v , Ε Ε El rp h r n aou , E aeoca aceae, l h t n fl v , Ε Ε S t l , Α l h t n l rp , Ε b r rp ( a . Cockay e & A a ,. El rp s ., E aeoca aceae, Ε Sc o u a iaceae, Ε Cr t ll t t rn t , Ε S t ff , A 2 4 d n & nd r n (2000 : C d (In t : pt r : C d b d r ( ook. . Cockay e, Sc o u a iaceae, Aqui o iaceae, A s ., Il x E A r t n r , K l r p r d pr , E S t l , [ ], A b p r (Cockay e & A a . . Moo e Ir r n ., I i aceae, A i A a , Sc o u a iaceae, E r t n r , A Aph n h t n pr n , E b tr t ( e . . . Moo e i A a , K nz r d (A. ic . oy om s., Sc o u a iaceae, E My aceae, C r pl t n n , A C r pl t n n , [ ], A b s ., Sc o u a iaceae, E Cr t ll t t l pt p r , E C r pl t n n , [ ], A Cr t ll t t rn t ll , E d r s ., A a iaceae, A l h t n p ll n , E C h p r d , A U b n h t n b ll t , E d r rb r . . o s . & G. o s ., Mo imiaceae, E Aph n h t n p b n , E n r p t r n (A . G. o , Ma aceae, A Aph n h t n bt l , E S t l , A Ct n h t n p r v r d , E pr nth s ., Ai oaceae, A Cr t ll t t rn t , E lv n r br nth , A Ep l d h t n p p r , E au e , au aceae, A E n l I p t ll , C h p r d , A l h t n l rp , E S t l , [ ], A l h t n fl v , E r l n v z l nd A. Cu ., Mo imiaceae, r p z l b t , E E n t b l , E Ct n h t n p r v r d , E U b n h t n h n nth r , E Ep l d h t n p p r , E b t ll ., Ma aceae, [ΚΕ] v t r s ., Ma aceae, A C h p r d , A S t l , A Ma aceae,s ., A b pt p r p r . . o s . & G. o s ., S t l , A My aceae, h r p p ln A. Cu ., Ma aceae, E C r pl t n n , [Ρ], A C r pl t n n , A Cr t ll t t l pt p r , E h r Ma aceae, E s ., Cr t ll t t rn t ll , E Ct n h t n p r v r d , E l h t n p ll n , E l l d s ., o ygo aceae, A U b n h t n b ll t , E S t ff , A pt p r s ., My aceae, dr n s ., y a geaceae, A Cr t ll t t l pt p r , E lv n r h dr n , A Cr t ll t t rn t ll , E S t l , A l h t n fl v , E l h t n p ll n , E n nth r s ., io aceae, E p n f l t (G. o s . A. ic ., Cr t ll t t f , E E ac i aceae, E U b n h t n h n nth r , E Cr t ll t t rn t ll , E p r s ., y e icaceae, A t h h n n So ., Sa i aceae, A C l n l , A S t l , A t l r (A. Cu . e . & ook . e Ki k, Il t l s ., o a aceae, E au aceae, E l h t n l rp , E Aph n h t n p b n , E Il x f l ., Aqui o iaceae, A Ct n h t n h l n, E C r pl t n n , A l h t n l rp , E C h p r d , A n r s ., Ca i o iaceae, A r t n r , A C r pl t n n , A n f l nd 4 2 l s ., u iaceae, Α n h t n n r, Ε C h p r d , Α M d l f l A. Cu ., Sa a aceae, Ε S t ff , Α Aph n h t n p b n , Ε Aph n h t n bt l , Ε M r p p r x l (G. o s . Miq., i e aceae, M hl nb tr l (G. o s . Meiss ., Ε o ygo aceae, Ε Ep l d h t n p p n , Ε Aph n h t n n n p , Ε M n l t ll t (Sie o & ucc. Ma im., M p r l t G. o s ., Myo o aceae, Ε Mag o iaceae, Α C h p r d , A C l n l , Α Cr t ll t t f , Ε M l x d t o k ., osaceae, Α S t ff , A C h p r d , Α S t l , A S t l , [ ], Α M r n tr lί (A. ic . A a , My si aceae, Ε M d t v ., a aceae, Α Ct n h t n p r v r d , Ε C h p r d , Α l h t n fl v , Ε M l p pl x A. Cu ., u aceae, Ε S t l , A C r pl t n n , A M r n d v r t A. Cu ., My si aceae, Ε C l n l , A U b n h t n h n nth r , Ε Ct n h t n h l n, Ε M r n l n ewa e ook. ., My si aceae, K l r p rf r t , Ε Ε M l p t rn t . . o s . & G. o s ., u aceae, Ct n h t n p r v r d , Ε Ε l h t n fl v , Ε C l n l , A M rt s ., My aceae, Ε M l t r fl r . . o s . & G. o s ., U b n h t n h n nth r , Ε io aceae, Ε Cr t ll t t f , Ε rl l nd r ., A ocy aceae, A l h t n fl v , Ε C h p r d , [ ], A U b n h t n h n nth r , Ε S t ff , [ ], A M l t s ., io aceae, Ε S t l , [ ], A Cr t ll t t f , Ε t l n l t ( ook. . .A.S. o so , Ep l d h t n p p r , Ε O eaceae, E M r t n l r ( ook. . Seem., A a iaceae, Ε S t l , A Aph n h t n bt l , Ε th f f ( ook. . Oe s ., agaceae, Ε Ct n h t n v r d , Ε Cr t ll t t f , Ε M br nth r t ll n ., Ai oaceae, Cr t ll t t n f , Ε Α th f nz ( ook. . Oe s ., agaceae, lv n r br nth , A Ε M br nth s ., Ai oaceae, A Cr t ll t t f , Ε lv n r br nth , Α th f tr n t (Co e so Cockay e, M tr d r x l So . e Gae ., My aceae, agaceae, Ε Ε Cr t ll t t n f , Ε n h t n t t , Ε Cr t ll t t rn t , Ε n h t n t ll r , Ε M tr d r r b t A. Cu ., My aceae, Ε n h t n t t , Ε Ol r p ., O eaceae, A M tr d r b ll t Ca ., My aceae, Ε S t l , Α n h t n tr d r , Ε Ol r r d nt ake , As e aceae, Ε n h t n n r, Ε l h t n p n t t , Ε M tr d r s ., My aceae, Ε Ol r n l r f l ook. ., As e aceae, Ε Cr t ll t t rn t ll , Ε Aph n h t n n n p , Ε Ep l d h t n p p r , Ε C h p r d , Α n h t n tr d r , Ε l h t n d d , Ε 2 6 d n & nd r n (2000 : C d (In t : pt r : C d Ol r p n l t ( . . o s . & G. o s . uce, tt p r l n ook. ., i os o aceae, E As e aceae, E In l p t ll , E S t ff , A tt p r n d A. Cu ., i os o aceae, S t l , A E Ol r tr v r ( . Mue . ook. ., As e aceae, E Ep l d h t n p p r , E C h p r d , A K l r p rf r t , E Ol r s ., As e aceae, E tt p r r ook. ., i os o aceae, E C r pl t n n , [Ρ], A K l r p rf r t , E S t l , A tt p r t n f l So . e Gae ., O c i s i os o aceae, E S t ff , [ ]. A Aph n h t n p b n , E Oz th n l pt ph ll (G. o s . ei w. & K l r p rf r t , E .M. Wa [sy . C n l pt ph ll (G. tt p r t rn r e ie, i os o aceae, E o s . . . i c u i g C n K l r p rf r t , E v v ll r ( om . & acq. ook. .], U b n h t n j b t , E As e aceae, E tt p r b ll t a ks & So . e Gae ., l h t n d d , E i os o aceae, E S t l , A K l r p rf r t , E tt p r s ., i os o aceae, E a ms Aph n h t n bt l , E S t ff , [ ], A C r pl t n n , [ ], A S t l , [ ], A C h p r d , A r h b l ll ( ook. . W. . . O i ., Ep l d h t n p p r , E Sc o u a iaceae, E In l p t ll , E S t ff , A K l r p rf r t , E r h b l n (Co e so W. . . O i ., l h t n fl v , E Sc o u a iaceae, E S t l , A l h t n d d , E l nth d v r t . . o s . & G. o s , r n s ., A ocy aceae, E Ma aceae, E K l r p rf r t , E Aph n h t n n n p , E fl r d l Sims, assi o aceae, A S t l , A S t ff , [ ], A l nth s ., Ma aceae, E fl r t tr ndr C, assi o aceae, E Ct n h t n h l n, E K l r p rf r t , E K l r d pr , E nn nt r b . . o s . & G. o s ., S t l , A Icaci aceae, E d rp h ll Ki k, o oca aceae, E Ct n h t n p r v r d , E Aph n h t n d r , E r x ll l n ( ook. . ieg ., o a aceae, d rp t t r G. e . e . o i am ., E o oca aceae, E Ct n h t n p r v r d , E Aph n h t n p b n , E r r n Mi . , au aceae, A Cr t ll t t rn t , E C h p r d , A r p z d r d , E h l n p s ., O c i aceae, A U b n h t n d l , E C h p r d , A U b n h t n b ll t , E o s ., o emo iaceae, A U b n h t n p ll p , E C h p r d , A d rp s ., o oca aceae, E u ge s E ge m. c . G auca, i aceae, A r p z d r d , E S t ff , A l s ., yme aeaceae, E n r tr f l t ., u aceae, A C h p r d , A C h p r d , A ln r d t . o , i aceae, A p l x ?n r , Sa icaceae, A C h p r d , A lv n r v t , A n f l nd 4 2 t r t t (E . ae i, Sa o aceae, E d p n x r f l (So . e A. Cu . C. S t ff , A Koc , A a iaceae, E S t l , A C h p r d , A r t ph l d (A.M. Cu . ems ., Ct n h t n v r d , E o e iaceae, E d p n x f r x Ki k, A a iaceae, E C h p r d , A Ct n h t n v r d , E S t ff , A d p n x l n ( C C. Koc , A a iaceae, E r t nth r n t e ., amiaceae, A C h p r d , A C h p r d , A Ct n h t n v r d , E r n p t f rr n ( . o e au ., n t b l , E o oca aceae, E l h t n fl v , E d p n x l n r ( . C. Koc , A a iaceae, r p z d r d , E E r p z l b t , E Ct n h t n v r d , E [ os s .?, eco as Ρ. r n p t t x f l ( . o e au ., ?l n r ] o oca aceae, E d p n x pl x see pl x, E Aph n h t n t , E d p n x s ., A a iaceae, E r n r n ., osaceae, A C h p r d , A C h p r d , A n t b l , E r t n r , A S t l , E rth n l n rn , A d nt r x ll r ( . . o s . & G. o s . lv n r v t , A a y, Wi e aceae, E S t l , [ ], A Ct n h t n p r v r d , E r n v ., osaceae, A In l p t ll , E C h p r d , A l h t n fl v , E rth n l n rn , A d nt r l r t ( aou a y, lv n r h dr n , A Wi e aceae, E r n l r r ., osaceae, A Aph n h t n n n p , E C h p r d , A Ct n h t n p r v r d , E r n p r ( . a sc , osaceae, A In l p t ll , E lv n r v t , A l h t n fl v , E r n p r ( . a sc a n p r n t b l , E (Suckow C.K. Sc ei ., osaceae, A d nt r s ., Wi e aceae, E C h p r d , A K l r p rf r t , E S t ff , A n t b l , E r n l n i ., osaceae, A d s ., My aceae, A C h p r d , A S t l , [ ], A S t ff , A t r d l nt (G. o s . Cockay e, r n rr l t i ., osaceae, A e s ae iaceae, lv n r h dr n , A C r pl t n n , A r n s ., osaceae, A S t l , A rth n l n rn , A r n ., osaceae, A lv n r v t , A lv n r v t , A S t l , [ ], A S t l , A d p n x rb r (Mu ay i i so , r p r f l ( u . . akai, osaceae, A A a iaceae, E S t l , A Cr t ll t t f , E Ct n h t n p r v r d , E Ct n h t n v r d , E Q r pl l tr Mü c ., agaceae, A n t b l , E rth n l n rn , A 2 8 d n & nd r n (2000 : C d (In t : pt r : C d pl x (G. o s . A. . Mi c e , . r p n ( u . S ac ., o i & M. ea s [sy d p n x ig o iaceae, A pl x (G. o s . i i so ], A a iaceae, S t l , [ ], A Ε nth p W. . . O i ., ig o iaceae, Aph n h t n bt l , Ε Ε n t b l , Ε C h p r d , A h p l t l p d . We . & u e, h nb r s ., u e giaceae, A A ecaceae, Ε C r pl t n n , A l h t n n , Ε S t ff , A b n r ., G ossu a iaceae, A r n t r (A. Cu . .A.S. o so & .G. C h p r d , A iggs, o eaceae, Ε rth n l n rn , A Ct n h t n t r , Ε lv n r v t , A ee e , ?Cya eaceae, Ε S t ff , [ ], A S t ff , A p n nd n . . o s . & G. o s ., i ogo aceae, Ε r n s ., Sc o u a iaceae, A Ct n h t n p r v r d , Ε C r pl t n n , [ ], A l h t n fl v , Ε t x l n Ki k, e e aceae, Ε s ., osaceae, A C r pl t n n , A S t l , A C h p r d , A r n s ., amiaceae, A Cr t ll t t rn t , Ε C h p r d , A Ct n h t n h l n, Ε S t l , A Ep l d h t n p p r , Ε b x [as oga e y], osaceae, A S t l , A C h p r d , A t v n f r ., i aceae, A b s ., osaceae, Α Ε C h p r d , A Ct n h t n v r d , Ε C l n l , A K l r p rf r t , Ε lv n r v t , A rth n l n rn , A S ntp l s ., Ges e iaceae, A rth n l n p r , A S t ff , A S t ff , A S h ffl r d t t . . o s . & G. o s ., S t l , A A a iaceae, Ε Ct n h t n p r v r d , Ε W h n t n r b t We ., A ecaceae, A Ep l d h t n p p r , Ε S t ff , A r p z l b t , Ε W n nn r . ., Cu o iaceae, Ε S h n s ., A aca iaceae, A Aph n h t n h , Ε S t l , [ ], A l h t n l rp , Ε S n s ., As e aceae, Ε l h t n fl v , Ε C h p r d , A U b n h t n b ll t , Ε S l n v l r G. o s ., So a aceae, Ε W n nn lv l So . e A. Cu ., C r pl t n n , A Cu o iaceae, Ε C h p r d , A Aph n h t n h , Ε S t l , A l h t n l rp , Ε S l n l n n ., So a aceae, A W n nn s ., Cu o iaceae, Ε C h p r d , [ ], A l h t n fl v , Ε S t ff , [ ], A W t r s ., a aceae, A S ll h t r ph ll i ., i os o aceae, A rth n l n p r , A C h p r d , A S t l , [ ], A Str bl h t r ph ll ( ume Co e , Mo aceae, Ε Ct n h t n h l n, Ε n f l nd 4 2 A e i Β. A a e ica is i g o e emic so f sca e ge e a a s ecies wi ei associa e os r h l tt r p nd (As e aceae a s a ami ies. C pr s . ( u iaceae n nth r s . ( io aceae Aph n h t n M l t r fl r ( io aceae h n hl M l t s . ( io aceae Ch n hl fl v n (G ami eae M p r l t (Myo o aceae d r d p n x rb r (A a iaceae r ph ll tr v r (E ac i aceae l pt p r r ph ll s . (E ac i aceae K nz r d (My aceae d rp h ll ( o oca aceae pt p r p r (My aceae r pt p r s . (My aceae r ph ll n l r (E ac i aceae n f r ph ll b l t (E ac i aceae th f f ( agaceae i n n p th f tr n t ( agaceae Ar t t l fr t (E aeoca aceae rn t C pr pr p n ( u iaceae l h d t r ( au aceae C pr s .( u iaceae El rp s . (E aeoca aceae C r t n t r (Esca o iaceae d r rb r (Mo imiaceae M hl nb tr l ( o ygo aceae th f tr n t ( agaceae Ol r n l r f l (As e aceae d rp t t r ( o oca aceae l nth d v r t (Ma aceae t x l n ( e e aceae d nt r l r t (Wi e aceae rn t ll h K nz r d (My aceae W n nn r (Cu o iaceae pt p r p r (My aceae W n nn lv l (Cu o iaceae pt p r s . (My aceae t p n f l t (E ac i aceae r n p t t x f l ( o oca aceae M tr d r s . (My aceae pr n C pr s . ( u iaceae Ct n h t n b p r (Sc o u a iaceae h l n p b n Cl t nn n h ( a u cu aceae l h d t ( au aceae C pr s . ( u iaceae C rd l n b n (As e iaceae C r n rp l v t (Co y oca aceae Gr l n l d (G ise i iaceae El n t s . (My si aceae d r rb r (Mo imiaceae t l r ( au aceae t l r ( au aceae M l p pl x ( u aceae M d l f l (Sa a aceae l nth s . (Ma aceae tt p r t n f l ( i os o aceae Str bl h t r ph ll (Mo aceae d rp t t r ( o oca aceae t x l n ( e e aceae bt l p r v r d C r n rp l v t (Co y oca aceae Al p fl v d ( o a aceae El n t s . (My si aceae d n v (Sa i aceae d r rb r (Mo imiaceae Gr l n l tt r l (G ise i iaceae M r t n l r (A a iaceae Gr l n l d (G ise i iaceae M d l f l (Sa a aceae d r rb r (Mo imiaceae tt p r s .( i os o aceae h r s . (Ma aceae pl x (A a iaceae r l n v z l nd (Mo imiaceae Cr t ll t t M r n tr l (My si aceae f M r n l n (My si aceae th f f ( agaceae nn nt r b (Icaci aceae th f nz ( agaceae r x ll l n ( o a aceae 240 d n & nd r n (2000 : C d (In t : pt r : C d d p n x rb r (A a iaceae C th s . (Cya eaceae d nt r x ll r (Wi e aceae l nth s . (Ma aceae d nt r l r t (Wi e aceae p r d pr p n nd n ( i ogo aceae b br h ph n (Sc o u a iaceae S h ffl r d t t (A a iaceae b d r (Sc o u a iaceae t r p rf r t r n t r ( o eaceae C pr l d ( u iaceae v r d C pr r p n ( u iaceae C pr s . ( u iaceae C pr p th l t ( u iaceae M r t n l r (A a iaceae C pr s . ( u iaceae d p n x rb r (A a iaceae r t t ( am aceae d p n x r f l (A a iaceae Gr l n l tt r l (G ise i iaceae d p n x f r x (A a iaceae M l p pl x ( u aceae d p n x l n (A a iaceae r n s . (A ocy aceae d p n x ?l n r (A a iaceae fl r t tr ndr ( assi o aceae b s . ( osaceae tt p r n d ( i os o aceae tt p r r ( i os o aceae Ep l d h t n tt p r t n f l ( i os o aceae p p r tt p r t rn r ( i os o aceae Al tr n x l (Sa i aceae tt p r b ll t ( i os o aceae Ar t t l rr t (E aeoca aceae tt p r s . ( i os o aceae Ch p r l n n (Cu essaceae d nt r s . (Wi e aceae C pr r rp ( u iaceae b s . ( osaceae C pr r b t ( u iaceae C r n rp l v t (Co y oca aceae n h t n x l p t b l (Me iaceae t t d r rb r (Mo imiaceae M tr d r x l (My aceae r l n v z l nd (Mo imiaceae M tr d r r b t (My aceae M r p p r x l ( i e aceae M tr d r s . [ a a] (My aceae M l t s . ( io aceae tr d r M tr d r s . [ a a] (My aceae M tr d r b ll t (My aceae tt p r n d ( i os o aceae M tr d r s . [ a a] (My aceae tt p r s . ( i os o aceae n r S h ffl r d t t (A a iaceae M tr d r b ll t (My aceae t x l n ( e e aceae M tr d r s . (My aceae t ll r In l M tr d r x l (My aceae p t ll El rp s . (E aeoca aceae l h t n d r rb r (Mo imiaceae d d tt p r l n ( i os o aceae G lth r s . (E icaceae tt p r s . ( i os o aceae Ol r n l r f l (As e aceae d nt r x ll r (Wi e aceae Oz th n l pt ph ll (As e aceae d nt r l r t (Wi e aceae r h b l n (Sc o u a iaceae l rp K l r l h d t ( au aceae d pr l h d t r ( au aceae C pr rb r ( u iaceae C pr s . ( u iaceae C pr l n ( u iaceae El rp h r n (E aeoca aceae C pr f t d ( u iaceae El rp s . (E aeoca aceae C pr rh n d ( u iaceae d r rb r (Mo imiaceae C pr s . ( u iaceae Il t l s . ( o a aceae n f Ν Ζ l nd 4 24 t l r ( au aceae d nt r s . (Cu o iaceae W n nn r (Cu o iaceae pl x (A a iaceae W n nn lv l (Cu o iaceae fl v U b n h t n l h d t r ( au aceae d l r h l tt s . (As e aceae d rp t t r ( o oca aceae El rp d nt t (E aeoca aceae b ll t b ll pt ? (Sc o u a iaceae K nz r d (My aceae d r rb r (Mo imiaceae pt p r p r (My aceae pt p r s . (My aceae d rp t t r ( o oca aceae M l t r fl r ( io aceae W n nn r (Cu o iaceae M r n tr l (My si aceae h n nth r M r n l n (My si aceae ?A t l b n (As e iaceae tt p r s . ( i os o aceae d r rb r (Mo imiaceae r n p t f rr n ( o oca aceae n nth r s . ( io aceae d nt r x ll r (Wi e aceae M l t r fl r ( io aceae d nt r l r t (Wi e aceae M r n d v r t (My si aceae p n nd n ( i ogo aceae M rt s . (My aceae W n nn r (Cu o iaceae b t W n nn s . (Cu o iaceae tt p r t rn r ( i os o aceae n p ll p h p l t l p d (A ecaceae d rp t t r ( o oca aceae p ll n K nz r d (My aceae pt p r p r (My aceae pt p r s . (My aceae p n t t Ol r r d nt (As e aceae r p z l b t l hn fr r ( ec aceae r d pr n ( o oca aceae d r rb r (Mo imiaceae S h ffl r d t t (A a iaceae d r d r rp d r d d ( o oca aceae r d pr n ( o oca aceae d rp t t r ( o oca aceae d rp s . ( o oca aceae r n p t f rr n ( o oca aceae n n t l hn fr r ( au aceae e t b l d r rb r (Mo imiaceae d p n x rb r (A a iaceae d p n x l n (A a iaceae d p n x s . (A a iaceae d nt r l r t (Wi e aceae 242 d n & nd r n (2000 : C d (In t : pt r : C d A e i C. yme o e ous a asi oi s eco e n n t om Cocci ae i ew ea a a a ge a a e i Ad l n rt d s . ( [m] ca y y ami y, a y ge us s ecies wi i ami y. Ct n h t n p r v r d e i e om a e i e & Wa ke ( a .A. En rt nf l x (Em e o e y (u u is e eco s . S t ff E n rn r (Wa ke A E I I AE C h p r d rd l s . M t ph r nt A ecke & My a In l p t ll C h p r d C ph hr owa M t ph l n b r ( owa C r pl t n n S t ff S t ff M t ph t b rl (Is ii S t l rth n l n p r C ph t ll r ( a ma M r t r fl v ( owa C h p r d C h p r d lv n r h dr n S t l En r tr n (C aw C r pl t n n A YGAS I AE C h p r d Err l s . (u esc i e E x nth ll ph llpp Si es i In l p t ll C r pl t n n C h p r d E OMA I AE C l n l Aph b t ll owa K l r p rf r t Ct n h t n h l n rth n l n p r Ct n h t n p r v r d lv n r h dr n Ct n h t n s . S t ff Cocci ae i r z ( sy i ae ga s S t l Aph b t n n ( oucek Aph n h t n h Cr t ll t t l pt p r E CY I AE Ct n h t n p r v r d Ad l n rt d bl t thr h oyes Ep l d h t n p p r Ct n h t n s . In l p t ll In l s . K l r d pr K l r p rf r t K l r p rf r t Ad l n rt d n n t n oyes n h t n t ll r Ct n h t n p r v r d M r n l l f rn ( owa Ct n h t n v r d S t l K l r d pr Ad l n rt d r oyes Ct n h t n s . Ad l n rt d v oyes Ct n h t n s . Ad l n rt d n l r oyes n h t n s . Ad l n rt d v r b l oyes Cr t ll t t l pt p r Cr t ll t t rn t Ct n h t n v r d s . In l K l r d pr K l r p rf r t n f l nd 4 24 A e i . Geog a ica coo i a es o oca i ies. u e i , 4 0 0 Co o i a es s ou e ea as 00°00 S 000°00 Ε. U i e I, S 40 0 0 e wo e e a ea co es o ow C os y t l. ( 8 . usky Sou , Sea I, 4 4 662 A au a, 422 2 Eas Ca e, G 4 8 Aka oa, MC 4 4 2 8 Eas Ca e, ig ouse k, G 4 8 Aka a awa Sa e, W 40 0 E es a ey E es us , 4 20 04 A e , a ie s k, 4220 2 Amo eo ay, C 64 26 ai y a s, Wai ake e a, KA 6 4 A c o I ( y ake , 4 4 66 2 e ow k, Wai ake e a, AK 6 4 A a aki, S 8, O 8 46 e c e C eek, I a ga ua, 4 0 A gowa , 4 04 0 8 ock ouse, WI 40 6 A u s ass, C 42 a ose , W 4 28 0 Auck a , Owai aka, AK 6 44 Auck a , S e ie s, AK 6 4 Ga e s a ey, e so , 4 2 0 Auck a , i i a gi, AK 6 6 440 Ga ey C eek, 420 Auck a U i e si y g ou s, AK 6 446 Gi e I 6, eaksea Sou , 4 6 6640 Auck a , Wa e ay, AK 0 4 Gi ies I, usky Sou , 4 44 66 6 Awakau oa , K 84 444 Gis o e, G 840 80 G e E e , Auck a , AK 6 4 au a I, 4 66 Go e ow s, 4 2 e e s, e e ga es, AK 6 4 0 Go o C eek, Go e ow s, ... 4 4 2 i i gs a , MC 4 0 242 Go e o s ay, MC 4 8 2 ig Sou , Wi a i es , 44 6 2 Gowa i ge, u e , 4 4 2 ue Mou ai s, S 4 4 6 2 G a i e o es , 42 u i , G o y ack, S 46 6820 G ays us , e o , G 8 6 a s aw Sou , eci ice Co e, 4 6 6 0 eaksea I, 4 66 8 a Moo ay, Kaikou a, KA 42 4 i a ei a s, ag a , WO 4 4 4 a moo ay, S ewa I, SI 46 4 680 uce a k Sce ic es, WI 8 2 au u ga oa a, O 84 4 u e Go ge, 4 4 icks ay, W a eka ika , 4 8 8 u e , Mi e o , 4 8 i ewai es, a ks e i su a, MC.. 4 4 0 o okaka, a gi ua a, 2 408 Ca es o , 4204 uia, AK 00 4 4 C a ky I e , . Ca ma I, 4 2 664 uia am, AK 00 4 4 C a es o A iso a , u ua a, AK 0 2 4 4 2 4 4 26 C a eys Co e , MC 4 2 2 I a ga ua a i g, 4 4 C a es Sou , E ae o I, 4 06 6 08 C is c u c , MC 4 0 24 Kaikou a, KA 4224 4 Co i gwoo , 404 24 Kai ga oa S , 824 6 4 Co i e i , C 6 8 28 Kaiwaka, ΑΚ 6 0 426 Cook Sa e, W 4 2 004 Kaka ui, ase o coas a c i , G 824 Co wa is, S aggs Mo ume , AK .. 0 4 6 Kaka ui, 2 0m, G 824 Co sai ay, y e o , MC 4 24 Kaka ui, 00m, G 824 Cu e Co e, ese a io I e , ... 4604 6640 Kaka ui eas wes sa e, G 824 Kamo, 4 4 8 a s C eek, u e Mai ai , 4 22 Ka aka uw e o a ey , Wai ia a, G agg Sou , A c o age A m, 4 2 66 2 3740 / 8 8 a ga i e, 6 2 Ka ama u a , Wai ake e a, AK 00 4 4 e is o , 4 44 48 Kauae a ga S , C 0 6 ese oa Waiou u, O 2 4 Kawau I, AK 62 4 244 d n & nd r n (2000 : C d (In t : pt r : C d Kaweka oa , ack i c a, ... / 62 Mu c iso , 4 48 / 220 Ko ai ai u , Ka amea, 4 06 / 206 Ko uko u, 2 / 2 e so , 4 6 8 Kokako ui S m, Ka a oa S a io , G 82 / 804 ew ig o , MC 4 0 / 244 Ko u ikuai [mi oi ], C 0 / 4 oises Is, Mo u o o a a I, AK 64 / 4 8 oises Is, O a a I, AK 642 / 4 8 ai g o m, AK 6 8 / 4 8 ake Sy es e , 4 06 / 2 8 Oama u, 4 06 / 0 ake eka o, MK 4 2 / 0 4 O a , u e, SI 46 4 / 680 ake Waika emoa a, G 846 / 0 O aku e, O 2 / 2 ees a ey , O o , C 4 0 / 2 2 O o e, / 0 i co , MC 4 / 22 Okiwi ay, KA 42 / 2 i e a ie I, ami o k, C 6 / 0 Oku a ,AK 640 444 i e a ie I, Summi k, C 6 / 0 Oku i a ey, Sce ic es, MC 4 4 / 2 0 i e a ie , e Ma ae oa, wes , C 6 / 0 O Coac oa k, Wai ake e a, AK i e a ie I, (u e a ey k, C . 6 / 04 6 4 i e I, C a ky I e , 4 / 66 Oma u a S , Kau i Sa c ua y, .... / o i oi , O a ga, 2 / 8 0 O ekaka, 4046 / 242 y e o , MC 4 6 / 242 O e o o ay c eek, icks ay, ... 6 / 8 8 O ou i a ey, Ma o oug , S 4 2 / 4 Maimai, 420 / 4 O e e o es , e uia u , 40 / 8 Mai ai , 4 20 O o go o go a ey, W 4 2 / 4 4 Maka u S u k, Kaweka a, / 624 O aki o ks, W 40 2 4 Maka o a ("Maka oa" , O 44 / 6 4 O a e i o es, a ks e i su a, MC 4 0 / 0 Mamaku, Aqua ius , 80 / 8 O i a, W 42 0 / 4 Ma a ou i, 4 4 6 6 O i a Go ge, W 4248 / 4 Ma ga o ue, a u a i ge, WO .... 8 6 / 4 4 Owai aka, Auck a , AK 6 / 44 Ma ga a gi ese oi , Wo kma k, AK 3707/17513 a me oa , S i gs u c io , .. 4224 / 208 Ma gawi i asi , W i i aki S , O 8 / 64 a gaki S m, aemako es, WO 8 0 / 4 6 Ma s e a ey, 4 20 / ao eo e, G / 82 Ma uia, 42 / 2 a a a a , o e y .S. a e, AK / 0 Ma uia S i gs, 422 / 220 a a ea, 40 / 2 Ma aka a I, / 60 a a Swam , ic o , S 4 22 / 6 Ma aki aki , Mu c iso , 4 48 2 e o us i ge, M 4 8 / 4 Ma uku es, a a ga k, AK 6 2 / 428 ic ic oi , Ma i a ua, W 4248 / 0 6 Maw e a S [ we e Mi e C eek], 4226 / 2 i o i o oa , au u ga oa a, O 8 / 26 Mi gi ui a ea, O 8 6 / 64 o a gi a a ey, o a a es, I 400 / Mo ueka, 4 06 / 00 o u u, Awa e e i ge, G 4 / 82 Mo ueka, ea o s us , 4 06 / 00 okaka, O 8 24 Mo umoku us , 6km W Mu u a a, oko o o, Mo ueka , 4 / 2 824 / 6 8 oo K ig s Is, aw i i a i, 28 / 444 Mo u iko, 4 2 / 24 o e s ass, MC 4 8 4 M A u , o a u , 4 / 24 rt Mo ueka, 4 08 / 0 M A u , o a k, 4 0 242 u eo a o es , O 8 / 6 M Egmo , K 8 / 404 u eo a o es o ge, Gu y , O 82 / 4 M o es o e, 4 0 24 u eo a o es , Wai a a o ge k, Ka ama ama M Ma ews, imu aka a, W 4 2 / 0 S m, O 82 4 M O o , C 4 / 20 u akaiki, 420 / 20 M ua e u, O 6 / 4 M S e e s k, ai am, 4048 / 22 ag a , WO 48 / 4 2 M e A o a, ui Mi e oa si e, .. 2 / 4 ai a ey, 4 4 n f Ν Ζ l nd 4 5 a gemo e k, Wai ake e a, AK 6 6 4 e Koau, us wa k k— ookou k, a gi o o I, AK 648 4 2 6 8 a gi o o I, Ki ey e G e , AK 648 4 e Koau, o e s Wa c i g og k, a gi o o S a io , Ma ga u u, ΤΟ 820 2 8 aou I, Ke ma ec Is, ΚE 2 6 e Koau, 24 m, 8 8 awe e, 2 e Koau, 60m, 8 8 e ma s C eek, ee o , 420 e a ae ,WA 4 0 4 ee o , 420 e uke, 4 620 e eaui a Swam , eec i ge, 804 e We a S , K 4 eso u io I, isa oi me Co e, e W ai i [ Mi gi ui], O 8 4 646 4 66 omso s us , I e ca gi , S 462 6822 eso u io I, aci e a ou , 4 4 66 ee Ki gs Is a s, G ea I, ΤΗ 4 0 208 icca o us , MC 4 2 2 6 ee Ki gs Is a s, Sou Wes I, ΤΗ 4 204 i e ea o es , a ow oa es, AK ee Ki gs Is a s, Wes I, ΤΗ 4 202 642 4 4 i i a gi, Auck a , AK 6 6 440 iwaka, 4 0 00 iwai oi , S 46 6 6822 ock i e, Ao e e , 4044 2 8 o a a ese e, o a gi a a ey, I 400 ocky C eek i ge, S 4 22 46 o a a, Oama u, 4 08 0 o oe u S a e o es , 800 6 io , S 40 0 400 o o ua, A iamu i , 8 2 6 4 wi eaks i ge, uia, AK 00 4 4 o o ua I u se y, 8 0 6 6 ua a u a, Mimi a S m, ΤΟ 840 6 6 Wae ga us , o i oi , 4 80 ua a u a, W aka a e , 8 6 8 Waia o o S m, 8 0 u y ay, e so , 4 4 0 Wai a a ( ack , au u ga oa a, O 3842 / 17540 Sec e a y I, 4 4 66 Wai o ai a ey [mi oi ], M 4 40 Sec e a y I, G o o ay k, 4 66 Wai o a, au u ga oa a, O 8 6 40 Sewe eak, G eymou , 4224 2 Waimi a, S ( e Kui i , O 8 20 S a us , Wai ake e a, AK 6 4 4 4 Wai a a, C 4 0 24 S e y a ey, 4 2 242 Wai ia a, Ka aka uw e o a ey , G Sig o e e i , o i s, MC 4 8 2 6 3740 / 17818 S a u C eek, 42 0 4 Wai oua, 40 S o sma Co e, Coo e I, 4 44 6648 Wai u a, 42 4 S i g ie , MC 4 20 6 Wai oa Go ge, 4 2 0 Sou i a eac , AK 6 8 428 Wai ake e a, AK 6 4 2 S a e o es 0, Kaima awa o , ΤΟ Wai ake e a, Sce ic i e, AK 6 6 4 4 3857 / 17614 Wai ui Sa e ("Wai ui Sa e" , O 8 2 2 S e ie s, Auck a , AK 6 4 Wa ke us ack, Wai ake e a, ΑK 6 4 4 S o e e ge, ock & i a a, CO 4 28 00 Wa e , ΚA 42 6 06 Wa kwo , AK 624 440 aikawakawa, Mau gakaka, G ... 40 82 Wa e ay, AK 0 4 ai ua S [ ookou owe W a gama a], C We i g o , W 4 44 3710/17549 Wes o , mi es o , 4 4 4 amaki , I 4008 60 Wes amaki a ey, ua i e a, I 4007 / 17602 a ako e, 40 0 2 4 W aka uaka [S m], 4 0 26 aw ai a s, W aka a a ui S m, ΤO 0 0 W aka ewa ewa ( I , 8 0 6 aw ai S [Me i igs], 42 4 4 W a gamoa Sa e, 4 26 aw i i a i I, oo K ig s Is, 28 444 W a ga oa, 0 4 e A ai Sa c ua y, ou o a, 442 2 6 W i i aki, O 8 4 642 e A a oa eac , [ u a uku S m], 6 6 8 W i e i e us ese e, 6 e Koau, us Wa k, 6 8 246 d n & nd r n (2000 : C d (In t : pt r : C d O S O E O S O E IS A S \z IS A S ❑ Ke ma ecs ❑ Ke ma ecs ee Ki gs ❑ ee Ki gs ❑ <, ι C a ams ❑ . C a ams ❑ a . ❑ S a es ❑ . , S a es φ e ❑ ou y ❑ Ί ou y A i o es ❑ A i o es ❑ Auck a s ❑ Auck a s ❑ Cam e ❑ Cam e ❑ i i : • ι( 4• Ma 0 Co ec io oca i ies, Aph n h t n h n hl . Ma 02 Co ec io oca i ies, Aph n h t n d r . O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ❑ ❑ ee Ki gs 4. ee Ki gs ❑ C a ams ❑ C a ams k ° q $ S a es ❑ . S a es ❑ ou y ❑ ou y ❑ ❑ ❑ A i o es A i o es ( Auck a s ❑ Auck a s ❑ • Cam e ❑ Cam e ❑ ΓΡ ,, 2 f Ma 0 Co ec io oca i ies, Aph n h t n r . Ma 04 Co ec io oca i ies, Aph n h t n n n p . n f l nd 4 24 O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ❑ ee Ki gs ❑ ee Ki gs C a ams ❑ C a ams ❑ S a es ❑ S a es ❑ ou y ❑ ou y ❑ A i o es ❑ A i o es ❑ λ Auck a s ❑ Auck a s ❑ • Cam e ❑ Cam e ❑ Ma 0 Co ec io oca i ies, Aph n h t n h . Ma 06 Co ec io oca i ies, Aph n h t n t . O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs ❑ ee Ki gs C a ams ❑ C a ams σ S a es ❑ S a es ou y ❑ ou y σ A i o es ❑ A i o es I Auck a s ❑ Auck a s I Cam e ❑ Cam e ❑ Ma 0 Co ec io oca i ies, Aph n h t n pr n . Ma 08 Co ec io oca i ies, Aph n h t n p b n . 248 d n & nd r n (2000 : C d (In t : pt r : C d O S O E O S O E IS A S IS A S Ke ma ecs ❑ ‚. Ke ma ecs ❑ ee Ki gs ee Ki gs ❑ $e C a ams ❑ ζΡ:,• Ι C a ams ❑ ❑ S a es ❑ " S a es •Q•. ou y ❑ ou y ❑ ❑ A i o es ❑ A i o es • 4 Auck a s ❑ Auck a s ❑ C Cam e ❑ n Cam e ❑ ςΡ, i ( Κ Ma 0 Co ec io oca i ies, Aph n h t n bt l . Ma 0 Co ec io oca i ies, Cr t ll t t f . O S O E O S O E IS A S IS A S ❑ Ke ma ecs ❑ Ke ma ecs Νuι ❑ ee Ki gs ee Ki gs • C a ams ❑ e C a ams ❑ S a es ❑ S a es ❑ ,mú ou y ❑ ζΡ ou y ❑ A i o es ❑ i A i o es ❑ ❑ Auck a s ❑ Auck a s Cam e ❑ C Cam e ❑ . Χ • Ma Co ec io oca i ies, Cr t ll t t f . Ma 2 Co ec io oca i ies, Cr t ll t t l pt p r . n f l nd 4 24 O S O E O S O E IS A S IS A S ❑ Ke ma ecs ❑ C ,μ Ke ma ecs u 2. ❑ ee Ki gs ❑ ee Ki gs ?, C a ams ❑ C a ams ❑ Ι q S a es ❑ ou y ❑ ou y ❑ C A i o es ❑ A i o es ❑ ❑ Auck a s ❑ • Auck a s ❑ ❑ Cam e C Cam e ς n ,i nς l , 0 l λ Ma Co ec io oca i ies, Cr t ll t t n f . Ma 4 Co ec io oca i ies, Cr t ll t t rn t . O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs ee Ki gs • ❑ C a ams ❑ C a ams S a es ❑ S a es ❑ ou y ❑ ou y ❑ ❑ A i o es ❑ A i o es Auck a s ❑ Auck a s ❑ Cam e ❑ Cam e ❑ 4 Ma Co ec io oca i ies, Cr t ll t t rn t ll . Ma 6 Co ec io oca i ies, Ct n h t n h l n. 2 0 d n & nd r n (2000 : C d (In t : pt r : C d O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ee Ki gs ❑ ee Ki gs C a ams ❑ C a ams S a es ❑ S a es ou y ❑ ou y A i o es ❑ A i o es Auck a s ❑ Auck a s Cam e ❑ Cam e ‚ Ma Co ec io oca i ies, Ct n h t n p r v r d . Ma 8 Co ec io oca i ies, Ct n h t n t r . O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs • ee Ki gs ❑ C a ams I C a ams ❑ S a es I S a es ❑ ou y I ou y ❑ A i o es I A i o es ❑ ❑ Auck a s ❑ Auck a s ❑ Cam e ❑ Cam e Ma Co ec io oca i ies, C e oc i o v r d . Ma 20 Co ec io oca i ies, E e i oc i o p p r . n f Ν Ζ l nd 4 2 O S O E O S O E IS A S IS A S Ke ma ecs Ke ma ecs ❑ C ee Ki gs ❑ ee Ki gs ❑ ° ε , C a ams ❑ C a ams ❑ ❑ S a es ❑ S a es a s ou y ❑ s ou y ❑ A i o es ❑ A i o es ❑ . ι Auck a s ❑ • Auck a s ❑ Cam e ❑ Cam e ❑ l V , Γ Γ Ma 2 Co ec io oca i ies, In l p t ll . Ma 22 Co ec io oca i ies, K l r d pr . O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs ❑ ee Ki gs ❑ o C a ams ❑ C a ams ❑ S a es ❑ S a es ❑ ❑ ou y ❑ ou y • A i o es ❑ A i o es ❑ • Auck a s ❑ Auck a s ❑ Cam e ❑ Cam e ❑ Ί f ι• i 4 Ma 2 Co ec io oca i ies, K l r p r d pr . Ma 24 Co ec io oca i ies, K l r p rf r t . 2 2 d n & nd r n (2000 : C d (In t : pt r : C d O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs I ee Ki gs ❑ ee Ki gs I C a ams ❑ C a ams I S a es S a es ❑ I ou y I ou y I A i o es A i o es I I Auck a s Auck a s I I Cam e Cam e ❑ ( Ma 2 Co ec io oca i ies, n h t n t t . Ma 26 Co ec io oca i ies, n h t n tr d r . O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs ❑ ee Ki gs ❑ C a ams ❑ C a ams ❑ S a es ❑ S a es ❑ ou y ❑ ou y ❑ A i o es ❑ A i o es ❑ Auck a s ❑ Auck a s ❑ Cam e ❑ Cam e ❑ Ma 2 Co ec io oca i ies, n h t n n r. Ma 28 Co ec io oca i ies, n h t n t ll r . n f l nd 4 2 O S O E O S O E IS A S IS A S ❑ Ke ma ecs ❑ Ke ma ecs C ❑ ❑ : ee Ki gs " y ee Ki gs • ς C a ams ❑ C a ams ❑ S a es ❑ ou y ❑ ou y ❑ , . A i o es ❑ A i o es ❑ ❑ ❑ Auck a s Auck a s , $ Cam e ❑ Cam e ❑ • Ma 2 Co ec io oca i ies, l h t n d d . Ma 0 Co ec io oca i ies, l h t n l rp . O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs ❑ ee Ki gs ❑ C a ams ❑ C a ams ❑ S a es ❑ S a es ❑ ou y ❑ ou y ❑ A i o es ❑ A i o es ❑ Auck a s ❑ ( Auck a s ❑ Cam e ❑ Cam e ❑ ζΡ Ma Co ec io oca i ies, l h t n fl v . Ma 2 Co ec io oca i ies, l h t n n . 5 d n & nd r n (2000 : C ld (In t : pt r : C d O S O E O S O E IS A S IS A S ❑ Ke ma ecs ❑ Ke ma ecs Ma Co ec io oca i ies, l h t n p ll n . Ma 4 Co ec io oca i ies, l h t n p n t t . O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ■ ee Ki gs ee Ki gs ❑ C a ams C a ams ❑ S a es ι , S a es ❑ ou y μ ou y ❑ A i o es A i o es ❑ Auck a s Auck a s ❑ Cam e Cam e ❑ Ma Co ec io oca i ies, r p z l b t . Ma 6 Co ec io oca i ies, r p z d r d . n f l nd 4 55 O S O E O S O E IS A S IS A S ti. ❑ Ke ma ecs ❑ Ke ma ecs f ❑ ee Ki gs ❑ ee Ki gs ❑ C a ams ❑ C a ams S a es S a es ❑ έϊ ou y ou y ❑ A i o es A i o es ❑ Auck a s Auck a s ❑ i Cam e Cam e ❑ Ma Co ec io oca i ies, n n t . Ma 8 Co ec io oca i ies, n t b l . O S O E O S O E IS A S IS A S ❑ Ke ma ecs ❑I Ke ma ecs ❑ ee Ki gs ❑ ee Ki gs ❑ C a ams ❑ 4 C a ams S a es ❑ S a es ❑ ou y ❑ ❑ A i o es ❑ rrr A i o es ❑ Auck a s ❑ Auck a s Γ Cam e ❑ Cam e ❑ r Γ ✓ ςΝ ς, . n S Ϋ Ma Co ec io oca i ies, U b n h t n d l . Ma 40 Co ec io oca i ies, U b n h t n b ll t . 5 d n & nd r n (2000 : C d (In t : pt r : C d O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs ❑ ee Ki gs ❑ C a ams ❑ C a ams ❑ S a es ❑ S a es ❑ ou y ❑ ou y ❑ A i o es ❑ A i o es ❑ Auck a s ❑ Auck a s ❑ Cam e ❑ Cam e ❑ Ma 4 Co ec io oca i ies, U b n h t n h n nth r . Ma 42 Co ec io oca i ies, U b n h t n j b t . O S O E IS A S Ι.. Ke ma ecs ❑ „y ee Ki gs ❑ C a ams ❑ S a es ou y ❑ A i o es ❑ Auck a s ❑ Cam e ❑ Ma 4 Co ec io oca i ies, U b n h t n p ll p . n f l nd 4 2 A O OMIC I E is i e co e s e omi a i e e a e a u ga a a me io e i e e , ega ess o ei cu e s a us i a o omy. a a i o y e a e ose i c u e i e c eck is . a a o e a Cocci ae a e is e i e o e ge us s ecies. age um e s i o y e e o e e s a o a esc i io , a i t l t p a esc i i e igu e. SEMs a e e i e M a co ou a es a e e i e C . e su i k i ica es a key, a e su i m a ma . AC E I AE h l n, Ct n h t n , 2, 2 , Μ2 , d pr , K l r 0, , C40, , t t , n h t n 0, 2 , Μ2 , Μ , 0 8, 4 , 8, , 0 k, 04, 8 , 04, 20, 2 , 24k, 26, 2 , C4 , 2, 0 2k, , 4, 8, 0 , 08, , 20, 2 ,2 2 2 6, 2 , 2 2, 2 6, 240, 242, 2 m 40, 2 , 240, 2 2m 2 8, 2 , 242, 24 m d pr , S t 20 Ad l n rt d bl t thr h 0, h n hl , Aph n h t n , , d pr , n 20 242 8k, , 68, 2, , , 2 2, 2 , d pr , Ct n h t n 2, 04, 24 Ad l n rt d n n t n 0 , , 246m 26 26, 242 h r l , n 8 d pr n r, Ct n h t n 24 Ad l n rt d r 242 CISSOCOCC AE 26 Ad l n rt d v 242 Chl r p lv n r 2 2, 2 d tr t r, C r pl t 0, 8, Ad l n rt d n l r 242 COCCI AE , , 2, , 0, , C 4, 8 k, 8 , , 2 , Ad l n rt d v r b l 2, , , 6, 2, 242 2 2 26, 0, 0, 242 COCCI AE 0, , , 84, 4, d tr t r, G rd Ad l n rt d s . 0 , 242 COCCI E I AE 6 d d , l h t n 0, , 6, 2 , d l , U b n h t n 0, 048, Cocci i 0, , 84, 4 Μ , 04 , 4 , 44k, 4 , , 2k, , 4, , 8 , 82, Coccoi ea , 2 , , 8, , 2 , 2 , 2 6, 2 6, 24 , 2 m C ph hr 2 ,242 240, 2 m A r t bb r 6, C42, 0 , , C ph t ll r 242 d r , Aph n h t n , , 8k, 4 C 0, , 2 k, 4, k 60, 6 , 0, , 2 , 2 6, 2 , An pl p tr d r 40 ff , n 208 246m A E I I AE , , 0, 2 , 242 ff , S t 0, , , C4 , IAS I I AE 2 Aph n h t n , 2, 6, 2 , 22k, 4 , , , k, 20 , 208 2 , , , , 8k, , 2 2 2 8, 242 l rp , Ct n h t n 2, , 2 Aph b t ll 04, 0 , 242 C l b p 2 l rp , l h t n 0, C4 , 4 , Aph b t n n 4, 2, 0 , 4, , l b t , r p z 0, , 6, 8 , 42 44k, 46, , , , 26, 0, 42, 242 60k, 6 , 6 , 6 , 2 , 2 , 6, 2 2 4, 2 8, 240, 2 m AS E O ECA II AE 2 2 4, 2 , 2 8, 24 , 2 4m l n t , C 8, tr l , C r pl t rn , E l n 206, 20 l n t , Ct n h t n , 2, 6, rn , n 206 84 rd l s . , 242 rn , n (E l n 206 En r tr n 242 b rb r d , n 208 rn , rth n l n 0, , , E CY I AE 2, , 0 , , 26, b rb r , C 20 C42, k, 20 , 206 208, 2 , 0, 0, 8, 242 b rb r , E l n 20 2 , 2 8 En rt nf l x 242 b ll t , U b n h t n 0, 048, , Cr t ll t t , 2, 2 k, 4 , 0, , Ep l d h t n , 0, 2, 22k, 4 , , 2k, , , 80, 82, 2 4, 6, , 80, 8 k, 88, 8, 20, 2 240 2 6, 2 8, 24 , 2 m Ct n h t n , 6, 22, 2 k, 4 , Er h t n 2 0, 2, 4, , 4, 80, , 8, E IOCOCCI AE 2, 0 l l , lv n r 2 2, 2 0 k, 08, 20, 2 Er r 2 CA IOCOCCI AE n t , n 0, , E IO E I AE n , n , 2 0, 2 22, Μ 2, 04 , 4 , 2, 66k, 6 , Err l s . , 242 n , n (S t 2 0 6 , , 2 , 2 , 24 , 242, EU ECA II AE n , S t 2 0 2 m E p lv n r 2 8 CECI OMYII AE 6 CY OCOCCI AE E n rn r 242 r f r , C r pl t 0, , 8 k, E x nth ll ph l pp , 0, 2 , 86, , 2 2 d r d , Ct n h t n 2, , 64, 242 r f r , C 8 6 CE OCOCCI AE 2 d r d , r p z 0, , 8, , f , Cr t ll t t , 2, , C r pl t 0, 2 k, 2, , 8 k C44, 04 , 4 , 60k, 62 64, C , 0 6, 80, 8 , 8 k 8 , 2, , CE O AS I AE 0, 2, , 8, 4 , 6 , 2, 2 , 2 6, 2 , 24 , 2 , 2 , 248m 0 , , 6, 84 2 4m f , In l , , 8 8 5 d n & nd r n (2000 : C d (In t : pt r : C d f l , Ch r 208 K l r , 0, 2, 4, 2 k, 4 , 0, , l , rn rd 2 f l , S t 208 , 4, 80, 8, , 20, 24k, 240 l , C 2 0 I I II AE hl Aph n h t n , , 6, l , n 2 0, 2 fl v , Ct n h t n 2, Μ , C , , 8k, 6 , 6 , 0, 2, l , n (S t 2 0 fl v , l h t n 0, , Μ 2, , 6, , 2 8, 2 , 242, 24 m l , S t , , 0, , , 8, C4 , 42, 4 k, 4 , , , C4 , 6, k, 204, 20 ,2 0 2 2, , 6, 8, 2 , 2 2 8, n h t n , 0, 2, 4, , 22k, 2 2 8, 242 24 , 2 m 4 , 0, 6, 0 2k, 240 rn t , Cr t ll t t , 2, Μ 0, fl f r , Chl r p lv n r 2 2, 2 l pt p r , Cr t ll t t , 2, C , 4 , 6, , 80, 8 , 8 k, 4 fl f r , lv n r 0, , , C4 , , 2 , 0 6, 80, 8 , 8 k, 0, 2, , 2 , 2 2 6, 2 8, 2 , 242, 0, 2 2k, 2 , 2 , 2 2 4, 2 , 242, 248m 24 m fl f r , C 2 2 l pt p r , In l 0 2, 6 rn t , In l 2, 4 f , Cr t ll t t , 2, , 80, EUCAS I I AE 2 rn t ll , Cr t ll t t , , C , 82, 8 k, 8 , 0, 2 , 2 2, 2 4, 2 6, , 80, 86, 8 k, , 2, , 2 , 2 , 2 , 248m l n l , n 8 2 4, 2 , 2 , 24 m f , Ct n h t n 2, 8 , 0 l n l , C 0, , C , k O E II AE 2 , 8, , 2 , 2 2, 2 4, 2 , G rd 8 , 2 8, 242 p r d pr , K l r 0, 20, r , Aph n h t n , 4 , , 22, 24k, 26, 2 , 2 , 2 , 8k, 6 , 6 , 0, , 4, 2 , 2 , l t , n 2 4, 240, 2 m 246m l t , C a a eca ii i MA GA O I AE 2 r t 0, , 2 k, 20 A IMOCOCCI AE 2 t , Aph n h t n , , 8k, 64, p r v r d , Ct n h t n , , 2, l h l b 6, 2 2, 4, , , 2 , 2 , 24 m 4 , Μ2 , C 8, C , C42, , r n nt p d 6 br nth , C 2 8 , 8, 00, 0 k 0 , 0 , h ph r , S t 20 br nth , lv n r 0, 2 , 2 2 , 242, 2 0m h ph r , n 208, 2 0 , C46, 0, 2 2k, 2 , 2 8, 2 , rth n l n 0, 2 k, 2, k, h ph r , n (S t 2 2 2 206 20 br nth , lv n r ll 2 8, p t ll , In l 0, 2, 22k, Μ 2, h p r d , C , 0, 8, 2 C , 4 , 0, 6, 4, 6, 2 , 0 , 4 , , 2, 4, , k, M t ph r nt 242 , 2 ,2 4,2 6,2 ,240, 6, 8, , 2 2 8, 242 M t ph l n b r 2 , 242 242, 2 m h p r d , n M t ph t b rl 242 p ll p , U b n h t n 0, Μ 0, h b l r , Ch r 208 tr d r , n h t n 0, 2, C48, 2, , 2k, 8, , 82, hlb rn l r , n 20 , 2 0 , C4 , 2, 0 2k, , 8 , 2 6, 24 , 2 6m h dr n , E p lv n r 2 8 8, 4 , 2 , 240, 2 2m p rf r t , Ct n h t n 2, , , h dr n , lv n r 0, , 046, MIC OCOCCI AE 2 2 2 0, 2 2k, 2 4, 2 , 2 4, 2 , 242 M r t r fl v 8, 242 p rf r t , K l r 0, 2, , h n nth r , Ct n h t n 2, , n r, n h t n 0, 2, , 6, Μ , C40, 4 , 20, 2 , 24k, 80 0 2k, 6, 40 42, 2 , 26, 2 , 2 , 0, 2 2, 2 , h n nth r , U b n h t n 0, 240, 2 2m 2 2 8, 240, 242, 2 m C48, , 2k, , 6, , r nd , S t 2 p r , Ch r 20 80 82, 2 , 2 4, 2 , 24 , M r n l l f rn 2 , 242 p r , E l n 20 , 208 2 6m r , n 20 p r , n 20 p r ll d pl x 6, 042, 8 , , MY O ECA II AE p r , n (E l n 20 6 p r , rth n l n , 0, n l t , S t 2 2, , k, 202, 20 , 208, 2 2, n n p , , 2, Aph n h t n n f , Cr t ll t t , , 0 6, 2 8, 242 Μ 0, C , , 8k, 62, , 2, , , 80, 84, 8 k, 88, 2, , 2 , p r p n , n 2, 80, 2 , 2 2, 2 2 , 2 , 2 , 24 m (E l n , 2, 20 , 208 246m n r , r t 0, 04 , k, p r p n , rth n l n n n p , In l , , 2 200, 20 , 2 2 4, 2 20 n l , 0, 2, , 22k, 4 , 4, n r , S t 20 E ACO EAC II AE 2 , 240 n r , n 20 p p r , Ct n h t n n r , n (S t 20 p p r , Ep l d h t n 0, 2, , j n r n , C (C r pl t 8 n r a . d pr , n 20 22k, Μ 0, 0 , 4 , , , , 8, j b t , U b n h t n 0, Μ 0, , n , l h t n 0, , 6, 044, 2, 4, , 2 2 6,2 8,240, 2, k, , 8 , 82, 4 k, 48, , , , 6, 242, 2 0m 2 6, 24 , 2 6m 8, 2 8, 24 , 2 m A YGAS E I AE , 242 n f Ν Ζ l nd 4 59 l h t n , 0, 2, 6, 2 k, 4 , 0, p n t t , l h t n 0, 6, th , rth n l n 208 , 42, 4 k, , , 6, 4 k, 0, , , , 8, t r , Ct n h t n , 2, 8, 0 , 8, , 2, 240 , 2 , 24 , 2 4m 0 k, 04, 0 , 08, 2 8, 240, p ll n , l h t n 0, 4 6, t 2 2 0m Μ , C44, , , 42 44k, 4 , r p 6 , , , 6, 8, , 2 4, r b , n 206, 20 t b l , n 0, 6, 24 , 2 4m h z b f r t r 6, 2 Μ 0, 4 , 66k, 68, 6 , 0, , r p z , 0, 2, 4 6, 22k, 4 r r , n 206 208 3 37 3 1 55m , , 6, , 60k, 24 r b n , C r pl t 0, 8 , 88, r ph p rn 6 n , 0, 2 4, 22k, 2 4 , 0, , 4, 6, 66k, 24 r , C r pl t 0, 8 , 8 , 2 U b n h t n , 0, 2, 2 k, 4 , 2, r l dv n 8, 6 r , C 2 , 4 , , 2k, , 24 pr n , Aph n h t n , , C 4, , , 8k, 6 , , 6, , 2 2, S t 0, , 2 k, 0, 2, k, rt ll l n , 042, 0 , 4 2 4, 2 , 24 m 208, 20 , 2 v r d , Ct n h t n , , 2, 4, 6, SEU OCOCCI AE 2 Saisse ii i 0, , , 84, 4, , 22k, Μ 2, C 8, 4 , , 8, p d , lv n r , 2 2, 2 t ll r , n h t n 0, 6, 02, 0 k 08, 0 , 2 2, E OMA I AE 4, 2, 04, 0 , C4 , 6, 0 2k, , 13 1 1 2 , 2 , 2 8, 240, 242, 2 0m 4, , 26, 0, 42, 2 , 242 1 35 5 m v t , C 212, 19 p b n , Aph n h t n , 4, Μ , n n , C r pl t 0, , , v t , lv n r 0, , , C46, 0, C , 4 , , 8k, 66, 68, 2, , C 4, 8 k, 0, , 4, 2 2 2k, 2 6, 2 , 220, 2 6 2 8 6 , 2 2 6, 2 , 24 m 2 8, 242 lv n r 0, 4, , 2 k, , p n , n (E l n , W x ll 8 2 2k, 220 12, 7 u i a ii i 0, , 0, 6, 84, 4, bt l , Aph n h t n , 4, , 2 2 C , 4 , 4, , 8k, 6 , 6 8, , 2 2 2 6, 2 8, 2 , 248m d n & nd r n (2000 : C d (In t : pt r : C d 260 2° ° 4° ° 6° ° 8° 2° ° 4° ° 6° ° 8° Υι ° ° • ι . ►Δι ι. aam πιιι 6° 6° ίπ i . 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M nd & Ann tt K W l r IS 0 4 0668 .2 ec 82. 20 . $2 . 2 Oso ii ae (I sec a: Co eo e a: S a y i i ae • . l n M C ll IS 0 4 06688 .2 ec 82 • 6 . $ 8.60 A i i ae (I sec a: Co eo e a • .A. ll IS 0 4 06 0 4 • 2 ec 82.2 2 . $4 .00 4 E io yoi ea e ce E io yi ae (A ac i a: Aca i • .C.M. M n n IS 0 4 06 4 • 2 o 84. 44 . $2 . E io yi ae (A ac i a: Aca i: E io yoi ea • .C.M. M n n IS 0 4 06 46 8. 4 o 84. 28 . $2 . 6 y ae i ae (I sec a: Co eo e a • .G. Ord h IS 0 4 06 4 6. 2 o 84.64 . $ 8.60 C y os igma a (A ac i a: Aca i a co cise e iew • M. xt n IS 0 4 06 62 • 8 ec 8 . 2 . $2 . 8 Ca i o i ae (I sec a: i e a • . r IS 0 4 06 64 6.24 e 86. 88 . $ 8.60 o u a (I sec a • S. . x n IS 0 4 06 6 4. 24 e 86 . 2 . $ 8.60 0 u u i e a (I sec a: ysa o e a • r n A. M nd & Ann tt K W l r IS 0 4 06 84 0 • 22 Se 86. 44 . $ 4.6 seu ococci ae (I sec a: emi e a • .M. C x IS 0 4 06 . A 8 . 2 2 . $4 . 2 om i i ae (I sec a: yme o e a • A.C. rr IS 0 4 02 0 . o 8 . 60 . $ . E cy i ae (I sec a: yme o e a • .S. 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M ll n IS 0 4 02 8 8. 2 May • 64 . $24. 2 Scia o i ae, Me e e i ae (I sec a: i e a wi a ge e ic e iew o e o ic o o i ae • . . l • IS 0 4 0262 . a 2 • 4 . $2 . 24 e e i ae (I sec a: i e a • . n b r IS 0 4 026 2 • 4 Ma 2. 40 . $ 4. 2 Ce co i ae (I sec a: omo e a • K.G.A. lt n & C. . M r l IS 0 4 026 6 2 • 2 May 2 • 40 . $ . 26 e e io i ae (I sec a: Co eo e a : ca a ogue o y es a keys o a a .C. W tt• IS 0 4 026 . u 2. 0 . $2 . 2 A a c o e i ae (I sec a: eco e a • I. . M ll n IS 0 4 0 644 8. 8 e • 0 . $2 . 28 a ae o Cu cu io oi ea (I sec a: Co eo e a : a sys ema ic o e iew r nd M. M . IS 0 4 8 04 0 0 • 4 u •226 . $ .00 2 C y o y c i ae (I sec a: Co eo e a: Cu cu io i ae C. .C. l • IS 0 4 8 04 8 2. 2 ec . 08 . $6 .00 0 e ia i ae (I sec a: e i o e a • .S. d l IS 0 4 8 04 24 . Ma 4. 64 . $42. 0 a i i ae (C us acea: Am i o s • K.W. n n IS 0 4 8 04 6. Oc 4. 28 . $ 6.00 2 S eci ae (I sec a: yme o e a • A.C. rr IS 0 4 8 04 4 4. Oc 4. 2 . $ . 0 Mo a i i i (I sec a: yme o e a • .A. rr IS 0 4 8 04 8 . 8 May . 82 . $2 . 4 A ici ae (I sec a: Co eo e a • .G. W rn r& .S. Ch ndl r IS 0 4 8 04 4 6. 2 u . 64 . $26. 0 Cy i ae, Aca osoma i ae, a e a omi ae (I sec a: e e o e a : sys ema ics, geog a ica is i u io , a ioeco ogy • M. C. r v èr IS 0 4 8 0 0 2. 2 o . 2 . $42. 0 6 e o e ii ae (I sec a: E eme o e a • . . n & W. . t r IS 0 4 8 0 0 • Aug 6. 44 . $ . 0 Co eo e a: ami y g ou e iew a keys o i e i ica io • . Kl z & .C. W tt• IS 0 4 8 0 2 8. Aug . . $4 . 0 8 a u a ise e es ia S y omma o o a (Mo usca: Gas o o a G.M. r r. IS 0 4 8 0 22 . 2 a • 2 . $ 2. 0 Mo y i i (I sec a: Co eo e a: Cu cu io i ae: Mo y i ae • .C. Cr IS 0 4 8 0 2 • 4 e • 68 . $2 . 0 40 Ci ii ae (I sec a: emi e a: Auc e o y c a • M. C. r v èr IS 0 4 8 0 4 . 2 o . . $ . 0 4 Cocci ae (I sec a: emi e a: Coccoi ea • C. . d n & . C. nd r n IS 0 4 8 0 • e 2000. 264 . $ 2. 0 isi e Ma aaki W e ua ess We si e a : www.mw ess.co. o u e i o ma io , a o gai access o o i e e ac s om ese u ica io s. 264 d n & nd r n (2000 : C d (In t : pt r : C d O ICES GĀ A UI This series of refereed publications has been established to Kua w aka ū ia ē ei ui ga uka uka ei w aka au au encourage those with expert knowledge to publish concise i gā o u ga w ai mā au a ga kia w aka u a i gā kō e o yet comprehensive accounts of elements in the New o o e ga i e w aikiko o u e ā a a ki gāā ai a ga Zealand fauna. 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Henderson O U A SUMMA Y KEYS O A A E E E CES A O OMIC I E IS - 7 - 9335-7 804 8 0 is is a acsimi e o e i e u ica io a is u y sea c a e I is su ie o i i i ua use o y a is o o e os e o we si es ( i ks s ou e ma e o e age om w ic i was ow oa e o a o is wo k co e e y co y ig may e e o uce o co ie i a y o m o y a y mea s (g a ic e ec o ic o mec a ica i c u i g o oco yi g eco i g a i g i o ma io e ie a sys ems o o e wise wi ou e w i e e missio o e u is e Fauna of New Zealand we si e co y 1 a ca e esea c co o gso C ; e e so C Cocci ae (I sec a emi e a Coccoi ea n f Ν l nd 4 , a e o u ica io 3 e ua y Fauna of New Zealand, ISS 111-53 3; 1 IS - 7 - 9335-7 ew ea a Cocci ae Sca e images o i e y S e e awso (www.bugz.org.nz). OC e e co ec e o is sea c a e y e o C os y FNZ se ies e i o u y 1 Use s may e ac e om is o ei ow use u mus c eck i agai s e o igi a ocume o e seque ce accu acy a o ma i g