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SERIES EDITOR Dr T. K. Crosby Private Bag 92170, Auckland, New Zealand Fauna of New Zealand Ko te Aitanga Pepeke o Aotearoa

Number / Nama 52

Raphignathoidea (Acari: Prostigmata)

Qing-Hai Fan College of Plant Protection, Fujian Agricultural and Forestry University, Fuzhou 350002, China [email protected] and Zhi-Qiang Zhang Landcare Research, Private Bag 92170, Auckland, New Zealand [email protected]

Manaaki W h e n u a P R E S S

Lincoln, Canterbury, New Zealand 2005 4 Fan & Zhang (2005): Raphignathoidea (Acari: Prostigmata)

Copyright © Landcare Research New Zealand Ltd 2005

No part of this work covered by copyright may be reproduced or copied in any form or by any means (graphic, electronic, or mechanical, including photocopying, recording, taping information retrieval systems, or otherwise) without the written permission of the publisher.

Cataloguing in publication FAN, QING-HAI Raphignathoidea (Acari: Prostigmata) /Qing-Hai Fan & Zhi-Qiang Zhang – Lincoln, Canterbury, N.Z. : Manaaki Whenua Press, 2005. (Fauna of New Zealand, ISSN 0111–5383 ; no. 52). ISBN 0-478-09371-3 1. Raphignathoidea — Classification. 2. Raphignathoidea — Identification. 3. — Classification. 4. Stigmaeidae — Identification. I. Zhang, Zhi-Qiang, 1963 II. Title III. Series UDC 595.42

Suggested citation: Fan, Q.-H.; Zhang, Z.-Q. 2005. Raphignathoidea (Acari: Prostigmata). Fauna of New Zealand 52, 400 pp.

Prepared for publication by the series editor using computer-based text processing and layout at Landcare Research, Private Bag 92170, Auckland, New Zealand

M~ori text by H. Jacob, Auckland.

Published by Manaaki Whenua Press, Landcare Research, P.O. Box 40, Lincoln, Canterbury, N.Z. Website: http://www.mwpress.co.nz/

Printed by PrintLink Ltd, Wellington

Front cover: Mecognatha hirsuta Wood (Illustrator: Qing-Hai Fan).

Publication of the Fauna of New Zealand series is the result of a research investment by the Foundation for Research, Science and Technology under contract number C09X0202. Fauna of New Zealand 52 5

POPULAR SUMMARY HE WHAKARAPOPOTOTANGA

Class Arachnida Subclass Acari Supraorder Prostigmata Superfamily Raphignathoidea

Raphignathoid Illustration / Whakaahua: Cryptognathus sp. Mites of the superfamily Raphignathoidea are biological control agents of spider mites, eriophyid mites, and scale insects in agriculture and forestry. The majority of the Whakamahia ai ng~ pãwereriki o te wh~nau nui are free-living predators, but a few are phytophages, Raphignathoidea hei patu ~-koiora i ng~ pãwereriki feeding on moss, and symbionts/parasites of insects. pãng~werewere, ng~ p ãwereriki eriophyid, me ng~ ng~rara The superfamily can be dated back at least 56 million unahi i roto i ng~ kawenga ahuwhenua, whakatipu r~kau. years. It belongs to the supraorder Acariformes, order Ko te nuinga o ng~ momo, he konihi h~ereere noa. Heoi Prostigmata, and comprises about 770 species, and 57 anÇ, ko ‘tahi he kai pãkohu, ko ‘tahi anÇ he hanga ka genera in eleven families. They are worldwide in piritahi, ka pirinoa r~nei ki t‘tahi momo pepeke. distribution, abundant in most of the geographical regions, E ora ana ng~ t ãpuna taketake o te wh~nau nui nei i te and are even found in the Antarctic region. 56 miriona tau ki muri. Ko Acariformes te pãtoi o runga, They pass through five or six stages to complete their ko Prostigmata te pãtoi, e 770 pea ng~ momo, e 57 pea life cycles. The development from the egg to adult can be ng~ puninga i roto i ng~ wh~nau tekau m~ tahi. Kua marara completed in 1 to 3 weeks. The males develop slightly ki ng~ t Çpito katoa o te ao, ~, kei te huhua tonu ki te nuinga faster than females. Their reproduction is arrhenotokous, o ng~ momo takiw~ o te ao. Ar~ hoki ‘tahi kei te meaning males result from unfertilised eggs. The sex ratio KÇpakatanga ki te Tonga. of offspring from mated females is female-biased. E rima, e ono r~nei Ç r~tou tã~tipu mai i te wh~nautanga As generalist predators, most raphignathoids show ~, mate noa. Kotahi ki te toru wiki e whanake ana mai i te some degrees of prey preference. In addition to mites and hua ki te hanga pakeke. He paku tere ake te pakari haere o small insects, they also feed on pollen, and can develop ng~ toa i ng~ uwha. MÇ te w~hi ki ng~ uri ka puta ake, ki and reproduce on various kinds of pollens. te whakatÇkia te hua, he uwha te putanga, ki te kore e Taxonomic studies on this superfamily in New Zealand whakatÇkia, he toa te putanga. Heoi, i roto i ng~ uri o ng~ were initiated by Wood in the mid 1960s. Subsequently, he uwha ka aitia, he maha ake ng~ uwha. published a series of studies on Stigmaeidae (1966, 1967, O ng~ hanga konihi, kei t‘n~ momo t~na tino kai, kei 1968, 1970, 1971b, 1971c, 1981). Luxton (1973) described t‘n~ anÇ t~na. I tua atu i te kai pãwereriki me ‘r~ atu three new species of the Cryptognathidae, and Bolland pepeke iti, ko te hae anÇ t‘tahi o ~ r~tou kai, ~, t‘r~ ka tipu (1991) described a new species of the . Fan haere, ka whakaputa uri i runga i ng~ momo hae huhua. & Zhang recently published two more articles on I t§mataria e Wood ng~ rangahau whakarÇpã i t‘nei Stigmaeidae (2002a, 2002b). wh~nau nui i Aotearoa i ng~ tau waenga o ng~ 1960. I muri

(continued overleaf) (haere tonu) 6 Fan & Zhang (2005): Raphignathoidea (Acari: Prostigmata)

In this contribution the superfamily mai, ka puta i a ia t‘tahi raupapa rangahau mÇ ng~i Raphignathoidea (Acari: Prostigmata) is comprehensively Stigmaeidae (1966, 1967, 1968, 1970, 1971b, 1971c, revised. Keys to world families and genera are provided. 1981). N~ Luxton (1973) i whakaahua ~-kupu ‘tahi momo The , biology, and ecology of world hou e toru o ng~i Cryptognathidae, n~ Bolland (1991) i Raphignathoidea are briefly reviewed so the 76 species, whakaahua t‘tahi momo Camerobiidae. I n~ tata nei, ka including 21 new species, now recognised from New whakaputaina e Fan r~ua ko Zhang ‘tahi atu tuhinga e rua Zealand can be placed in context. The species belong to 20 mÇ ng~i Stigmaeidae (2002a, 2002b). genera (including 1 new ) and 5 families. All species I t‘nei o ng~ putanga, ka ~ta whiriwhiria anÇ te wh~nau are diagnosed, keyed, described, and illustrated, and notes nui Raphignathoidea (Acari: Prostigmata). Ka takoto hoki are provided on the taxonomic references, habitats, and he ara tautohu mÇ ng~ wh~nau me ng~ puninga o te ao. Ka distribution of each species. tirohia anÇ te whakarÇpãtanga, te koiora, me te taupuhi kaiao o ng~i Raphignathoidea puta noa i te ao, kia noho ai ng~ momo e 76 o Aotearoa, tae atu ki ng~ momo hou e 21 kua kitea, ki tÇna horopaki e tika ana. NÇ ‘tahi puninga e 20 (ko t‘tahi he puninga hou) me ‘tahi wh~nau e rima ng~ momo nei. Katoa ng~ momo kua ~ta tohua Ç r~tou ~huatanga matua e rerek‘ ai t‘tahi i t‘tahi, kua tuhia he ara tautohu, kua whakaahuatia ~-kupu, ~-pikitia, kua tuhia anÇ he kÇrero mÇ ng~ tohutoro whakarÇpã, ng~ k~inga noho, me te t§taringa o t‘n~ momo, o t‘n~.

I wh~nau mai a Qing-Hai Fan i Haina ki te Raki, ka kuraina ki Haina ki te Tonga, me te whiwhi i tana T~kutanga m~tai pepeke i te Whare W~nanga Ahuwhenua Fujian i te tau 1996. Mai i te tau 1985 ki te 2001, ka noho ia ki ng~ tãranga o te pãkenga ~whina, te pãkenga, me te pãkenga tÇrua i taua Whare W~nanga anÇ. Mai i te tau 2002, he Contributor Qing-Hai Fan was born in North China and ahorangi m~tai pepeke ia i te Whare W~nanga Ahuwhenua, educated in South China, graduating with a PhD in ento- Whakatipu R~kau Fujian. Ko te Tau~rai Tipu, te M~tai mology from Fujian Agricultural University in 1996. From Pepeke Ahuwhenua, te M~tai Pepeke Noho T~one me te 1985 to 2001 he served as an assistant lecturer, lecturer, M~tai Pãwereriki ‘tahi o ng~ kaupapa kua whakaakona e and associate professor in Fujian Agricultural University. ia. I te tau 2001 me te 2002, i a ia e toro ana i te Whare He has been a professor of entomology at Fujian Agricul- W~nanga o Queensland i Ahitereiria, ka mahi tahi r~ua ko tural and Forestry University since 2002. He has taught T~kuta David E. Walter ki te tirotiro i ng~ pãwereriki o courses including Plant Quarantine, Agricultural Entomol- Ahitereiria. I te tau 2003 ka rere mai ki Aotearoa, ka ogy, Urban Entomology, and Acarology. From 2001 to rangahau i ng~ pãwereriki ‘p~taka porotaka’ i te taha o 2002, as a visiting scientist in Queensland University, T~kuta Zhi-Qiang Zhang, i raro i te maru o Manaaki Australia, he worked on Australian mites with Dr David Whenua. K~tahi ia ka mahi hei kairangahau i Te Kunenga E. Walter. He came to New Zealand in 2003 to study bulb ki Pãrehuroa, he ~ta tirotiro t~na i te pãwereriki Var roa e mites with Dr Zhi-Qiang Zhang as an acarologist in patupatu ana i ng~ p§-miere. Ko ia te •tita Waihanga o Landcare Research, and then worked on the devastating Systematic and Applied Acarology. He nui ake i te 50 ng~ honeybee , Var roa mite, as a research associate at tuhinga hautaka kua oti i a ia e p~ ana ki ng~ whakapapa, Massey University. He is the Production Editor of Sys- te koiora, me te here i ng~ pãwereriki me ‘tahi atu pepeke. tematic & Applied Acarology. He has written more than Kua whakaputaina e r~ua ko T~kuta Zhi-Qiang Zhang 50 journal papers on the systematics, biology, and con- t‘tahi pukapuka e p~ ana ki ng~ pãwereriki p~taka porotaka trol of mites and insects. He published a book on the o Ahitereiria me Te Moana-nui-a-Kiwa. Ko ng~ kaupapa Australasia and Oceania bulb mites in collaboration with e ng~kau nuitia ana e ia, ko ng~ whakapapa pãwereriki (me Dr Zhi-Qiang Zhang. His main interests are the systemat- tino kÇrero i konei ko ‘ r~ o ng~ wh~nau nui ics of mites (especially the superfamilies Raphignathoidea, Raphignathoidea, , me Acaroidea), me te Tetranychoidea, and Acaroidea) and pest management. here i ng~ rauropi kino. (continued overleaf) (haere tonu) Fauna of New Zealand 52 7

I wh~nau mai a Zhi-Qiang Zhang i Shanghai, i Haina, ka whai i te m~tauranga i te Whare W~nanga Fudan (Shang- hai). NÇ te tau 1985 ka whiwhi ia i tana Tohu Paetahi, ko te M~tauranga Kararehe te kaupapa. Ka t§mata tana rangahau i ng~ whakapapa me te koiora pãwereriki i te Kura Paerua, i te Whare W~nanga Fudan, i te tau 1985, ~, ka haere tonu ana akoranga paerua mai i te 1988 ki te 1992 i te Whare W~nanga o Cornell, i Ithaca, Te }poro Nui. I reira ka riro i a ia tana T~kutatanga m~tai pepeke, ko te kaupapa wh~iti, ko te taupuhi kaiao o ng~ pãwereriki konihi me ng~ hanga ka kainga e r~tou. Mai i te tau 1992 ki te 1994, ka mahi ia hei kaim~tai pepeke taupuhi kaiao i te Whare W~nanga o Oregon, i Corvallis, Oregon, i runga i t‘tahi kaupapa here ~-koiora i te tarutaru, he mea whakataki n~ T~kuta Peter McEvoy. Mai i te tau 1994 ki te 1999, he kaim~tai pãwereriki ia m~ te CAB Pãtahi M~tai Pepeke o te Ao, i te Whare Pupuri Taonga o te Ao Tãroa, i R~nana. I a ia e mahi ana m~ CAB International, ko ia anÇ te }piha Hangarau o te BioNET-INTERNATIONAL mai i te 1998 ki te 1999. I te tau 1999, ka neke ia ki Aotearoa, ~, mai i t‘r~ w~, ko ia te kaim~tai pãwereriki o Manaaki Whenua, e whakapau kaha ana ki ng~ whakapapa me te koiora Contributor Zhi-Qiang Zhang was born in Shanghai, pãwereriki. China and educated at Fudan University (Shanghai), gradu- He paewai rangahau hÇnore a T~kuta Zhang i te Whare ating in 1985 with a BSc in Zoology. He began his studies Pupuri Taonga o te Ao Tãroa, he ahorangi turuki ia i te on mite systematics and biology at the Graduate School, Whare W~nanga Fudan, he ahorangi hÇnore anÇ ia i te Fudan University, in 1985, and then continued his post- Kura Tiketike Fujian mÇ ng~ M~tauranga Ahuwhenua graduate studies between 1988 and 1992 at Cornell Uni- (Fujian, China). In~ k ‘ te maha o ng~ tuhinga aronga wh~iti versity, Ithaca, New York, where he received his PhD in kua puta i a ia e p~ ana ki te pãwereriki, he neke atu i te entomology for research on mite predator-prey ecology. 100 ana tuhinga, he mea arotake e t‘tahi atu, e p~ ana ki Between 1992 and 1994 he worked as a postdoctoral ng~ whakapapa, te taupuhi kaiao me te here i ng~ hanga insect ecologist at Oregon State University, Corvallis, Or- kino o te ao angawaho. Ko ia te ‘tita, me t‘tahi o ng~ egon, on a biological weed control project headed by Dr mema o ng~ poari ‘tita o te maha atu o ng~ hautaka o te ao Peter McEvoy. From 1994 to 1999 he was the acarologist e aro wh~iti ana ki te m~tai pãwereriki, te m~tai pepeke, with CAB International Institute of Entomology based in me te m~tauranga kararehe. Ko ia anÇ te Perehitene o te the Natural History Museum in London. While employed Systematic & Applied Acarology Society, ~, kei runga ia i at CAB International he also served as a Technical Officer te Komiti Wh~iti o te Whakarauikatanga M~tai Pãwereriki of the BioNET-INTERNATIONAL from 1998 to 1999. o te Ao. In 1999, he moved to New Zealand and has since been the acarologist for Landcare Research, working on mite sys- Translation by H. Jacob tematics and biology. Dr Zhang holds an honorary research fellowship at the Natural History Museum, an adjunct professorship at Fudan University, and an honorary professorship at Fujian Academy of Agricultural Sciences (Fujian, China). He has published several monographs on mites and more than 100 refereed papers on systematics, ecology, and pest management. He is the editor and an editorial board member of several international journals of acarology, entomology, and zoology. He is the President of the Systematic & Applied Acarology Society and is also on the Executive Committee of the International Congress of Acarology. 8 Fan & Zhang (2005): Raphignathoidea (Acari: Prostigmata) Fauna of New Zealand 52 9

ABSTRACT

The mite superfamily Raphignathoidea (Acari: Prostigmata) is comprehensively revised. Keys to world families and genera of Raphignathoidea are included. The taxonomy, biology, and ecology of world Raphignathoidea are briefly reviewed. 76 species belonging to 20 genera and 5 families recognised as occurring in New Zealand, are diagnosed, keyed, and described. Known stages (if specimens avail- able) of New Zealand raphignathoid species are described and illustrated with line drawings, and notes are provided on the taxonomic references, habitats, and distribution of each species. The following 21 species are described as new: Tycherobius aotearoa, Mecognatha parilis, Mecognatha rara, Raphignathus atomatus, Raphignathus crustus, Agistemus mecotrichus, Eustigmaeus eburneus, Eustigmaeus edentatus, Eustigmaeus ptilosetus, Mediolata delicata, Mediolata polylocularis, Mediolata whenua, Mediolata woodi, Mediolata xerxes, Mediolata zonaria, Mullederia procurrens, Mullederia scutellaris, Pseudostigmaeus schizopeltatus, Storchia hendersonae, Zetzellia biscutata, and Zetzellia spiculosa. A new genus, Scutastigmaeus gen. n., is described. The following 3 new combinations are proposed for three species that were previously placed in Stigmaeus: Scutastigmaeus confusus (Wood), Scutastigmaeus longisetis (Wood), and Scutastigmaeus montanus (Wood).

Keywords: Acari, Prostigmata, Raphignathoidea, taxonomy, keys, New Zealand.

Fan, Qing-Hai; Zhang, Zhi-Qiang 2005. Raphignathoidea (Acari: Prostigmata). Fauna of New Zealand 52, 400 pp.

Received: 2 September 2003. Accepted: 1 March 2004.

CONTENTS Checklist of taxa ...... 9 Acknowledgments ...... 10 Genus Tycherobius Bolland ...... 25 Introduction ...... 11 Tycherobius aotearoa sp. n...... 25 Cryptognathidae Oudemans ...... 25 Characters, biology, techniques and conventions ...... 12 Genus Cryptognathus Kramer ...... 26 Descriptions ...... 21 Cryptognathus striatus Luxton ...... 26 References ...... 111 Cryptognathus vulgaris Luxton ...... 27 Appendices ...... 118 Genus Favognathus Luxton ...... 28 Favognathus leopardus Luxton ...... 28 Illustrations ...... 126 Family Mecognathidae Gerson & Walter ...... 29 Distribution maps ...... 376 Genus Mecognatha Wood ...... 29 Taxonomic index ...... 385 Mecognatha hirsuta Wood ...... 30 Habitat and host index ...... 391 Mecognatha parilis sp. n...... 31 Mecognatha rara sp. n...... 32 Family Raphignathidae Kramer ...... 33 Genus Raphignathus Dugés ...... 34 CHECKLIST OF TAXA Raphignathus atomatus sp. n...... 34 Superfamily RAPHIGNATHOIDEA ...... 21 Raphignathus collegiatus Atyeo, Baker & Family Camerobiidae Southcott ...... 22 Crossley ...... 35 Genus Neophyllobius Berlese ...... 23 Raphignathus crustus sp. n...... 36 Neophyllobius sturmerwoodi Bolland ...... 23 Raphignathus gracilis (Rack) ...... 37 10 Fan & Zhang (2005): Raphignathoidea (Acari: Prostigmata)

Family Stigmaeidae Oudemans ...... 38 Pseudostigmaeus longisetis Wood ...... 85 Genus Agistemus Summers ...... 40 Pseudostigmaeus schizopeltatus sp. n...... 86 Agistemus collyerae González-Rodríguez ..... 41 Pseudostigmaeus striatus Wood ...... 87 Agistemus longisetus González-Rodríguez ... 42 Genus Scutastigmaeus gen. n...... 88 Agistemus mecotrichus sp. n...... 44 Scutastigmaeus confusus (Wood) ...... 89 Agistemus novazelandicus González-Rodríguez Scutastigmaeus longisetis (Wood) ...... 90 ...... 45 Scutastigmaeus montanus (Wood) ...... 90 Agistemus subreticulatus (Wood) ...... 46 Genus Stigmaeus Koch ...... 91 Genus Cheylostigmaeus Willmann ...... 47 Stigmaeus arboricola Wood ...... 92 Cheylostigmaeus luxtoni Wood ...... 47 Stigmaeus brevisetis Wood ...... 93 Genus Eryngiopus Summers ...... 48 Stigmaeus campbellensis Wood ...... 94 Eryngiopus arboreus Wood ...... 48 Stigmaeus luxtoni Wood ...... 94 Eryngiopus bifidus Wood ...... 50 Stigmaeus novazealandicus Wood ...... 95 Eryngiopus nelsonensis Wood ...... 51 Stigmaeus rotundus Wood ...... 97 Eryngiopus similis Wood ...... 52 Stigmaeus rupicola Wood ...... 97 Genus Eustigmaeus (Berlese) ...... 53 Stigmaeus summersi Wood ...... 98 Eustigmaeus brevisetosus (Wood) ...... 54 Genus Storchia Oudemans ...... 100 Eustigmaeus clavigerus (Wood) ...... 55 Storchia hendersonae sp. n...... 100 Eustigmaeus corticolus (Wood) ...... 56 Storchia robustus (Berlese) ...... 101 Eustigmaeus distinctus (Wood) ...... 57 Genus Summersiella Gonzalez ...... 101 Eustigmaeus dumosus (Wood) ...... 58 Summersiella coprosmae (Wood) ...... 102 Eustigmaeus eburneus sp. n...... 59 Genus Zetzellia Oudemans ...... 103 Eustigmaeus edentatus sp. n...... 60 Zetzellia antipoda Wood ...... 103 Eustigmaeus granulosus (Wood) ...... 61 Zetzellia biscutata sp. n...... 105 Eustigmaeus manapouriensis (Wood) ...... 62 Zetzellia gonzalezi Wood ...... 105 Eustigmaeus mixtus (Wood) ...... 63 Zetzellia maori González-Rodríguez ...... 106 Eustigmaeus ptilosetus sp. n...... 64 Zetzellia oudemansi Wood ...... 108 Eustigmaeus simplex (Wood) ...... 65 Zetzellia spiculosa sp. n...... 110 Genus Ledermuelleriopsis Willmann ...... 66 Ledermuelleriopsis incisa Wood ...... 67 Ledermuelleriopsis spinosa Wood ...... 67 ACKNOWLEDGEMENTS Genus Mediolata Canestrini ...... 68 The senior author thanks the College of Plant Protection, Mediolata brevisetis Wood ...... 69 Fujian Agricultural and Forestry University for providing Mediolata delicata sp. n...... 70 facilities during the early stages of this study, and Dr David Mediolata favulosa Wood ...... 71 E. Walter and the Department of Zoology and Entomol- Mediolata mollis Wood ...... 71 ogy, of The University of Queensland for hosting and pro- Mediolata oleariae Wood ...... 72 viding space and facilities when this paper was finalised. Mediolata polylocularis sp. n...... 73 He is also grateful to Dr Eddie A. Ueckermann (Plant Pro- Mediolata robusta González-Rodríguez ...... 73 tection Research Institute, South Africa) for sending litera- Mediolata simplex Wood ...... 75 ture, Dr Ricardo Palma (Museum of New Zealand Te Papa Mediolata whenua sp. n...... 76 Tongarewa, Wellington, New Zealand), and Dr Hans R. Mediolata woodi sp. n...... 76 Bolland (Department of Pure and Applied Ecology, Uni- Mediolata xerxes sp. n...... 77 versity of Amsterdam, The Netherlands) for loaning speci- Mediolata zonaria sp. n...... 78 mens. Rosa Henderson (Landcare Research, Auckland) Genus Mullederia Wood ...... 79 prepared some specimens and typed the list of specimens, Mullederia arborea Wood ...... 79 and Birgit Rhode (Landcare Research, Auckland) prepared Mullederia procurrens sp. n...... 80 the maps. We thank Dr Uri Gerson (Hebrew University of Mullederia scutellaris sp. n...... 80 Jerusalem), Dr David E. Walter, and Dr Eddie A. Genus Primagistemus Fan & Zhang ...... 81 Ueckermann for review and criticism of the manuscript. Primagistemus loadmani (Wood) ...... 81 This project was funded in part by a contract from the Genus Pseudostigmaeus Wood ...... 82 Foundation for Research, Science and Technology, New Pseudostigmaeus collyerae Wood ...... 83 Zealand (Contract C09X0202). Fauna of New Zealand 52 11

INTRODUCTION new family, Homocaligidae, from elements previously placed in Stigmaeidae. Gerson (1972b) synonymised the Historical review Neophyllobiidae with Camerobiidae. Robaux (1975) erected The Raphignathoidea is one of the old groups of the Acari, a new family, Barbutiidae, from elements previously placed which can be dated back at least 56.5 million years. A in Stigmaeidae. Gonzalez (1978) erected a new family, camerobiid mite, Neophyllobius succineus Bolland & Xenocaligonellididae (= Xenocaligonellidae), which was Magowski, was discovered in Baltic amber of the Upper previously included in Caligonellidae. Krantz (1978) (Bolland & Magowski 1990). The first existent included Pomerantziidae in the superfamily again, but his species, Stigmaeus siculus Berlese, 1883 (= rubens opinion has not been followed by most acarologists. Gerson Schrank 1781) was described two centuries ago. & Walter (1998) added the tenth family and provided a key Raphignathoidea belongs to the mite superorder to the families. More recently, a new family, Dasythyreidae Acariformes, order Prostigmata. The superfamily was was erected by Walter & Gerson (1998). Thus the total proposed by Grandjean (1944) to accommodate three number of families became eleven (Table 1) and the families: Raphignathidae Kramer, 1877, Stigmaeidae superfamily currently consists of about 770 valid species Oudemans, 1931, and Caligonellidae Grandjean, 1944. in 57 genera. Baker & Wharton (1952) placed the Stigmaeidae and Modern research on the superfamily was initiated by Caligonellidae into synonymy with the Raphignathidae. Summers and his students (1957–1966) and Wood (1964– Cunliffe (1955) first characterised the superfamily based 1981). A series of reviews/revisions at the level of the on an analysis of leg tarsi, gnathosoma, genitalia, palps, genus or family, especially on the family Stigmaeidae, were and dorsal idiosomal setation, and then added published. Cryptognathidae Oudemans, 1902 and Pomerantziidae In New Zealand, the taxonomic studies on this Baker, 1949 to the superfamily; he also mentioned that the superfamily started in the mid 1960s (Wood 1964a, 1964b). latter might belong elsewhere. Southcott (1957) emendated Wood subsequently published a series of studies on the definition of the superfamily and added two families, Stigmaeidae (1966, 1967, 1968, 1970, 1971b, 1971c, 1981). Camerobiidae and Neophyllobiidae. Meyer & Ryke (1960) Luxton (1973) described three new species of the also made emendations to the superfamily. Summers Cryptognathidae and Bolland (1991) described a new (1966a) included three families, Eupalopsellidae Willmann, species of the Camerobiidae. Two more articles on 1952, Camerobiidae Southcott, 1957, and Neophyllobiidae Stigmaeidae were recently published by Fan & Zhang Southcott, 1957, excluded Pomerantziidae and gave the (2002a, 2002b). first practical key to the families. Wood (1969) erected a

Table 1. Systematic systems of Raphignathoidea by authors. * = Walter & Gerson (1998).

Family Grandjean Cunliffe Summers Krantz Gerson & Walter (1944) (1955) (1966a) (1978) (1998)

Raphignathidae + + + + + Caligonellidae + + + + + Stigmaeidae + + + + + Cryptognathidae + + + + Eupalopsellidae + + + Camerobiidae + + + Homocaligidae + + Barbutiidae + + Xenocaligonellididae + Mecognathidae + Dasythyreidae +* 12 Fan & Zhang (2005): Raphignathoidea (Acari: Prostigmata)

MORPHOLOGICAL CHARACTERS Stigmaeidae), sometimes with more than 2 pairs (neotrichy, Xenocaligonellididae, one genus of Dasythyreidae, and one ADULT FEMALE species of Camerobiidae); f-series with 1 pair of setae, Gnathosoma (Fig. 1–3). Projecting in front of prodosoma, sometimes with 2 or more pairs (in Camerobiidae, or covered by prodosoma (Camerobiidae), rarely retract- Dasythyreidae, and Xenocaligonellididae); h-series with 2 able (Cryptognathidae). Chelicerae (Fig. 1) basally fused, pairs of setae, sometimes with 3 pairs (Raphignathidae, separate (most genera of Stigmaeidae) or conjunct some genera of Calignonellidae, and Stigmaeidae) or more (Homocaligidae, some genera of Stigmaeidae); conical, rarely pairs (one genus of Dasythyreidae). stumpy (Camerobiidae); peritreme present, or absent Ventral idiosoma (Fig. 5). Coxae II and III separate, (Eupalopsellidae, Homocaligidae, Mecognathidae, and sometimes contiguous (Cryptognathidae and Stigmaeidae). Palps (Fig. 2) stout, or slender Raphignathidae); ventral setae 1a and 3a present, 4a present (Eupalopsellidae and Mecognathidae); tibial claws promi- or absent (Xenocaligonellididae, a few Stigmaeidae); ventral nent, or reduced (Raphignathidae), vestigial, or absent opisthosoma with 1–5 pairs of aggenital setae; genital and (Cryptognathidae, Eupalopsellidae, Mecognathidae, and anal valves separate (Barbutiidae, Calignonellidae, Xenocaligonellididae); palptarsus commonly with 4 Cryptognathidae, and Raphignathidae), contiguous eupathidia, 3 of them (ul’ζ, ul”ζ, and sulζ) may be basally (Camerobiidae, Dasythyreidae, Xenocaligonellididae, and fused (Eupalopsellidae, Homocaligidae, Mecognathidae, a few genera of Stigmaeidae), or fused (Eupalopsellidae, and Stigmaeidae); counts of setae (excluding solenidia and Homocaligidae, Mecognathidae, and most genera of eupathidia) from palpcoxa to palptarsus: 1elcp, 0, 1–3, 1– Stigmaeidae), with 1–3 pairs of genital setae, and commonly 2, 3 + 0–1claw, 4 (1–3 in Barbutiidae, Camerobiidae, and with 3 pairs of pseudanal setae, rarely with 1 or 2 pairs (a some genera of Calignonellidae). (Fig. 3) few species of Caligonellidae and Camerobiidae); genital stumpy, sometimes basally elongate (Cryptognathidae) or and anal opening longitudinal; genital folds present. terminally elongate (Eupalopsellidae and Mecognathidae), Leg. Tarsal claws present, sometimes absent (some genera with 2 pairs of rostral setae, and 1or 2 pairs of subcapitular of Stigmaeidae), rarely with tenent hairs (in Barbutiidae); setae (Eupalopsellidae, Homocaligidae, Mecognathidae, empodium with tenent hairs directly arising from axis (Fig. Raphignathidae, most genera of Stigmaeidae, and some gen- 12, 40) or from shafts (Homocaligidae, Mecognathidae, era of Calignonellidae). and Stigmaeidae, Fig. 30, 48) or from vestigial axis Idiosoma (Fig. 4–5). Oval or round in dorsoventral view. (Eupalopsellidae); tarsal stalk sometimes prominent Prodorsum with 2 pairs of vertical setae, rarely with 3 or (Camerobiidae and Dasythyreidae); counts of solenidia on more pairs (neotrichy, some of Camerobiidae and one ge- genua I–III: 1, 0–1, 0; on tibiae I–III: 0–3, 0–2, 0–1; on tarsi nus of Dasythyreidae); with 2 pairs of scapular setae, I–III: 1–2, 1–2, 0–1; counts of setae on legs I–IV: coxae sometimes with only 1 pair (some of Mecognathidae and (including 1a, 3a, and 4a) 2–3 + 1elcp, 0–2, 1–3, 1–3; Stigmaeidae), rarely with 3 or more pairs (neotrichy, one trochanters 0–1, 0–1, 0–2, 0–1; femora 2–6, 1–6, 1–4, 1–4; genus of Dasythyreidae); setae pdx (neotrichy) only present genua 1–5, 0–5, 0–4, 0–4; tibiae 3–9, 2–8, 2–8, 2–7; tarsi in Dasythyreidae and some genera of Camerobiidae; eyes 7–23, 6–21, 5–13, 1–13. present, sometimes absent (some Calignonellidae and Stigmaeidae); postocular bodies (pob) present, sometimes ADULT MALE absent (some Calignonellidae and Stigmaeidae). Dorsal Similar to adult female, but differs in: hysterosoma often hysterosoma with 5 series of dorsal idiosomal setae: c, d, e, somewhat tapered; first and second pseudanal setae often f, and h (pseudanal setae ps are associated with the anal reduced; genital and anal openings fused; having aedeagus; 1–3 ω ω opening and often ventrally located); c-series commonly solenidia 2 (= male [male], Fig. 22) on tarsi I–II absent with 2 pairs of setae, rarely with 1 pair (one genus of (Barbutiidae, Calignonellidae, Camerobiidae, Mecognathidae and a few genera of Stigmaeidae), some- Cryptognathidae, Dasythyreidae, Raphignathidae, and times with 3 or more pairs (neotrichy, Dasythyreidae and Xenocaligonellididae) or present (Eupalopsellidae, Xenocaligonellididae); d-series with 1 pair of setae, or with Homocaligidae, Mecognathidae, and Stigmaeidae); solenidia ω ω 2 pairs (Barbutiidae, Camerobiidae, Eupalopsellidae, or 1 enlarged. Homocaligidae, Mecognathidae, and most genera of Stigmaeidae), sometimes with more than 2 pairs (neotrichy, TRITONYMPH Dasythyreidae, Xenocaligonellididae, and one species of Only members of Raphignathidae and Xenocaligonellididae Camerobiidae); e-series with 1 pair of setae, or 2 pairs are known to have this stage. It can be separated from the (Barbutiidae, Cryptognathidae, Camerobiidae, adult female by the absence of genital valves and folds, and Eupalopsellidae, Homocaligidae, Mecognathidae, and the presence of 1 pair of genital setae in the female. Fauna of New Zealand 52 13

DEUTONYMPH Table 3. Prey associations of genera of Raphignathoidea Similar to adult but without genital setae in both sexes; (* = Unpublished data). absence of genital folds in female and aedeagus in male. Predators Prey

PROTONYMPH Caligonellidae Molothrognathus Acari: Prostigmata: With 1 pair of subcapitular setae; ventral setae 4a and Tetranychidae genital setae absent; with fewer setae in aggenital area and Paraneognathus Acari: Astigmata: * on segments of legs than deutonymph. Prostigmata: * Camerobiidae LARVA Neophyllobius Acari: Prostigmata: Subcapitular setae, ventral setae 4a, genital, and aggenital * Insecta: Homoptera: Diaspididae, setae absent; without leg IV; with fewer setae on segments Margarodidae of palps and legs than protonymph; leg I with 1 (tc’ ) or 2 Eupalopsellidae (tc’ and tc’’) tactile setae (Calignonellidae, Cryptognathidae, Eupalopsellus Acari: Prostigmata: Eupalopsellidae, Homocaligidae, Mecognathidae, Tetranychidae, Raphignathidae, and Stigmaeidae), or without tactile seta Insecta: Homoptera: Diaspididae Eupalopsis Insecta: Homoptera: Diaspididae (Camerobiidae, Dasythyreidae, and Xenocaligonellididae). Exothorhis Insecta: Homoptera: Diaspididae Saniosulus Insecta: Homoptera: Diaspididae, Phoenicococcidae (Coccoidea) DEVELOPMENTAL STAGES Stigmaeidae Agistemus Acari: : Generally, there are 5 known stages in most members of Raphignathoidea: the egg, larva, protonymph, deutonymph, Prostigmata: , and adult. Only one species, Agistemus exsertus Gonzalez, Stigmaeidae, Tarsonemidae, has a known prelarval stage (Hanna et al. 1984). Mites of Tenuipalpidae, Tetranychidae, the genus Raphignathus have 3 nymphal stages: Insecta: Homoptera: protonymph, deutonymph, and tritonymph (Meyer & Aleyrodidae, Coccoidea Ueckermann 1989, and authors’ unpublished observation). Lepidoptera: Pyralidae Three nymphal stages are also present in Pollen of plant: Caprifoliaceae, Xenocaligonellididae (Fan 2000). Euphorbiaceae, Palmae, Poaceae, Typhaceae Eryngiopus Insecta: Homoptera: Diaspididae Mediolata Acari: Prostigmata: FEEDING HABITS Tetranychidae According to our current knowledge, the majority of the Insecta: Homoptera: Coccoidea Raphignathoidea are free-living predators (Table 2). Some Zetzellia Acari: Astigmata: Saproglyphidae species of Stigmaeidae and Dasythyreidae are found on Prostigmata: Eriophyidae, insects, and a few species of Stigmaeidae are phytophages, Stigmaeidae, Tarsonemidae, feeding on moss. A couple of species of Xenocaligo- Tenuipalpidae, Tetranychidae nellididae are possibly microphytophages, probably feed- Mesostigmata: Phytoseiidae ing on substances on the outer layer of bark. Insecta: Homoptera: Coccoidea Lepidoptera: Pyralidae Table 2. Feeding pattern of raphignathoid mites Family Predators Phytophages Parasites Microphytophages

Barbutiidae ? Caligonellidae + Camerobiidae + Cryptognathidae + Dasythyreidae ? Eupalopsellidae + Homocaligidae ? Mecognathidae ? Raphignathidae + Stigmaeidae + + + Xenocaligonellididae ? 14 Fan & Zhang (2005): Raphignathoidea (Acari: Prostigmata)

Table 4. Insect hosts of Stigmaeidae

Stigmaeid species Host Author

Eustigmaeus dyemkoumai Abonnenc Phlebotomine sandflies Abonnenc 1970 Eustigmaeus gamma (Chaudri) Phlebotomus pius Chaudhri 1965 Eustigmaeus gorgasi (Chaudri) Sandfly Chaudhri 1965 Eustigmaeus johnstoni Zhang & Gerson Phlebotomus longicuspis Nitzulescu, Zhang & Gerson 1995 Phlebotomus papatasi (Scopoli), Sergentomyia africana (Newstead), Sergentomyia dreyfussi (Parrot), Sergentomyia magna (Sinto), Sergentomyia spp. Phlebotomus bergeroti, P. sergenti Shehata & Baker 1996 P. sergenti Ozbel et al. 1999 Eustigmaeus parasiticus (Chaudri) Phlebotomus Chaudhri 1965 Eustigmaeus sp. Phlebotomine sandflies Martinez et al. 1983 Stigmaeus smithi (Mitra & Mitra) Phlebotomus papatasii Mitra & Mitra 1953 Stigmaeus sinai Swift Phlebotomus papatasii Swift 1987 Stigmaeus youngi (Hirst) “Flies” (probably sandflies) Hirst 1926; Wood 1972

Table 5. Microhabitats of raphignathoid mites

Foliage and Moss and Litter and nest branches Trunk lichen soil or house

Barbutiidae + Caligonellidae + + + + + Camerobiidae + + + + Cryptognathidae + + + + + Dasythyreidae + + Eupalopsellidae + + + + Homocaligidae + + Mecognathidae + + + + Raphignathidae + + + + + Stigmaeidae + + + + + Xenocaligonellididae + +

Predators. Relatively little is known about the predatory (Table 4). Whether these species have any negative effects habit of the Raphignathoidea (Table 3). Species of the two on insect hosts is yet to be shown, although feeding scars genera of Caligonellidae, Molothrognathus and were commonly observed and the nature of the mite-insect Paraneognathus, are known to prey on spider mites on relationship was assumed to be by most au- and on acarid mites and cheyletid mites in stored thors (Table 4). A species of Dasythyreus was found on the products. Species of the Camerobiidae are known to feed pronotal-mesonotal interface of the eyed-click beetle Alaus on crawlers of the scale insects and tarsonemid mites (Ta- myops (F.) and it is not known whether this mite feeds on ble 3). Members of Eupalopsellidae are mainly predators the beetle (Walter & Gerson 1998). of mites and scale insects (Table 3). Species of Stigmaeidae Phytophages and microphytophages. Some members of feed on spider mites and other mites, as well as on small Eustigmaeus (Stigmaeidae) were observed feeding on insects such as crawlers of scale insects (Table 3). mosses (Gerson 1972a). Xenocaligonellidus ovaerialis De Leon (Xenocalignoellididae) was observed probing a tree Parasites. Five species of Eustigmaeus, three species of bark surface (De Leon 1959). Another species, Stigmaeus (both Stigmaeidae), and one species of Xenocaligonellidus smileyi Hu & Liang, was observed Dasythyreus (Dasythyreidae) have been found on insects. scrubbing the substance on the outer layer of oliver tree Stigmaeids have so far been recorded on sandflies only bark (Fan 2000). Fauna of New Zealand 52 15

MICROHABITATS female/day at 25°C when feeding on eggs of Panonychus citri (Yue & Childers 1994) and 0.150 individuals/female/ Members of the superfamily are found on foliage, branches, day at 27–29°C when feeding on trunks (tree bark and holes), moss and lichen, litter, soil, (Abou-Awad & Elsawi 1993), and that of Z. mail is 0.109 animal nests (birds, possum, and honeybee), stored prod- female offspring/female/day at 24±1°C when feeding on ucts, and house dust (Table 5). A few are aquatic or sub- Aculus schlechtendali (White & Laing 1977). aquatic: Annerosella and Homocaligus of Homocaligidae; Adult females of Agistemus industani live about as and some species of Caligohomus, Cheylostigmaeus, long as adult males but consume three times as many prey Eustigmaeus, and Ledermuelleriopsis of Stigmaeidae. as males (Arbabi & Singh 2002). Diapause. There appears to be a lack of diapause in BIOLOGY raphignathoid mites, at least for the limited number of Compared with the Phytoseiidae, relatively little is known studies reported so far. Eustigmaeus frigidus apparently about the biology of the Raphignathoidea. Published records reproduces under both long-day (16 h) and short-day (9 h) concern a handful of species from two main families, photoperiodic regimes, without a reproductive diapause Stigmaeidae and Eupalopsellidae. The biology of the (Gerson 1972a). In Auckland, Agistemus longisetus breeds Stigmaeidae was reviewed by Santos & Laing (1985) in throughout the year on non-deciduous plants, without an relation to their role as predators of Tetranychidae. overwintering phase (Collyer 1964). Thistlewood et al. (1996) discussed the biology of Stigmaeidae in relation to their role as predators of the Feeding behaviour and . The biology and application of Stigmaeidae, Unlike the phytoseiids, which can respond to kairomones Camerobiidae, and Eupalopsellidae in biological control associated with prey, Zetzellia mali does not detect were reviewed by Gerson & Smiley (1990) and updated kairomones and appears to search for prey by random by Gerson et al. (2003). encounters (Santos 1991). Once inside a prey patch or Life history. The life cycle has been studied for only a few , Z. mali increases its residence time in response to the species: Agistemus exsertus, A. floridanus, A. industani, presence of prey, but it also a patch before all prey and Zetzellia mali. In general, development from egg to are consumed (Lawson & Walde 1993). Saniosulus nudus adult can be completed in 1–3 weeks, although the dura- holds its prey by its anterior legs while inserting the cheli- tion is affected by abiotic factors such as temperature and cerae into the body of the prey; it then sucks the body biotic factors such as the type and quantity of food (Collyer fluids for 30–40 min or more and when finished pushes the 1964; ElBadry et al. 1969a; Gerson & Blumberg 1969; shrivelled prey off the chelicerae with its long palps (Gerson Muma & Selhime 1971; Gerson 1972a; Inoue & Tanake & Blumberg 1969). Agistemus exsertus punctures 1983; Osman & Zaki 1986; Yue & Childers 1994; White & tetranychid eggs but does not necessarily suck their con- Laing 1977; El-Laithy 1998; Jamali et al. 2001; Arbabi & tents completely (ElBadry et al. 1969b). Singh 2002). The egg stage is invariably the longest among As generalist predators, most stigmaeids show some immature stages and often takes at least twice as long as degrees of prey preference. Zetzellia mali tends to prefer the eriophyids over the economically more significant the larval or each nymphal stage. The males develop slightly tetranychids (Santos 1976a; Clements & Harmsen 1993; faster than females. Walde et al. 1995). Agistemus exsertus prefers immatures Reproduction is arrhenotokous and the offspring sex of Tenuipalpus granati to those of Tetranychus urticae ratio of mated females is female-biased (Gerson 1972a; (Yousef et al. 1982), and Tetranychus cinnabarinus to Arbabi & Singh 2002). Unmated females produce males Eutetranychus orientalis (ElBadry et al. 1969b). When only and start to lay eggs one day later than mated females feeding on tetranychids, Zetzellia mali prefers eggs and (Rasmy & Hussein 1996). Multiple-mated females have only occasionally attacks resting and nymphal stages; this shorter life spans but consume more prey and lay more species never attacks adult spider mites (Santos 1976b; eggs than single-mated females (Abou-Awad & Reda 1992; Clements & Harmsen 1990). Agistemus exsertus also Rasmy & Hussein 1995). After a pre-ovipositional period develops faster and produces more eggs when feeding on of a few days, most females start to lay eggs for 1–2 weeks. eggs than it doeswhen feeding on larvae or nymphs of Female reproductive rates (mostly between 1 to 4 eggs per Tetranychus urticae and T. cucurbitacearum (Hafez et al. day) are strongly affected by food type/quantity (ElBadry 1983); it also develops faster when feeding on the eggs of et al. 1969a; Yousef et al. 1982; Nawar 1992) and T. urticae than it does on the eggs of T. cucurbitacearum, temperature (Inoue & Tanake 1983). The intrinsic rate of although the eggs of the latter prey were more attractive to increase of Agistemus exsertus is 0.229 female offspring/ the predator. 16 Fan & Zhang (2005): Raphignathoidea (Acari: Prostigmata)

Table 6. Distribution of raphignathoid families and genera according to geographical regions and New Zealand. + = present; * = unpublished data.

Family/genus Palaearctic Nearctic Neotropical Afrotropical Oriental Australian Antarctic New Zealand

Barbutiidae + +* + Barbutia ++ +*+ Caligonellidae + + + + + Caligonella ++ + ++ Coptocheles ++++ Molothrognathus ++ + ++ Neognathus ++ + ++ Paraneognathus +++ Camerobiidae + + + + + + + Bisetulobius + Camerobia +++ Decaphyllobius ++ + ++* Neophyllobius ++ + + ++ + Tillandsobius ++* Tycherobius ++ + + + Cryptognathidae + + + + + + + Cryptognathus ++ + + Favognathus ++ + + ++ + Dasythyreidiae + + Dasythyreus + Xanthodasythyreus + Eupalopsellidae + + + + + Eupalopsellus ++ + +* Eupalopsis +++ Exothorhis ++ + ++ Peltasellus + Saniosulus ++ + ++ Homocaligidae + + + + + Annerossella +++ Homocaligus ++ + Mecognathidae + + + + + Mecognatha ++ Paraeupalopsellus +++ Raphignathidae + + + + + + + + Neoraphignathus + Raphignathus ++ + + ++++ Stigmaeidae + + + + + + + + Agistemus ++ + + ++ + Caligohomus + Cheylostigmaeus ++ + ++* + Eryngiopus ++ + + ++++ Eustigmaeus ++ + + ++ + Ledermulleriopsis ++ + ++ + Macrostigmaeus + Makilingeria + Mediolata ++ + ++ + Mendanaia + Mullederia +++ + Mullederiopsis + Neilstigmaeus + Parastigmaeus + Paravillersia + Pilonychiopus + Fauna of New Zealand 52 17

Table 6 (continued).

Family/genus Palaearctic Nearctic Neotropical Afrotropical Oriental Australian Antarctic New Zealand

Postumius + Primagistemus ++ Prostigmaeus ++ Pseudostigmaeus +++*+ Scutastigmaeus + Stigmaeus ++ + + ++ + Storchia +++++ Summersiella ++ Villersia + Villersiella + Zetzellia ++ + + ++ + Zetzelliopsis + Xenocaligonellididae + + + + Echinopsis + Xenocaligonellidus +++ +

Predation rates vary with a number of biotic and abiotic son than T. pyri, and the interference between these preda- factors. Within a certain range, the number of prey tor species is only occasionally strong enough to affect A. consumed increases with temperature (Afify et al. 1969) schlechtendali population dynamics (Walde et al. 1997). In and prey density (Nawar 1992; Yue & Tsai 1995). the Northern U.S.A., where the phytoseiid mites Agistemus exsertus, for example, can consume 5.8 larvae pyri and occidentalis are com- per day of Tetranychus urticae at a prey density of 7 mon in apple orchards, Z. mali has a stronger impact on M. larvae; very high levels of prey decreased predator occidentalis than on T. pyri only (Croft & MacRae 1993), oviposition and feeding capacity (Nawar 1992). At very because M. occidentalis lays significantly more eggs in the low prey densities, females of Zetzellia mali primary foraging area of adult female Z. mali than T. pyri disproportionately reduce their predation and oviposition does (MacRae & Croft 1996). rates compared with high densities (Santos 1982). This response, as well as its ability to become cannibalistic, Spatial distribution and seasonal fluctuations allows Z. mali to persist on apple leaves when few prey Stigmaeids are unevenly distributed in orchards. The pat- are present. terns of aggregation vary among different predator species and change with the season and population densities of Intraguild predation and competition their prey, competitors and predators (Holdsworth 1972; Stigmaeids feed on and are fed upon by phytoseiids, espe- Hu et al. 1994; Slone & Croft 1998, 2001). Agistemus cially when phytophagous mites are scarce — this may terminalis, for example, was more aggregated in the lower have both positive and negative impacts on their interac- and western portions of the tree than in other portions (Hu tions and their roles in biological control (Clements & et al. 1994). Zetzellia mali multiplied on the fruit-cluster leaves to become more numerous on the outside of the tree Harmsen 1992; Croft & MacRae 1993; Croft 1994; Slone than on watersprouts (Holdsworth 1972). It is unknown & Croft 2001). At low prey densities, stigmaeids are more how these mites move from tree to tree and disperse from effective than phytoseiids because of their higher prefer- orchard to orchard. ence for prey eggs, higher oviposition relative to prey con- Raphignathoid seasonal fluctuations have been studied sumption, and the ability to consume their own eggs, for Zetzellia mali (Rice et al. 1976; Hu et al. 1996), and for whereas at high prey densities the higher maximum preda- Agistemus longisetus in orchards (Collyer 1964). In apple tion rate of phytoseiids gives them a higher efficacy orchards in Massachusetts, Z. mali was present in early (Clements & Harmsen 1992); a combination of stigmaeids spring and increased slowly until reaching peak levels in and phytoseiids has greater efficacy than either alone over autumn (Hu et al. 1996). In apple and plum orchards in a wide range of prey densities. Zetzellia mali is usually less Auckland, Agistemus longisetus first appears in late important than the phytoseiid Typhlodromus pyri in the December or early January, becomes abundant in February direct reduction of the population growth rate of the and sometimes reaches densities as high as 100 mites per eriophyid Aculus schlechtendali and acts later in the sea- leaf (Collyer 1964). 18 Fan & Zhang (2005): Raphignathoidea (Acari: Prostigmata)

Diet and rearing METHODS AND TECHNIQUES The diets of most raphignathoids are too poorly known to allow rearing. Most stigmaeids that have been studied have Collecting relatively broad ranges of food and are generalist predators Leaves and twigs were examined under a microscope and (Table 3). In addition to mites and small insects, some mites living on them were picked off. Those living in litter, stigmaeids can also develop and reproduce on pollens of moss, nests, soil, tree barks, and stored products were some plants (Abo Elghar et al. 1969; Wafa et al. 1969; extracted by using Berlese funnels. Rasmy 1975; Rasmy et al. 1996). When feeding on the Specimens were stored in 70% ethanol with 3% glycerin pollen of Phoenix dactylifera, Zea mays, and Ricinus com- for light microscope study, or in 95–100% ethanol for munis, Agistemus exsertus does not develop as well as scanning electron microscope study. when feeding on Tetranychus cinnabarinus, but lays more eggs when feeding on the pollen of dates than on T. Preparation for study cinnabarinus (Wafa et al. 1969). This species can also develop normally on artificial diets composed of yeast, Slide mounting. milk, amino acids, and sugar, but the number of eggs laid Mites were cleared in Nesbitt’s fluid (chloral hydrate 40 g, per female per day is two-thirds of that for mites reared on concentrated HCl 2.5 ml, distilled water 25 ml) for a few a natural diet of pollen (Reda 1990). The adult female hours or days depending on the size and sclerotisation of lifespan on the artificial diet is equal to that on the standard specimens. Specimens were mounted in Hoyer’s medium diet. (chloral hydrate 200 g, crystalline gum arabic 30 g, glycerol 20 ml, distilled water 50 ml) or Heinze-PVA (chloral hy- Economic importance and role in biological control drate 100 g, glycerol 10 ml, polyvinyl alcohol 10 g, dis- Raphignathoid mites are important biological control agents tilled water 60 ml, 85–92% lactic acid 35 ml). Slides were of spider mites, eriophyid mites, and scale insects in agri- labelled with the collecting data and then placed in an oven culture and forestry. Most species of the families at 45–50°C for several weeks until the medium was dry. Eupalopsellidae, Stigmaeidae, Caligonellidae, and Detailed methods for mite extracting, preparing and mount- Camerobiidae are free-living predators (Meyer & ing are available in Walter & Proctor (2001). Ueckermann 1989; Gerson & Smiley 1990). Among them the genera Agistemus and Zetzellia of the Stigmaeidae and Illustrations and measurements Saniosulus of the Eupalopsellidae are well-known biologi- Drawings were made in pencil using a camera lucida under cal control agents on plants. Gerson et al. (2003) reviewed a microscope and inked with Rotring Rapidograph Pens. the role and application of Eupalopsellidae and Stigmaeidae Measurements were made from slide-mounted in biological control. specimens using stage-calibrated ocular micrometers. Chelicerae were measured from basal articulations to tips of movable digits. Palps were measured from bases of GEOGRAPHICAL DISTRIBUTION trochanters to tips of palpal tarsi. Idiosomal lengths were measured from the anterior to the posterior margins. Mites of the superfamily are worldwide in distribution, Idiosomal widths were measured from maximum width of and abundant in the Palaearctic, Nearctic, Neotropical, the idiosoma between leg II and III. Setae and solenidia Afrotropical, Oriental, and Australian Regions. were measured from alveoli to tips. Legs were measured Raphignathus johnstoni Womersley was even discovered from bases of trochanters to tips of claws. Femora of leg I in the Antarctic region (Womersley 1937). The were measured from ventral junction between trochanter raphignathoid faunas of the Palaearctic, Nearctic, and femur to junction between femur and genu. Genua I Afrotropical, and Oriental Regions are relatively well were measured from junction between femur and genu to known, but the Neotropical Region has only a few species junction between genu and tibia. Tibiae I were measured recorded or described (Table 6). from junction between genu and tibia to junction between tibia and tarsus. Tarsi I were measured from posterior margins to tips of claws. In the material examined, n/n indicates number of slides/ number of specimens. Measurements x (y–z): x is the measurement of the specimens (mostly are holotype or paratype) from which figure was drawn; y–z is the range of measurements. Female or male means adult unless indicated. Fauna of New Zealand 52 19

Terminology. The terminology of palp and leg chaetotaxy f2 2nd pair of setae in 4th series or row on follows that of Grandjean (1944, 1946) and the terminol- hysterosoma ogy of idiosomal chaetotaxy follows Kethley (1990). All h1 innermost (1st) pair of setae in 5th series or row measurements are given in micrometers (µm). on hysterosoma

h2 2nd pair of setae in 5th series or row on hysterosoma

h3 3rd pair of setae in 5th series or row on LIST OF ABBREVIATIONS hysterosoma Gnathosoma ia anterior pair of cupules (lyrifissures) on hysterosoma acc accessory claw of palpal tibia im middle pair of cupules (lyrifissures) on ω solenidion on palptarsus hysterosoma elcp supracoxal setae of palp ip poterior pair of cupules (lyrifissures) on ro 1st or internal pair of rostral setae 1 hysterosoma ro 2nd or external pair of rostral setae 2 ih caudal pair of cupules (lyrifissures) on m anterior or innermost pair of subcapitular setae hysterosoma n posterior pair of subcapitular setae 1a innermost (1st) pair of setae associated with bases of legs I Idiosoma 1b 2nd pair of setae associated with bases of legs I C 1st hypothesised segment of hysterosoma, re- 1c 3rd pair of setae associated with bases of legs I vealed by a row of setae c 2b 2nd pair of setae associated with bases of legs II D 2nd hypothesised segment of hysterosoma, re- 2c 3rd pair of setae associated with bases of legs II vealed by a row of setae d 3a innermost (1st) pair of setae associated with bases E 3rd hypothesised segment of hysterosoma, re- of legs III vealed by a row of setae e 3b 2nd pair of setae associated with bases of legs III F 4th hypothesised segment of hysterosoma, re- 3c 3rd pair of setae associated with bases of legs III vealed by a row of setae f 4a innermost (1st) pair of setae associated with bases H 5th hypothesised segment of hysterosoma, re- of legs IV vealed by a row of setae h 4b 2nd pair of setae associated with bases of legs IV PS 6th hypothesised segment of hysterosoma, re- 4c 3rd pair of setae associated with bases of legs IV vealed by a row of setae ps ag anterior (1st) pair of aggenital setae vi internal pair of vertical setae 1 ag2 2nd pair of aggenital setae ve external pair of vertical setae ag 3rd pair of aggenital setae vx vertical setae (neotrichy) 3 ag4 4th pair of aggenital setae pdx prodorsal setae (neotrichy) ag 5th pair of aggenital setae sci internal pair of scapular setae 5 g1 anterior (1st) pair of genital setae sce external pair of scapular setae g 2nd pair of genital setae pob postocular body 2 g3 3rd pair of aggenital setae c1 innermost (1st) pair of setae in 1st series or row ps1 1st pair of pseudanal setae on hysterosoma ps 2nd pair of pseudanal setae c 2nd pair of setae in 1st series or row on 2 2 ps 3rd pair of pseudanal setae hysterosoma 3 Legs d1 innermost (1st) pair of setae in 2nd series or row on hysterosoma Iω solenidion on tarsus I ω d 2nd pair of setae in 2nd series or row on I 1 anterior (1st) solenidion on tarsus I in male 2 ω hysterosoma I 2 posterior (2nd) solenidion on tarsus I in male ω e1 innermost (1st) pair of setae in 3rd series or row I p proximal solenidion on tarsus I in female on hysterosoma Iϕ solenidion on tibia I ϕ e2 2nd pair of setae in 3rd series or row on I ’ anteriorly located solenidion on tibia I hysterosoma Iϕ” posteriorly located solenidion on tibia I ϕ f1 innermost (1st) pair of setae in 4th series or row I p proximal solenidion on tibia I on hysterosoma Iκ sensillum on genu I 20 Fan & Zhang (2005): Raphignathoidea (Acari: Prostigmata)

dFI dorsalmost seta on femur I dGI dorsalmost seta on genu I elcp supracoxal setae of leg I IIω solenidion on tarsus II ω II 1 anterior (1st) solenidion on tarsus II in male ω II 2 posterior (2nd) solenidion on tarsus II in male IIϕ solenidion on tibia II IIϕ’ anteriorly located solenidion on tibia II IIϕ” posteriorly located solenidion on tibia II IIϕp proximal solenidion on tibia II IIκ sensillum on genu II IIIω solenidion on tarsus III ω III 1 anterior (1st) solenidion on tarsus III in male ω III 2 posterior (2nd) solenidion on tarsus III in male IIIϕp proximal solenidion on tibia III IVω solenidion on tarsus IV IVω anterior (1st) solenidion on tarsus IV in male ω1 IV 2 posterior (2nd) solenidion on tarsus IV in male IVϕp proximal solenidion on tibia IV

Abbreviations for museums and collections BMNH The Natural History Museum, London, U.K. ISZA Istituto Sperimentale per la Zoologia Agraria, Firenze, Italia. MONZ Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand. NZAC New Zealand Arthropod Collection, Landcare Research, Auckland, New Zealand USNM National Museum of Natural History, Smithsonian Institution, Washington DC, USA. ZMH Zoologischen Museum, Hamburg, Germany. Fauna of New Zealand 52 21

Superfamily Raphignathoidea Grandjean — Peritremes absent (Fig. 1 E–H); males with an additional Raphignathoidea Grandjean, 1944: 104. solenidion on tarsi I and II (Fig. 22 A–B); leg empodial axis bearing 1–3 shafts, each of them producing 1 pair Diagnosis. Adult female. Chelicerae basally fused or separate; peritreme present or absent; palptibial claw of tenent hairs (Fig. 30 E), sometimes empodial axis prominent, reduced, or absent, palptarsus commonly with reduced and directly bearing 2 bunches of long tenent 4 eupathidia, eupathidia separated or basally fused, counts hairs ...... 8 of setae (excluding solenidia and eupathidia) from palpcoxa 3 Dorsum with at least 14 pairs of setae in adult (d2 and to palptarsus: 1elcp, 0, 1–3, 1–2, 3 + 0–1 claw, 1–4; f2 present) (Fig. 4 I–K); tarsi I without setal complex subcapitulum with 2 pairs of rostral setae and 1–2 pairs (similar to duplex setae in Tetranychidae) in larva .. 4 of subcapitular setae. Propodosoma commonly with 2 — Dorsum with 11–12 pairs of setae in adult (d2 and f2 series of setae and dorsal hysterosoma with 5 series of absent) (Fig. 4 B–D); tarsi I with setal complex in larva setae; each series often with 2 pairs of setae; eyes and ...... 6 postocular bodies (pob) present or absent. Coxae II and III often separate; ventral opisthosoma with 1–5 pairs of 4 Legs not stilt-like; tibiae and genua subequal; most dorsal aggenital setae; genital and anal openings longitudinal, geni- idiosomal setae in marginal area whip-like; genital valves tal folds present, genital valves with 1–3 pairs of setae with 3 pairs of genital setae (Fig. 5 I–J); palptarsus and anal valves with 3 pairs of pseudanal setae (rarely with 3–4 setae (excluding solenidion and eupathidia) with 1–2 pairs). Tarsal claws present, sometimes absent, (Fig. 2 I–J) ...... 5 rarely with tenent hairs; empodium with tenent hairs; — Legs stilt-like; tibiae at least 3 times length of genua counts of solenidia on genua I–III: 1, 0–1, 0; on tibiae I– (Fig. 7 A–D); dorsal idiosomal setae in marginal area III: 0–3, 0–2, 0–1; on tarsi I–III: 1–2, 1–2, 0–1; counts of not whip-like (Fig. 6 A); genital valves with 1 pair of setae on legs I–IV: coxae (excluding 1a, 3a and 4a) 1–2 + genital setae (Fig. 5 K); palptarsus with 1–2 setae (Fig. 1elcp, 0–2, 1–2, 1–2; trochanters 0–1, 0–1, 0–2, 0–1; 2 K) ...... (p. 22)... Camerobiidae Southcott femora 2–6, 1–6, 1–4, 1–4; genua 1–5, 0–5, 0–4, 0–4; tibiae 3–9, 2–8, 2–8, 2–7; tarsi 7–23, 6–21, 5–13, 1–13. 5 Prodorsal and dorsal hysterosomal shields separate; palptibial claw present (Fig. 2 J); palptarsi mound- Adult male. Similar to adult female but: first and second like, clearly shorter than palptibiae ...... pseudanal setae often reduced; genital and anal openings ω ω ...... Dasythyreidae Walter & Gerson fused; having an aedeagus; solenidia or 1 on legs often enlarged. — Prodorsal and dorsal hysterosomal shields fused; Tritonymph. Only known in Raphignathidae. Similar to palptibial claw absent (Fig. 2 I); palptarsi subequal to adult but without genital folds in female or aedeagus in palptibiae ...... Xenocaligonellididae Gonzalez male. 6 Gnathosoma not retractable (Fig. 1 B, D); chelicerae Deutonymph. Similar to adult but without genital folds fused to the level of bases of stylets (Fig. 1 B, D); and setae in both sexes. neither dorsal nor ventral idiosoma covered by a single Protonymph. With 1 pair of subcapitular setae; ventral shield; prodorsum without a hood-like projection (Fig. setae 4a and genital setae absent; with fewer setae in 33 A) ...... 7 aggenital area and on segments of legs. — Gnathosoma retractable (Fig. 1 C); only the bottom of Larva. Subcapitular setae, ventral setae 4a, genital and chelicerae fused (Fig. 1 C); dorsal and ventral idiosoma aggenital setae absent; without leg IV; with fewer setae on each covered by a single shield; prodorsum forming a segments of palps and legs. hood-like projection (Fig. 13 A–B) ...... (p. 25)... Cryptognathidae Oudemans Key to families of Raphignathoidea 7 Peritremes situated on chelicerae (Fig. 1 B); coxae II and III separate (Fig. 5 B) ...... 1 Leg tarsal claws (if present) nude; palptibial claw (if present) without a ventral tooth (Fig. 2 B–K) ...... 2 ...... Caligonellidae Grandjean — Leg tarsal claws with tenent hairs; palptibial claw with — Peritremes situated between chelicerae and prodorsum a ventral tooth (Fig. 2 A) ...... Barbutiidae Robaux (Fig. 1 D); coxae II and III contiguous (Fig. 5 D) ...... (p. 33)... Raphignathidae Kramer 2 Peritremes present on chelicerae (Fig. 1 B–C, I–K) or between chelicerae and prodorsum (Fig. 1 D); both 8 Cheliceral bases completely fused (Fig. 1 G–H); palps female and male with same number of solenidia on tarsi prominently elongate, palptibial claw small or vestigial, I–II; leg empodial axis directly producing 2 rows of no more than 1/3 length of palptarsus (Fig. 2 G–H) . tenent hairs (Fig. 12 G) ...... 3 ...... 10 22 Fan & Zhang (2005): Raphignathoidea (Acari: Prostigmata)

— Cheliceral bases separate (Fig. 1 E), rarely fused or with 2 pairs of scapular setae; pdx (neotrichy) usually conjunct (Fig. 1 F); palps not elongate, palptibial claw present; eyes present; pob present. Dorsal hysterosoma developed, at least 1/3 length of palptarsus (Fig. 2 E– with 5 series of setae: c, d, e, f, and h (not including F) ...... 9 pseudanal setae which are associated with anal opening); c-series with 2 pairs of setae; d-series with 2 pairs of 9 Prodorsum with a transversal groove leading internally setae, rarely 3 pairs (neotrichy); e-series with 2 pairs, to 1 pair of sacs (female) or tubes (male); suranal and rarely 3 pairs of setae (neotrichy); f-series with 2 pairs of aggenital shields fused ...... Homocaligidae Wood* setae; h-series with 2 pairs of setae. Coxae II and III — Prodorsum without transversal groove, sacs or tubes; narrowly separate; ventral setae 4a present; ventral suranal and aggenital shields separate (Fig. 43 A–B) . opisthosoma with 1 pair of aggenital setae; genital and ...... (p. 38)... Stigmaeidae Oudemans anal valves contiguous, with 1 pair of genital setae and 3 pairs, rarely 2 pairs, of pseudanal setae. Leg tarsal claws 10 Palptarsi elongate, longer than palptibiae (Fig. 2 H); present, nude; empodial axis without shafts, directly pro- leg empodial axis minute, peg-like, bearing 2 bunches ducing 2 rows of tenent hairs; tarsal stalk prominent; of long tenent hairs ...... Eupalopsellidae Willmann coun