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Phylogeny and Classification of Armored Scale Insects (Hemiptera: Coccomorpha: Diaspididae)

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Zootaxa 4616 (1): 001–098 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Monograph ZOOTAXA Copyright © 2019 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4616.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:1B9DFBC9-2BA8-4619-8F70-E372DCBD7411 ZOOTAXA 4616 Phylogeny and classification of armored scale insects (Hemiptera: Coccomorpha: Diaspididae) BENJAMIN B. NORMARK1,2,7, AKIKO OKUSU1, GEOFFREY E. MORSE3, DANIEL A. PETERSON1,2,4, TAKAO ITIOKA5 & SCOTT A. SCHNEIDER1, 2, 6 1Department of Biology, University of Massachusetts, Amherst, Massachusetts, U.S.A. 2Graduate Program in Organismic and Evolutionary Biology, University of Massachusetts, Amherst, Massachusetts, U.S.A. 3Department of Biology, University of San Diego, San Diego, California, U.S.A. E-mail: [email protected] 4Present address: Finch Therapeutics, Somerville, Massachusetts, U.S.A. 5Graduate School of Human and Environmental Studies, Kyoto University, Japan. E-mail: [email protected] 6Present address: USDA, Agricultural Research Service, Henry A. Wallace Beltsville Agricultural Research Center, Systematic Ento- mology Laboratory, Beltsville, Maryland, U.S.A. E-mail: [email protected] 7Corresponding author. E-mail: [email protected] Magnolia Press Auckland, New Zealand Accepted by G. Watson: 5 Feb. 2019; published: 17 Jun. 2019 Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0 BENJAMIN B. NORMARK, AKIKO OKUSU, GEOFFREY E. MORSE, DANIEL A. PETERSON, TAKAO ITIOKA & SCOTT A. SCHNEIDER Phylogeny and classification of armored scale insects (Hemiptera: Coccomorpha: Diaspididae) (Zootaxa 4616) 98 pp.; 30 cm. 17 Jun. 2019 ISBN 978-1-77670-683-9 (paperback) ISBN 978-1-77670-684-6 (Online edition) FIRST PUBLISHED IN 2019 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] https://www.mapress.com/j/zt © 2019 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition) 2 · Zootaxa 4616 (1) © 2019 Magnolia Press NORMARK ET AL. Table of contents Abstract . 3 Introduction . 5 Materials and methods . 6 Results . 35 Discussion . 44 Keys . 46 Key to groups of scale insects that secrete a waxy scale cover . 46 Key to subfamilies of Diaspididae . 46 Key to tribes of Diaspidinae . 47 Key to subtribes of Diaspidini . 47 Key to tribes of Aspidiotinae . 48 NOTES ON HIGHER TAXA . 48 Family DIASPIDIDAE Targioni Tozzetti . 48 Subfamily ANCEPASPIDINAE Borchsenius, new rank . 49 Subfamily FURCASPIDINAE Balachowsky, new rank . 50 Subfamily DIASPIDINAE Targioni Tozzetti . 51 Tribe LEPIDOSAPHIDINI Shimer . 54 Tribe DIASPIDINI Targioni Tozzetti . 64 Subtribe DIASPIDINA Targioni Tozzetti . 66 Subtribe FIORINIINA Leonardi . 66 Subtribe CHIONASPIDINA Brues & Melander . 71 Subfamily ASPIDIOTINAE Westwood . 76 Tribe GYMNASPIDINI Balachowsky, new rank . 76 Tribe LEUCASPIDINI Atkinson . 77 Tribe AONIDIINI Balachowsky, new rank . 79 Tribe SMILACICOLINI Takagi . 82 Tribe ODONASPIDINI Ferris . 82 Tribe PARLATORIINI Leonardi. 83 Tribe ASPIDIOTINI Westwood . 85 DESCRIPTIONS OF NEW GENERA . 86 Hendersonaspis Normark, gen. n. 86 Brimblecombia Normark, gen. n. 90 Acknowledgements . 93 References . 93 Supplementary material . See DOI link Abstract Armored scale insects (Hemiptera: Coccomorpha: Diaspididae) are major economic pests and are among the world's most invasive species. Here we describe a system of specimen and identification management that establishes a basis for well- vouchered molecular identification. We also present an expanded Bayesian phylogenetic analysis based on concatenated fragments of 4 genetic loci: the large ribosomal subunit (28S), elongation factor-1 alpha (EF-1α), cytochrome oxidase I and II (COI‒II), and the small ribosomal subunit (16S) of the primary endosymbiont, Uzinura diaspidicola (Bacteroidetes: Flavobacteriales). Our sample includes 1,389 individuals, representing 11 outgroup species and at least 311 described and 61 undescribed diaspidid species. The results broadly support Takagi's 2002 classification but indicate that some revisions are needed. We propose a revised classification recognizing 4 subfamilies: Ancepaspidinae Borchsenius, new rank, Fur- caspidinae Balachowsky, new rank, Diaspidinae Targioni Tozzetti, and Aspidiotinae Westwood. Within Aspidiotinae, in addition to the existing tribes Aspidiotini Westwood, Parlatoriini Leonardi, Odonaspidini Ferris, Leucaspidini Atkinson, and Smilacicolini Takagi, we recognize as tribes Gymnaspidini Balachowsky, new rank, and Aonidiini Balachowsky, new rank. Within Diaspidinae we recognize the 2 tribes Lepidosaphidini Shimer and Diaspidini Targioni Tozzetti, and within Diaspidini we recognize three subtribes: Diaspidina Targioni Tozzetti, Fioriniina Leonardi, and Chionaspidina Brues & Melander. We regard Kuwanaspidina Borchsenius as a junior synonym of Fioriniina, Thysanaspidini Takagi as a junior synonym of Leucaspidini, and Protodiaspidina Takagi and Ulucoccinae Takagi as junior synonyms of Chionaspi- dina. To clarify the composition of the higher taxa we describe 2 new genera for Australian species heretofore misplaced in the genus Ancepaspis Ferris: Brimblecombia Normark (Aonidiini) and Hendersonaspis Normark (Leucaspidini). We also propose many additional minor modifications to the taxonomy of Diaspididae, including the following new combi- nations, revived combinations, and replacement names: Aonidia edgerleyi (Mamet), new combination (from Bigymnas- pis Balachowsky); Aonidomytilus espinosai Porter, revived combination (from Porterinaspis González); Aspidiotus PHYLOGENY AND CLASSIFICATION OF DIASPIDIDAE Zootaxa 4616 (1) © 2019 Magnolia Press · 3 badius (Brain), new combination (this and the next 5 Aspidiotus species all from Aonidia Targioni Tozzetti); Aspidiotus biafrae (Lindinger), new combination; Aspidiotus chaetachmeae (Brain), new combination; Aspidiotus laticornis (Bala- chowsky), new combination; Aspidiotus rhusae (Brain), new combination; Aspidiotus sclerosus (Munting), new com- bination; Brimblecombia asperata (Brimblecombe), new combination (this and the next 5 Brimblecombia species all from Ancepaspis); Brimblecombia longicauda (Brimblecombe), new combination; Brimblecombia magnicauda (Brim- blecombe), new combination; Brimblecombia reticulata (Brimblecombe), new combination; Brimblecombia rotundi- cauda (Brimblecombe), new combination; Brimblecombia striata (Brimblecombe), new combination; Cooleyaspis pseudomorpha (Leonardi), new combination (from Dinaspis Leonardi); Cupidaspis wilkeyi (Howell & Tippins), new combination (from Paracupidaspis Howell & Tippins); Cupressaspis isfarensis Borchsenius, revived combination (this species, the next 2 species in Cupressaspis Borchsenius, revived genus, and the next 9 species in Diaspidiotus Cockerell all from Aonidia); Cupressaspis mediterranea (Lindinger), revived combination; Cupressaspis relicta (Balachowsky), new combination; Diaspidiotus atlanticus (Ferris), new combination; Diaspidiotus marginalis (Brain), new combina- tion; Diaspidiotus maroccanus (Balachowsky), new combination; Diaspidiotus mesembryanthemae (Brain), new com- bination; Diaspidiotus opertus (De Lotto), new combination; Diaspidiotus shastae (Coleman), new combination; Diaspidiotus simplex (Leonardi), new combination; Diaspidiotus visci (Hall), new combination; Diaspidiotus.
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    ARTHROPODA Subphylum Hexapoda Protura, Springtails, Diplura, and Insects

    NINE Phylum ARTHROPODA SUBPHYLUM HEXAPODA Protura, springtails, Diplura, and insects ROD P. MACFARLANE, PETER A. MADDISON, IAN G. ANDREW, JOCELYN A. BERRY, PETER M. JOHNS, ROBERT J. B. HOARE, MARIE-CLAUDE LARIVIÈRE, PENELOPE GREENSLADE, ROSA C. HENDERSON, COURTenaY N. SMITHERS, RicarDO L. PALMA, JOHN B. WARD, ROBERT L. C. PILGRIM, DaVID R. TOWNS, IAN McLELLAN, DAVID A. J. TEULON, TERRY R. HITCHINGS, VICTOR F. EASTOP, NICHOLAS A. MARTIN, MURRAY J. FLETCHER, MARLON A. W. STUFKENS, PAMELA J. DALE, Daniel BURCKHARDT, THOMAS R. BUCKLEY, STEVEN A. TREWICK defining feature of the Hexapoda, as the name suggests, is six legs. Also, the body comprises a head, thorax, and abdomen. The number A of abdominal segments varies, however; there are only six in the Collembola (springtails), 9–12 in the Protura, and 10 in the Diplura, whereas in all other hexapods there are strictly 11. Insects are now regarded as comprising only those hexapods with 11 abdominal segments. Whereas crustaceans are the dominant group of arthropods in the sea, hexapods prevail on land, in numbers and biomass. Altogether, the Hexapoda constitutes the most diverse group of animals – the estimated number of described species worldwide is just over 900,000, with the beetles (order Coleoptera) comprising more than a third of these. Today, the Hexapoda is considered to contain four classes – the Insecta, and the Protura, Collembola, and Diplura. The latter three classes were formerly allied with the insect orders Archaeognatha (jumping bristletails) and Thysanura (silverfish) as the insect subclass Apterygota (‘wingless’). The Apterygota is now regarded as an artificial assemblage (Bitsch & Bitsch 2000).
  • Peña & Bennett: Annona Arthropods 329 ARTHROPODS ASSOCIATED

    Peña & Bennett: Annona Arthropods 329 ARTHROPODS ASSOCIATED

    Peña & Bennett: Annona Arthropods 329 ARTHROPODS ASSOCIATED WITH ANNONA SPP. IN THE NEOTROPICS J. E. PEÑA1 AND F. D. BENNETT2 1University of Florida, Tropical Research and Education Center, 18905 S.W. 280th Street, Homestead, FL 33031 2University of Florida, Department of Entomology and Nematology, 970 Hull Road, Gainesville, FL 32611 ABSTRACT Two hundred and ninety-six species of arthropods are associated with Annona spp. The genus Bephratelloides (Hymenoptera: Eurytomidae) and the species Cerconota anonella (Sepp) (Lepidoptera: Oecophoridae) are the most serious pests of Annona spp. Host plant and distribution are given for each pest species. Key Words: Annona, arthropods, Insecta. RESUMEN Doscientas noventa y seis especies de arthrópodos están asociadas con Annona spp. en el Neotrópico. De las especies mencionadas, el género Bephratelloides (Hyme- noptera: Eurytomidae) y la especie Cerconota anonella (Sepp) (Lepidoptera: Oecopho- ridae) sobresalen como las plagas mas importantes de Annona spp. Se mencionan las plantas hospederas y la distribución de cada especie. The genus Annona is confined almost entirely to tropical and subtropical America and the Caribbean region (Safford 1914). Edible species include Annona muricata L. (soursop), A. squamosa L. (sugar apple), A. cherimola Mill. (cherimoya), and A. retic- ulata L. (custard apple). Each geographical region has its own distinctive pest fauna, composed of indigenous and introduced species (Bennett & Alam 1985, Brathwaite et al. 1986, Brunner et al. 1975, D’Araujo et al. 1968, Medina-Gaud et al. 1989, Peña et al. 1984, Posada 1989, Venturi 1966). These reports place emphasis on the broader as- pects of pest species. Some recent regional reviews of the status of important pests and their control have been published in Puerto Rico, U.S.A., Colombia, Venezuela, the Caribbean Region and Chile (Medina-Gaud et al.