Ontogenesis, morphology and higher classification of archaeococcids (Homoptera: Coccinea: Orthezioidea)

I.A. Gavrilov-Zimin

Abstract. The monograph summarizes original research data and published literature data on reproduction, life cycles, individual development and morphology of scale insects of the superfamily Orthezioidea (archaeococcids). The superfamily system is accepted mainly in its traditional concept, i.e. with four well-defined families: Margarodidae s. l., Ortheziidae, Carayonemidae, and Phenacoleachiidae. The tribe Matsucoccini (Margarodidae s. l.: Xylococcinae s. l.) is considered as a most archaic group of scale insects according to morphological, reproductive and ontogenetic characters. A complicated ontogenesis with an alternation of movable/immovable instars and with arostrate imago of both sexes (as in Matsucoccus Cockerell, 1909 and many other Margarodidae s. l.) is presumed to be initial in scale insect evolution and such ontogenesis is supposed to be an apomorphy of suborder Coccinea. Distribution of different variants of ovoviviparity/viviparity amongst scale insect families is overviewed. It is demonstrated that the evolution of scale insects shows multiple cyclic conversions of oviparous reproduction pattern to ovoviviparous/viviparous ones with the appearance of new and new peculiar adaptations to eggs protection; the most ancient scale insects (Matsucoccini and their ancestor) were probably facultatively ovoviviparous, whereas the origin of the whole neococcid phylogenetic line (Coccoidea s. s.) was probably connected with obligate complete ovoviviparity, which also appeared in some “derived” archaeococcids of the tribe Iceryini (Margarodidae s. l.), in the families Phenacoleachiidae and Carayonemidae. New taxonomic additions and changes in generic composition of some tribes are provided for the family Margarodidae s. l., in its subfamilies Monophlebinae and Callipappinae s. l. The tribe Labioproctini tr. nov. (Monophlebinae) is erected for six genera possessing peculiar quadrilocular wax pores: Aspidoproctus Newstead, 1901, Hemaspidoproctus Morrison, 1927, Labioproctus Green, 1922, Lecaniodrosicha Takahashi, 1930, Misracoccus Rao, 1950, and Walkeriana Signoret, 1876. The presence of quadrilocular pores are considered as a synapomorphic character of the Labioproctini tr. nov. and Ortheziidae. Disputable taxonomic position of Xenococcidae Tang, 1992 is discussed and this family is also placed in Orthezioidea. New genera and species are described and illustrated, based mainly on material collected in the Oriental region: Eremostoma klugei gen. et sp. nov., Crambostoma largecicatricosum gen. et sp. nov. (both in Callipappinae s. l.: Coelostomidiini s. l.), Buchnericoccus reynei sp. nov., Monophlebus neglectus sp. nov. (both in Monophlebinae: Monophlebini), Crypticerya ovivivipara sp. nov., Icerya oculicicatricata sp. nov., I. siamensis sp. nov. (all three in Monophlebinae: Iceryini).
Key words: life cycles, reproductive biology, ovoviviparity, viviparity, comparative morphology, scale insects, giant scale insects, new taxa

Zoosystematica Rossica, 2018, Supplementum 2: 1-260  ▪  Published in print 31 May 2018

https://doi.org/10.31610/zsr/2018.supl.2.1  ▪  Open full article  


Anisimov A.P. 1992. Simple method of preparing slides with using of cellophane. Tsitologiya, 34(11–12): 110–112. (In Russian).

Baer K.M. 1866. About Professor Wagner’s discovery of agamic reproduction of larvae and about additional observations of this phenomenon by Mr. Ganin and about paedogenesis in general. Zapiski Imperatorskoy Akademii Nauk (St Petersburg), 10 (Supplement 1): 1–77. (In Russian). [Version in Germain: Baer K.H. 1866. Ueber Prof. Nic. Wagners’s Entdeckung von Larvae die sich fortpflanzen. Herr Ganin’s verwandte und ergänzende Beobachtungen und über dei Paedogenese überhaupt. Bulletin de l’Académie Impériale des Sciences de St. Petersburg, 9: 64–137.]

Beardsley J.W. 1964. Insects of Campbell Island. Homoptera: Coccoidea. Pacific Insects, 7: 238–252.

Beardsley J.W. 1969. A new fossil scale insect (Homoptera: Coccoidea) from Canadian amber. Psyche, 76: 270–279. https://doi.org/10.1155/1969/82354

Beardsley J.W. 1970. Three new species of Chavesia Balachowsky from tropical America (Homoptera: Coccoidea). Proceedings of the Hawaiian entomological Society, 20: 509–520.

Becker-Migdisova E.E. 1959. Some new members of Sternorrhyncha from Permian and Mesozoic depositions of USSR. Materialy k “Osnovam paleontologii”, 3: (In Russian).

Ben-Dov Y. 2005. A systematic catalogue of the scale insect family Margarodidae (Hemiptera: Coccoidea) of the world. Wimborne: Intercept Ltd. 400 p.

Ben-Dov Y. 2011. An updated checklist of the scale insects (Hemiptera: Coccoidea) of the Margarodidae sensu lato group. Zootaxa, 2859: 1–62.

Bazarov B.B. & Shmelev G.P. 1971. Shchitovki (Homoptera, Coccoidea) Tadzhikistana i sopredel’nykh rayonov Sredney Azii [Scale insects (Homoptera, Coccoidea) of Tadzhikistan and the adjoining regions of the Central Asia]. Dushanbe: Akademiya Nauk Tadzhikskoy SSR, Institut Zoologii i Parazitologii. 238 p. (In Russian).

Bhatti S. 1990. A new monophlebine genus (Homoptera: Coccoidea: Margarodidae: Monophlebinae) on Melaleuca L. in Australia. Invertebrate Taxonomy, 3: 495–516. https://doi.org/10.1071/IT9890495

Bhatti S. & Gullan P.J. 1990. New Margarodid species (Homoptera: Coccoidea: Margarodidae: Monophlebinae) from New Guinea. Invertebrate Taxonomy, 3: 877–911. https://doi.org/10.1071/IT9890877

Blackman R.L. 1987. Reproduction, cytogenetics and development. In: Minks A.K. & Harrewijn P. (Eds). Aphids, their biology, natural enemies and control, 2A: 163–195. Amsterdam: Elsevier.

Blackman R.L. 1995. Sex determination in insects. In: Leather S.R. & Hardie J. (Eds). Insect Reproduction: 57–94. Boca Raton: CRC Press. https://doi.org/10.1201/9781351073608-3

Bodenheimer F.S. & Harpaz I. 1955. Description of the various stages of Matsucoccus josephi n. sp. In: Bodenheimer F.S. & Neumark S. (Eds). The Israeli Pine Matsucoccus: 12–22. Jerusalem.

Börner C. 1910. Parametabolie und Neotenie bei Cocciden. Zoologischer Anzeiger, 35: 553–561.

Boratynski K.L. 1952. Matsucoccus pini (Green, 1925) (Homoptera, Coccoidea: Margarodidae). Bionomics and external anatomy with reference to the variability of some taxonomic characters. Transactions of the Royal Entomological Society of London, 103: 285–326. https://doi.org/10.1111/j.1365-2311.1952.tb01067.x

Borchsenius N.S. 1956. The question of the course of evolution of Coccoidea (Insecta, Homoptera). Zoologicheskiy Zhurnal, 35: 546–553. (In Russian).

Borchsenius N.S. 1960. Podotryad chervetsy i shchitovki (Coccoidea). Semeystva Kermococcidae, Asterolecaniidae, Lecanidodiaspididae, Aclerdidae [Coccoidea. Families Kermococcidae, Asterolecaniidae, Lecanidodiaspididae, Aclerdidae]. Fauna of Russia and neighbouring countries. New series, 77. Insecta: Hemiptera. Leningrad: Zoologicheskiy Institut. 282 p. (In Russian).

Borchsenius N.S. & Ter-Grigorian M.A. 1956. Mealybugs parasitising wheat and other grains in Armenian SSR. Izvestiya Akademii Nauk Armyanskoy SSR. Biologicheskikh i Sel’skokhozyaystvennykh Nauk, 9: 17–27. (In Russian).

Brain C.K. 1915. The Coccidae of South Africa. Transactions of the Royal Society of South Africa, 5: 65–194. https://doi.org/10.1080/00359191509519718

Brimblecombe A.R. 1960. Studies of the Coccoidea. 11. New genera and species of Monophlebidae. Queensland Journal of Agricultural Science, 17: 183–193.

Brown L.R. & Eads C.O. 1965. A technical study of insects affecting the sycamore tree in southern California.

Bulletin of the California Agricultural Experiment Station, 818: 3–38.

Brown S.W. 1958. The chromosomes of an Orthezia species (Coccoidea–Homoptera). Cytologia, 23: 429–434. https://doi.org/10.1508/cytologia.23.429

Brown S.W. 1967. Chromosome systems of the Eriococcidae (Coccoidea–Homoptera). 1. A survey of several genera. Chromosoma, 22: 126–150. https://doi.org/10.1007/BF00326725

Brown S.W. 1977. Adaptive status and genetic regulation in major evolutionary changes of coccid chromosome systems. Nucleus, 20: 145–157.

Brown S.W. & Cleveland C. 1968. Meiosis in the male of Puto albicans (Coccoidea–Homoptera). Chromosoma, 24: 210–232. https://doi.org/10.1007/BF00285204

Brown S.W. & Nelson-Rees W. 1961. Radiation analysis of a lecanoid genetic system. Genetics, 46: 983–1007.

Brożek J., Szwedo D., Gaj S. & Pilarczyk S. 2003. Former and current views on the classification of the bugs (Insecta, Hemiptera). Genus, Supplement 1: 85–100.

Buchner P. 1954. Endosymbiosestudien an Schildläusen. I. Stictococcus sjoestedti. Zeitschrift für Morphologie und Ökologie der Tiere, 43: 262–312. https://doi.org/10.1007/BF00391845

Buchner P. 1955. Endosymbiosestudien an Schildläusen. II. Stictococcus diversiseta. Zeitschrift für Morphologie und Ökologie der Tiere, 43: 397–424. https://doi.org/10.1007/BF00407487

Buchner P. 1957. Endosymbiosestudien an Schildläusen. 6. Die nicht in Symbiose lebende Gattung Apiomorpha und ihre ungewöhniche Embryonalentwicklung. Zeitschrift für Morphologie und Ökologie der Tiere, 46: 481–528. https://doi.org/10.1007/BF00410242

Buchner P. 1963. Endosymbiosestudien an Schildläusen. 7. Weitere Beiträge zur Kenntnis der Stictococcinensymbiose. Zeitschrift für Morphologie und Ökologie der Tiere, 52: 401–458. https://doi.org/10.1007/BF00408569

Buchner P. 1965. Endosymbiosis of animals with plant-like microorganisms. New York: John Wiley & Sons. 909 p.

Buchner P. 1966. Endosymbiosestudien an Schildläusen. 8. Die Symbiosen der Palaeococcoidea. Pt. I. Zeitschrift für Morphologie und Ökologie der Tiere, 56: 275–362. https://doi.org/10.1007/BF00409442

Buchner P. 1967. Endosymbiosestudien an Schildläusen. 8. Die Symbiosen der Palaeococcoidea. Pt. 2. Zeitschrift für Morphologie und Ökologie der Tiere, 59: 211–317. https://doi.org/10.1007/BF00406754

Buchner P. 1969. Endosymbiosestudien an Schildläusen. 8. Die Symbiosen der Palaeococcoidea. Pt. 3. Zeitschrift für Morphologie und Ökologie der Tiere, 64: 201–308. https://doi.org/10.1007/BF00298359

Burmeister H. 1835. Handbuch der Entomologie. 2. Berlin: T.F. Inslin. 400 p.

Campbell B.C., Steffen–Campbell J.D., Sorensen J.T. & Gill R.J. 1995. Paraphyly of Homoptera and Auchenorrrhyncha inferred from 18 S rDNA nucleotide sequences. Systematic Entomology, 20: 175–194. https://doi.org/10.1111/j.1365-3113.1995.tb00090.x

Carayon J. 1961. La viviparité chez les Hétéroptères. Verhanglungen: XI Internationaler Kongress für Entomologie, Wien, 1: 711–714.

Cockerell T.D.A. 1894. A check list of Nearctic Coccidae. Canadian Entomologist, 26: 31–36. https://doi.org/10.4039/Ent2631-2

Cockerell T.D.A. 1902a. A contribution to the classification of the Coccidae. The Entomologist, 35: 232–233. https://doi.org/10.5962/bhl.part.19973

Cockerell T.D.A. 1902b. What is Monophlebus Leach? The Entomologist, 35: 317–319.

Cox J.M. 1983(1984). Relationships of the Phenacoleachiidae (Homoptera: Coccoidea). Proceedings of the 10th International Symposium of Central European Entomofaunistics, Budapest, 15–20 August 1983: 339–341.

Danzig E.M. 1978. Scale insect fauna of South Sakhalin and Kunashir. Trudy Biologo-Pochvennogo Instituta Akademii Nauk SSR, 50: 3–23. (In Russian).

Danzig E.M. 1980. Koktsydy Dal’nego Vostoka SSSR (Homoptera, Coccinea) s analizom filogenii koktsyd mirovoy fauny. Leningrad: Nauka. 367 p. (In Russian). [English edition: Danzig, E.M. 1986. Coccids of the Far-Eastern USSR (Homoptera: Coccinea). Phylogenetic analysis of coccids in the world fauna. New Delhi. 450 p.]

Danzig E.M. 2000. A new species of gall-forming armored scale insect from Israel (Homoptera, Coccinea: Diaspidi-dae). Zoosystematica Rossica, 8(2): 287–289.

Danzig E.M. 2007. Mealybugs of the genus Fonscolombia Licht. (Homoptera, Pseudococcidae) of the fauna of Russia and adjacent countries. Entomologicheskoe Obozrenie, 86(2): 363–377. (In Russian). https://doi.org/10.1134/S0013873806020084

Danzig E.M. 2007. Mealybugs of the genus Heliococcus Šulc (Homoptera, Pseudococcidae) of the fauna of Russia and neighbouring countries. Entomologicheskoe Obozrenie, 86(3): 567–609.

Danzig E.M. & Gavrilov–Zimin I.A. 2014. Palaearctic mealybugs (Homoptera: Coccinea: Pseudococcidae). Part 1. Subfamily Phenacoccinae. Fauna of Russia and neighbouring countries. New series, 148. Insecta: Hemiptera: Arthroidignatha. St Petersburg: ZIN RAS. 678 p.

Darlington Ph.J. 1957. Zoogeography: the geographical distribution of animals. New York: John Wiley & Sons, Inc. 675 p.

Delabie J.H.C., Mariano C.S.F., Serrao J.E. & Pompolo S.G. 2004. The karyotype of the aberrant obligate ant-attended mealybug, Neochavesia caldasiae (Coccoidea: Pseudococcidae: Rhizoecinae). Caldasia, 26(1): 185–190.

Delavault R., Royer M. & Duvigneau C. 1969. Chronologie de l’embryogenèse d’Icerya purchasi Mask. (Insecte, Coccide). Annales d’Embryologie et de Morphogénèse, 3(1): 71–80.

De Klerk C.A. 1980. Biology of Margarodes vredendalensis De Klerk (Coccoidea: Margarodidae) in South Africa. South African Journal of Enology and Viticulture, 1: 47–58. https://doi.org/10.21548/1-1-2413

De Marzo L., Romano V. & Tranfaglia A. 1990. Types of the female reproductive system in some scale insects (Homoptera: Coccoidea). In: Koteja J. (Ed.). Proceedings of the Sixth International Symposium of Scale Insect Studies, Part II. Cracow, Poland: August 6–12, 1990: 41–45. Cracow: Agricultural University Press.

Depa L., Kanturski M., Junkiert L. & Wieczorek K. 2015. Giant females vs dwarfish males of the genus Stomaphis Walker (Hemiptera: Aphididae) – an aphid example of the ongoing course to permanent parthenogenesis? Arthropod Systematics & Phylogeny, 73(1): 19–40.

Derbeneva N.N. 1959. Life-cycle of Haplothrips yuccae Sav. (Thysanoptera, Phloeothripidae). Entomologicheskoe Obozrenie, 38(1): 64–81 (In Russian).

Derbeneva N.N. 1962. The biology and postembryonal development of Taeniothrips fedorovi Priesner (Thysanoptera, Thripidae). Entomologicheskoe Obozrenie, 41(2): 322–333. (In Russian).

Derbeneva N.N. 1967. New data on the biology and structure of preimaginal phases and stages of the predatory thrips Aeolothrips intermedius Bagnall (Thysanoptera, Aeolothripidae). Entomologicheskoe Obozrenie, 46(3): 629–644. (In Russian).

Dikshith T.S.S. 1964. Chromosome behaviour in Laccifer lacca (Kerr) Lacciferidae–Coccoidea. Cytologia, 29: 337–345. https://doi.org/10.1508/cytologia.29.337

Dohlen C.D., von & Moran N.A. 1995. Molecular phylogeny of the Homoptera: a paraphyletic taxon. Journal of Molecular Evolution, 41: 211–223. https://doi.org/10.1007/BF00170675

D’Urso V. 2002. The wing-coupling apparatus of Hemiptera Auchenorrhyncha: structure, function, and systematic value. Denisia, 4: 401–410.

Eichler W. 1946. Parthenogenese und Ovoviviparie als Entwicklungseigentümlichkeiten bei Läusen und Federlingen. Tierärztliche Umschau, 1(1): 10.

Emeljanov A.F. 1987. Phylogeny of cicads (Homoptera, Cicadina) based on comparative morphological studies. Trudy Vsesoyuznogo Entomologicheskogo Obschestva, 69: 19–109. (In Russian).

Eskov K.Yu. 2002. Geographical history of insects. In: Rasnitsyn A.P. & Quicke D.L.J. (Eds). History of Insects: 427–435. Dordrecht–Boston–London: Kluwer Academic Press.

Essig E.O. 1948. Mounting aphids and other small insects on microscopic slides. Pan-Pacific Entomologist, 24: 9–22.

Favret C., Blackman R.L., Miller G.L. & Victor B. 2016. Catalog of the phylloxerids of the world (Hemiptera, Phylloxeridae). ZooKeys, 629: 83–101. https://doi.org/10.3897/zookeys.629.10709

Fernando W. 1935. The early embryology of viviparous Psocid. Quarterly Journal of Microscopic Science, 77: 99–120.

Ferris G.F. 1919. A remarkable case of longevity in insects (Hem., Hom.). Entomological News, 30: 27–28.

Ferris G.F. & Murdock G.E. 1936. Contributions to the knowledge of the Coccoidea (Homoptera). III. (Contribution No. 4). Microentomology, 1: 115–121.

Forero D. 2008. The systematics of the Hemiptera. Revista Colombiana de Entomologia, 34(1): 1–21.

Foldi I. 1990. 1.1.3 Internal anatomy. In: Rosen D. (Ed.). Armored scale insects, their biology, natural enemies and control, 4A: 65–80. Amsterdam–Oxford–New York–Tokyo: Elsevier.

Foldi I. 1997a. Internal anatomy of the Adult Female. In: Ben-Dov Y. & Hodgson Ch.J. (Eds). Soft scale insects: their biology, natural enemies and control, 7A: 73–90 Amsterdam–Oxford–New York–Tokyo: Elsevier. https://doi.org/10.1016/S1572-4379(97)80045-0

Foldi I. 1997b. Defense strategies in scale insects: phylogenetic inference and evolutionary scenarios (Hemiptera, Coccoidea). Mémoires du Museum National d’Histoire Naturelle (N.S.) Serie A, Zoologie, 173: 203–230.

Foldi I. 2004. The Matsucoccidae in the Mediterranean basin with a world list of species (Hemiptera: Sternorrhyncha: Coccoidea). Annales de la Société Entomologique de France, 40(2): 145–168. https://doi.org/10.1080/00379271.2004.10697412

Foldi I. 2005. Ground pearls: a generic revision of the Margarodidae sensu stricto. (Hemiptera: Sternorrhyncha: Coccoidea). Annales de la Société Entomologique de France, 41: 81–25. https://doi.org/10.1080/00379271.2005.10697442

Foldi I. 2006. Deux nouvelles Cochenilles du Brésil et de l’Équateur (Hemiptera, Sternorhyncha, Coccoidea). Bulletin de la Societe Entomologique de France, 111: 101–113.

Foldi I. 2009. Archaeococcoid scale insects (Hemiptera: Coccoidea) from the tropical high mountains of the Andean Cordillera, South America. Zootaxa, 2300: 1–38. https://doi.org/10.11646/zootaxa.2300.1.1

Foldi I. 2011. Archéococcoïdes du Mexique : une espèce nouvelle de Cryptokermes Hempel, 1900, et description des stades de développement de C. mimosae Foldi, 1995 (Hemiptera, Coccoidea, Coelostomidiidae.) Bulletin de la Société entomologique de France, 116(4): 483–493.

Foldi I. 2016. New archaeococcoids from Guyana and Malaysia, with discussion of the tribes Llaveiini and Drosichini (Hemiptera, Coccomorpha, Monophlebidae). Bulletin de la Societe Entomologique de France, 121(3): 289–308

Foldi I. & Gullan P.J. 2014. Review of the Neotropical scale insects formerly assigned to Coelostomidiidae and here transferred to a new tribe within the Monophlebidae (Hemiptera: Sternorrhyncha: Coccoidea). Zootaxa, 3903(1): 1–62. https://doi.org/10.11646/zootaxa.3903.1.1

Foldi I. & Watson G.W. 2001. A new pest scale insect on avocado trees in Colombia, Laurencella colombiana, sp. n. (Hemiptera: Coccoidea: Margarodidae). Annales de la Société Entomologique de France, 37(3): 367–374.

Fuller C. 1899. XIV. Notes and descriptions of some species of Western Australian Coccidae. Transactions of the Entomological Society of London, 1899: 435–473. https://doi.org/10.1111/j.1365-2311.1899.tb00988.x

Gabritschesky E.G. 1923. Postembrional development, parthenogenesis and “pedogamy” in scale insects (Coccidae). Russkiy Zoologicheskiy Zhurnal, 3(3–4): 295–332. (In Russian).

García Morales M., Denno B.D., Miller D.R., Miller G.L., Ben-Dov Y. & Hardy N.B. 2016. ScaleNet: A literature-based model of scale insect biology and systematics. Database. Available from: http://scalenet.info; https://doi.org/10.1093/database/bav118 [viewed 1 March 2018].

Gavrilov I.A. 2007. A catalogue of chromosome numbers and genetic systems of scale insects (Homoptera: Coccinea) of the world. Israel Journal of Entomology, 37: 1–45.

Gavrilov I.A. 2007b. A revision of the mealybug genus Mirococcopsis Borchsenius (Homoptera: Coccinea: Pseudococcidae) based on the structure of the adult females. Zootaxa, 1586: 1–23. https://doi.org/10.11646/zootaxa.1586.1.1

Gavrilov-Zimin I.A. 2017. Contribution to the cytogenetics of Kuwaniini scale insects (Homoptera, Coccinea, Margarodidae s.l.). Comparative Cytogenetics, 11(4) : 659–663. https://doi.org/10.3897/CompCytogen.v11i4.20168

Gavrilov-Zimin I.A. & Danzig E.M. 2012. Taxonomic position of the genus Puto Signoret (Homoptera: Coccinea: Pseudococcidae) and separation of higher taxa in Coccinea. Zoosystematica Rossica, 22(1) : 97–111.

Gavrilov-Zimin I.A., Stekolshikov A.V. & Gautam D.C. 2015. General trends of chromosomal evolution in Aphidococca (Insecta, Homoptera, Aphidinea + Coccinea). Comparative Cytogenetics, 9(3): 335–422. https://doi.org/10.3897/CompCytogen.v9i3.4930

Gavrilov I.A. & Trapeznikova I.V. 2008. Cytogenetic studies of European Pulvinariini (Homoptera: Coccinea). Comparative Cytogenetics, 2(2): 123–131.

Gómez-Menor Ortega J. 1957. Adiciones a los cóccidos de España (4a nota). EOS, 33: 39–86.

Gorochov A.V. 2001. On some theoretical aspects of taxonomy (remarks by the practical taxonomist). Acta Geologica Leopoldensia, 24(52/53): 57–71.

Gounari S. 2004. Seasonal development and ovipositing behavior of Marchalina hellenica (Hemiptera: Margarodidae). Entomologia Hellenica, 15: 27–37. https://doi.org/10.12681/eh.14045

Green E.E. 1922. The Coccidae of Ceylon, Part V. London: Dulau & Co. 472 p.

Grozeva S. & Nokkala S. 1996. Chromosomes and their behavior in two families of the primitive infraoder Dipsocoromorpha (Heteroptera). Hereditas, 125: 31–36. https://doi.org/10.1111/j.1601-5223.1996.t01-1-00031.x

Guérin-Méneville F.E. 1827. Dictionnaire Classique d’Histoire Naturelle, 11. Paris: Rey et Gravier, Libraires-Èditeurs. 615 p.

Gullan P.J. 1999(2001). Why the taxon Homoptera does not exist. Entomologica (Bari), 33: 101–104.

Gullan P.J. & Brookes H.M. 1998. Giant females and bird-of-paradise flies: Notes on the biology of Callipappus Guerin-Meneville (Hemiptera: Margarodidae). Australian Journal of Entomology, 37: 2–7. https://doi.org/10.1111/j.1440-6055.1998.00007.x

Gullan P.J. & Cook L.G. 2001(2002). Phenacoleachia Cockerell (Hemiptera: Coccoidea: Phenacoleachiidae) revisited. Bollettino di Zoologia Agraria e di Bachicoltura (Milano), 33(3): 163–173.

Gullan P.J. & Sjaarda A.W. 2001. Trans-Tasman Platycoelostoma Morrison (Hemiptera: Coccoidea: Margarodidae) on endemic Cupressaceae, and the phylogenetic history of margarodids. Systematic Entomology, 26: 257–278. https://doi.org/10.1046/j.1365-3113.2001.00145.x

Gullan P.J. & Williams D.J. 2016. A new pupillarial scale insect (Hemiptera: Coccoidea: Eriococcidae) from Angophora in coastal New South Wales, Australia. Zootaxa, 4117(1): 85–100. https://doi.org/10.11646/zootaxa.4117.1.4

Hadzibejli Z.K. 1958. A new genus and species of mealybugs (Homoptera, Pseudococcidae) from Georgia, Transcaucasia. Entomologicheskoe Obozrenie, 37: 903–908. (In Russian).

Hadzibejli Z.K. 1963. Coccids of the family Ortheziidae (Homoptera, Coccoidea) in the fauna of Georgia. Entomologicheskoe Obozrenie, 42: 611–620. (In Russian).

Hadzibejli Z.K. 1966. On the biology and morphology of the genera Neomargarodes Green and Porphyrophora Brandt (Homoptera, Coccoidea). Entomologicheskoe Obozrenie, 45: 693–711. (In Russian).

Hadzibejli Z.K. 1968. Ecology of Laccifer lacca Kerr (Homoptera, Coccoidea) under the conditions of the Black Sea coast of Georgia. Entomologicheskoe Obozrenie, 47: 10–18. (In Russian).

Hadzibejli Z.K. 1969a. On Marchalina caucasica, sp. n. (Homoptera, Coccoidea) from the Caucasus. Entomologicheskoe Obozrenie, 48: 391–398. (In Russian).

Hadzibejli Z.K. 1969b. Fauna i ekologiya koktsyd (Insecta, Homoptera, Coccidea) Gruzii [Fauna and ecology of scale insects (Insecta, Homoptera, Coccidea) of Georgia]. Abstract of doctor of science (biology) dissertation. Tbilisi: University of Tbilisi. 36 p. (In Russian).

Hadzibejli Z.K. 1983. Koktsydy subtropicheskoy zony Gruzii [Coccids of the subtropical zone of Georgia]. Tbilisi: Metsniereba. 293 p. (In Russian).

Hafez M. & Salama H.S. 1967. Polymorphism in the male of the mealy bug Saccharicoccus sacchari (Cockerell). Proceedings of the Royal Entomological Society of London, Series A: General Entomology, 42: 25–29. https://doi.org/10.1111/j.1365-3032.1967.tb00684.x

Hagan H.R. 1951. Embryology of viviparous insects. New York. 472 p.

Hambleton E.J. 1946. Studies of hypogeic mealybugs. Revista de Entomologia (Rio de Janeiro), 17: 1–77.

Hamon A.B. & Williams M.L. 1984. The soft scale insects of Florida (Homoptera: Coccoidea: Coccidae). Arthropods of Florida and Neighboring Land Areas, 11. Gainesville: Florida Dept. of Agriculture and Consumer Services, Division of Plant Industry. 194 p.

Hathaway C.R. 1938. Verificação da viviparidade em Thysanoptera. Memoirias do Instituto Oswaldo Cruz, 33: 357–358. https://doi.org/10.1590/S0074-02761938000300002

Heie O.E. 1994. Why are there so few aphid species in the temperate areas of the southern hemisphere? European Journal of Entomology, 91: 127–133.

Hempel A. 1920. Descripçoes de coccidas novas e pouco conhecidas. Revista do Museu Paulista (São Paulo), 12: 329–377.

Hempel A. 1932. Descripção de vinte a duas espécies novas de coccideos (Hemiptera–Homoptera). Revista de Entomologia, 2: 310–339.

Heriot A.D. 1934. The renewal and replacement of the stylets of sucking insects during each stadium, and the method of penetration. Canadian Journal of Research, 11: 602–612. https://doi.org/10.1139/cjr34-117

Hille Ris Lambers D. 1950. An apparently unrecorded mode of reproduction in aphididae. Proceedings of the eighth International Congress of Entomology. Axel R. Elfströms Boktryckeri, Stockholm, Sweden, p. 235.

Hodgson C.J. 2012. Comparison of the morphology of the adult males of the rhizoecine, phenacoccine and pseudococcine mealybugs (Hemiptera: Sternorrhyncha: Coccoidea), with the recognition of the family Rhizoecidae Williams. Zootaxa, 3291: 1–79. https://doi.org/10.11646/zootaxa.3291.1.1

Hodgson C.J. 2014. Phenacoleachia, Steingelia, Pityococcus and Puto – neococcids or archaeococcids? An intuitive phylogenetic discussion based on adult male characters. Acta Zoologica Bulgarica, Supplement 6: 41–50.

Hodgson C.J. & Foldi I. 2006. A review of the Margarodidae sensu Morrison (Hemiptera: Coccoidea) and some related taxa based on the morphology of adult males. Zootaxa, 1263: 1–250. https://doi.org/10.11646/zootaxa.1263.1.1

Hodgson C.J, Gamper H., Bogo A. & Watson G. 2007. A taxonomic review of the Margarodid genus Stigmacoccus Hempel (Hemiptera: Sternorrhyncha: Coccoidea: Stigmacoccidae), with some details on their biology. Zootaxa, 1507: 1–55. https://doi.org/10.11646/zootaxa.1507.1.1

Hodgson C.J. & Henderson R.C. 2000. Coccidae (Insecta: Hemiptera: Coccoidea). Canterbury: Manaaki Whenua Press Lincoln. 259 p.

Hołyński R.B. 2005. Philosophy of science from taxonomist’s perspective. Genus, 16(4): 469–502.

Hovasse R. 1930. Marchalina hellenica essai de monographie d’une cochenille. Bulletin Biologique de la France et de la Belgique, 64: 389–449.

Howell J.O. & Tippins H.H. 1990. Pupillarial forms. In: Rosen D. (Ed.). Armored scale insects, their tiology, natural enemies and control., 4A: 55–57. Amsterdam–Oxford–New York–Tokyo: Elsevier.

Hughes-Schrader S. 1925. Cytology of hermaphroditism in Icerya purchasi (Coccidae). Zeitschrift für Zellforschung und Mikroskopische Anatomie, 2: 264–292. https://doi.org/10.1007/BF00376393

Hughes-Schrader S. 1930a. Contributions to the life history of the iceryine coccids, with special reference to parthenogenesis and hermaphroditism. Annals of the Entomological Society of America, 23: 359–380. https://doi.org/10.1093/aesa/23.2.359

Hughes-Schrader S. 1930b. The cytology of several species of iceryine coccids with special reference to parthenogenesis and haploidy. Journal of Morphology and Physiology, 50: 475–495. https://doi.org/10.1002/jmor.1050500208

Hughes-Schrader S. 1931. A study of the chromosome cycle and the meiotic division-figure in Llaveia bouvari – a primitive coccid. Zeitschrift für Zellforschung und Mikroskopische Anatomie, 13: 742–769. https://doi.org/10.1007/BF02450250

Hughes-Schrader S. 1940. The meiotic chromosomes of the male Llaveiella taenechina Morrison (Coccidae) and the question of the tertiary split. Biological Bulletin, 78: 312–337. https://doi.org/10.2307/1537781

Hughes-Schrader S. 1942. The chromosomes of Nautococcus schraderae Vays. and the meiotic division figure of male Llaveiine coccids. Journal of Morphology, 70: 261–299. https://doi.org/10.1002/jmor.1050700207

Hughes-Schrader S. 1944. A primitive coccid chromosome cycle in Puto sp. Biological Bulletin, 87: 167–176. https://doi.org/10.2307/1537954

Hughes-Schrader S. 1948. Cytology of coccids (Coccoidea–Homoptera). Advances in Genetics, 2: 127–203. https://doi.org/10.1016/S0065-2660(08)60468-X

Hughes-Schrader S. 1955. The chromosomes of the giant scale Aspidoproctus maximus Louns. (Coccoidea-Margarodidae) with special reference to asynaspis and sperm formation. Chromosoma (Berlin), 7: 420–438. https://doi.org/10.1007/BF00329735

Hughes-Schrader S. 1963. Hermaphroditism in an African coccid, with notes on other Margarodids (Coccoidea–Homoptera). Journal of Morphology, 113: 173–184. https://doi.org/10.1002/jmor.1051130205

Hughes-Schrader, S. Monahan, D.F. 1966 Hermaphroditism in Icerya zeteki Cockerell, and the mechanism of gonial reduction in iceryine coccids (Coccoidea: Margarodidae Morrison). Chromosoma (Berlin), 20: 15–31. https://doi.org/10.1007/BF00331895

Hughes-Schrader S. & Ris H. 1941. A diffuse spindle attachment of coccids, verified by the mitotic behavior of induced chromosome fragments. Journal of Experimental Zoology, 87: 429–456. https://doi.org/10.1002/jez.1400870306

Hughes-Schrader S. & Schrader F. 1961. The kinetochore of the Hemiptera. Chromosoma (Berlin), 12: 327–350. https://doi.org/10.1007/BF00328928

Hughes-Schrader S. & Tremblay E. 1966. Gueriniella and the cytotaxonomy of iceryine coccids (Coccoidea: Margarodidae). Chromosoma (Berlin), 19: 1–13. https://doi.org/10.1007/BF00332790

Jakubski A.W. 1965. A critical revision of the families Margarodidae and Termitococcidae (Hemiptera, Coccoidea). London: Trustees of the British Museum (Natural History). 187 p.

Jashenko R.V. 1989. Two new species of genus Neomargarodes Green (Coccinea, Margarodidae) from Kazakhstan. Vestnik Zoologii, 6: 14–19. (In Russian).

Jashenko R.V. 1994 Some little known and two new Palearctic species of the genus Drosicha Walker (Coccinea, Margarodidae). Selevinia, 1: 26–39.

Jashenko R.V. 2008. Chervetsy semeystv Ortheziidae i Margarodidae Palearktiki [Palaearctic scale insects of the families Ortheziidae and Margarodidae]. Abstract of Doctor of Science dissertation. Almaty: Istitute of Zoology. 47 p. (In Russian).

Jentsch S. 1936. Ovoviviparie bei einer einheimischen Copeognathenart (Hyperetes guestphalicus). Zoologicher Anzeiger, 116: 287–289.

John O. 1923. Fakultative Viviparität bei Thysanopteren. Entomologische Mitteilungen, 12(5–6): 227–232.

Kalicka-Fijalkowska J. 1928. Le développement of embryonnaire de Margarodes polonicus Ckll. Comptes Rendus des Séances de la Société de Biologie, 99: 1047–1048.

Kerzhner I. & Danzig E. 2001(2002). Hemiptera, Homoptera, Sternorrhyncha. Bollettino di Zoologia Agraria e di Bachicoltura (Milano), 33(3): 217–218.

Kishimoto-Yamada K., Itioka T. & Kawai S. 2005. Biological characterization of the obligate symbiosis between Acropyga sauteri Forel (Hymenoptera: Formicidae) and Eumyrmococcus smithii Silvestri (Hemiptera: Pseudo-coccidae: Rhizoecinae) on Okinawa Island, southern Japan. Journal of Natural History, 39(40): 3501–3524. https://doi.org/10.1080/00222930500393145

Kluge N.J. 2000. Sovremennaya sistematika nasekomykh. I. [Modern systematics of insects. I.] St. Petersburg: Lan’. 336 p. (In Russian).

Kluge N.J. 2005. Larval/pupal leg transformation and a new diagnosis for the taxon Metabola Burmeister, 1832 = Oligoneoptera Martynov, 1923. Russian Entomological Journal, 13(4): 189–229.

Kluge N.J. 2010a. Circumscriptional names of higher taxa in Hexapoda. Bionomina, 1: 15–55. https://doi.org/10.11646/bionomina.1.1.3

Kluge N.J. 2010b. Paradoxical molting process in Orthezia urticae and other coccids (Arthroidignatha, Gallinsecta). Zoosystematica Rossica, 19(2): 246–271.

Köhler G. 1987. Redescription and life cycle properties of Colobopyga coperniciae Ferris, 1952 (Coccoidea: Halimococcidae) – an endemic palm scale from Cuba. Zoologischer Anzeiger, 219: 113–125.

Kollnann J. 1884(1885). Das Überwintern von europäischen Frosch – und Tritonlarven und die Umwandlung des mexicanischen Axolotle. Verhandlungen Naturforschende Gesellschaft (Basel), 7: 387–398.

Kosztarab M. 1996. Scale insects of Northeastern North America. Identification, biology, and distribution. Martinsburg: Virginia Museum of Natural History. 650 p.

Kosztarab M. & Hale D.L. 1968. A new Cryptococcus species from North America, with a key to the species of the genus (Homoptera: Coccoidea). Virginia Journal of Science, 19: 7–11.

Kosztarab M. & Kozár F. 1988. Scale Insects of Central Europe. Budapest. 456 p. https://doi.org/10.1007/978-94-009-4045-1

Kosztarab M. & Watson E. 1994. Phenological, behavioral, and biological observations on a giant scale, Neosteingelia texana Morrison, in Blacksburg, Virginia (Homoptera: Coccinea: Margarodidae). Banisteria, 4: 24–25.

Koteja J. 1974 Comparative studies on the labium in the Coccinea (Homoptera). Zeszyty Naukowe Akademii Rolniczej w Warszawie, Rozprawy Naukowe, 89: 1–162.

Koteja J. 1985 Essay on the prehistory of the scale insects (Homoptera, Coccinea). Annales Zoologici (Warsaw), 38: 461–504.

Koteja J. 1990. Sections: 1.1.6. Paleontology; 1.3.1. Embryonic Development; Ovipary and Vivipary. 1.3.2. Life history. In: Rosen D. (Ed.). Armored scale insects, their tiology, natural enemies and control., 4A: 149–163; 233– 241; 243–254. Amsterdam–Oxford–New York–Tokyo: Elsevier.

Koteja J. 1996. Scale insects (Homoptera: Coccinea) a day after. Thomas Say Publications in Entomology. Proceedings. Studies on Hemipteran Phylogeny. Entomological Society of America Lanham, MD. 244 p.

Koteja J. 2000. Advances in the study of fossil coccids (Hemiptera: Coccinea). Polskie Pismo Entomologiczne, 69: 187–218.

Koteja J. & Azar D. 2008. Scale insects from Lower Cretaceous amber of Lebanon (Hemiptera: Sternorrhyncha: Coccinea). Alavesia, 2: 133–167.

Koteja J., Pyka-Fosciak G., Vogelgesang M. & Szklarzewicz T. 2003. Structure of the ovary in Steingelia (Sternorrhyncha: Coccinea), and its phylogenetic implications. Arthropod Structure & Development, 32: 247–256. https://doi.org/10.1016/S1467-8039(03)00046-X

Koteja J. & Zak-Ogaza B. 1981. The life history of Steingelia gorodetskia Nassonov (Homoptera, Coccinea). Annales Zoologici (Warsaw), 36: 171–186.

Kozár F. 2004. Ortheziidae of the World. Budapest: Plant Protection Institute, Hungarian Academy of Sciences. 525 p.

Kozár F. & Foldi I. 2002. The water scale insects of the mysterious family Carayonemidae (Hemiptera: Coccoidea). Bollettino di Zoologia Agraria e di Bachicoltura (Milano), 33(3): 197–201.

Kozár F., Kaydan M.B., Konczné Benedicty S. & Szita É. 2013. Acanthococcidae and Related Families of the Palaearctic Region. Budapest: Plant Protection Institute, Hungarian Academy of Sciences. 680 p.

Kozár F. & Konczné Benedicty Z. 2007. Rhizoecinae of the world. Budapest: Plant Protection Institute, Hungarian Academy of Sciences. 617 p.

Kozár F. & Konczné Benedicty Z. 2000. Carayonemidae of the Neotropical Region with the descriptions of new genera and species (Homoptera: Coccoidea). Folia Entomologica Hungarica, 61: 71–82.

Kozarzhevskaya E.F. 1968. Technique of preparing slides for scale insect (Homoptera: Coccoidea) identification. Entomologicheskoe Obozrenie, 47: 248–253. (In Russian).

Krassilshchik J. 1893. Zur Entwicklungsgeschichte der Phytophthires. Zoologischer Anzeiger, 413: 69–76.

Lambdin P.L. 2001(2002). Discourse on the classification of the scale insects. Bollettino di Zoologia Agraria e di Bachicoltura (Milano), 33(3): 209–213.

LaPolla J.S., Burwell C., Brady S. & Miller D.R. 2008. A new ortheziid (Hemiptera: Coccoidea) from Australia associated with Acropyga myops Forel (Hymenoptera: formicidae) and a key to Australian Ortheziidae. Zootaxa, 1946: 55–68. https://doi.org/10.11646/zootaxa.1946.1.3

List J.H. 1887. Orthezia cataphracta Shaw. Zeitschrift für Wissenschaft Zoologie, 45: 1–85.

Liu W., Xie Y., Dong J. et al. 2014. Ultra- and microstructure of the female reproductive system of Matsucoccus matsumurae. Arthropod Structure & Development, 43: 243–253. https://doi.org/10.1016/j.asd.2014.02.001

MacGillivray A.D. 1921. The Coccidae. Tables for the Identification of the Subfamilies and Some of the More Important Genera and Species, together with Discussions. Urbana. 502 p. https://doi.org/10.5962/bhl.title.30504

Mukhopadhyay A.K. & Ghose S.K. 1999. Biology of the mealybug Planococcus lilacinus (Cockerell) (Pseudococcidae: Homoptera). Environment & Ecology, 17(2): 464–466.

Mamet J.R. 1942. Pedronia strobilanthis Green redescribed (Homoptera, Coccoidea: Pseudococcidae). Proceedings of the Royal Entomological Society of London, Series B: Taxonomy, 11: 149–152. https://doi.org/10.1111/j.1365-3113.1942.tb01387.x

Mamontova V.A. 2008. Evolutsia, filogenez, sistema tley semeistva lyakhnid (Homoptera, Aphidoidea, Lachnidae) [Evolution, phylogeny and system of Lachnidae (Homoptera, Aphidoidea)]. Kiev: Naukova Dumka. 207 p. (In Russian).

Mamontova V.A. 2012. Tli semeystva lyakhnid (Homoptera, Aphidoidea, Lachnidae) fauny Vostochnoy Evropy I sopredel’nykh territoriy [Aphids of the family Lachnidae (Homoptera, Aphidoidea) of the fauna of Eastern Europe and neighboring territories]. Kiev: Naukova Dumka. 390 p. (In Russian).

Matesova G.Ya. 1957. New species of coccids (Homoptera, Coccoidea) from southeastern Kazakhstan. Entomologicheskoe Obozrenie, 36: 163–174. (In Russian).

Matesova G.Ya. 1958. Notes on the biology of soft and hard scales (Homoptera, Coccoidea) in southeastern Kazakhstan. Trudy Instituta Zoologii Akademii Nauk Kazakhskoy SSR, 8: 130–137. (In Russian).

Matesova G.Ya. 1966. A new genus and species of soft scales (Homoptera, Coccoidea, Pseudococcidae) from Kazakhstan. Vestnik Sel’skohozhaistvennoi Nauki, 9: 98–102. (In Russian).

Matesova G.Ya. 1967. Mealybugs (Homoptera, Coccoidea, Eriococcidae) damaging willows in Kazakhstan. Zoologicheskii Zhurnal, 46: 1193–1202. (In Russian).

Matesova G.Ya. 1968. Coccids (Homoptera, Coccoidea) of east Kazakhstan. Trudy Instituta Zoologii Akademii Nauk Kazakhskoy SSR, 30: 102–129. (In Russian).

Matevossyan A.K. 1977. Karyological peculiarities of the Ararat cochineal, Porphyrophora hamelii Brandt (Coccoidea: Margarodidae). Tsitologiya, 19(6): 704–707. (In Russian).

Mayr E. 1974. Cladistic analysis of cladistic classification? Zeitschrift Zoologische Systematik und Evolutionsforschung, 12(2): 94–128. https://doi.org/10.1111/j.1439-0469.1974.tb00160.x

Mayr E. & Ashlock N.Y. 1991. Principles of systematic zoology, 2nd ed. NewYork: McGrow Hill Book Co. 475 p.

Matsuura Yu., Koga R. & Nikoh N. et al. 2009. Huge symbiotic organs in giant scale insects of the genus Drosicha (Coccoidea: Monophlebidae) harbor flavobacterial and enterobacterial endosymbionts. Zoological Science, 26: 448–456. https://doi.org/10.2108/zsj.26.448

McConnell H.S. 1954. A classification of the coccid family Aclerdidae (Coccoidea, Homoptera). Bulletin of the Maryland Agriculture Experiment Station, A75: 1–121.

McKenzie H.L. 1942a. New species of pine-infesting Margarodidae from California and southwestern United States (Homoptera; Coccoidea; Margarodidae). Microentomology, 7(1), contribution No. 30: 1–18.

McKenzie, H.L. 1942b. Seasonal history of the margarodid scale, Matsucoccus bisetosus Morrison, occurring on ponderosa and jeffrey pines in California (Homoptera; Coccoidea; Margarodidae). Microentomology, 7(1), contribution No. 31: 19–24.

McKenzie H.L. 1943. Notes on Matsucoccus vexillorum Morrison (Homoptera: Coccoidea: Margarodidae). Microentomology, 8(2), contribution No. 40: 53–57.

McKenzie, H.L. 1967 Mealybugs of California with taxonomy, biology, and control of North American species (Homoptera: Coccoidea: Pseudococcidae). Berkeley: University of California Press. 526 p.

Mecznikow E. 1866. Embryologie von Aspidiotus nerii, nebst Bemerkungen über die Entwicklung einiger anderen Hemipteren. Zeitschrift für Wissenschaft Zoologie, 41: 468–500.

Meier R., Kotrba M. & Ferrar P. 1999. Ovoviviparity and viviparity in the Diptera. Biological Reviews, 74: 199–258. https://doi.org/10.1017/S0006323199005320

Mendel Z., Assael F., Zeidan S. & Zehavi A. 1998. Classical biological control of Palaeococcus fuscipennis (Burmeister) (Homoptera: Margarodidae) in Israel. Biological Control, 12: 151–157. https://doi.org/10.1006/bcon.1998.0621

Meyen S.V. 1987. Fundamental of palaeobotany. London; NewYork: Chapman & Hall. 432 p. https://doi.org/10.1007/978-94-009-3151-0

Miller D.R. & Miller G.L. 1993. A new species of Puto and a preliminary analysis of the phylogenetic position of the Puto group within the Coccoidea (Homoptera: Pseudococcidae). Jeffersoniana: Contributions from the Virginia Museum of Natural History, 4: 1–35.

Miller D.R. & Williams D.J. 1995. Systematic revision of the Family Micrococcidae (Homoptera: Coccoidea), with a discussion of its relationships, and a description of a gynandromorph. Bollettino del Laboratorio di Entomologia Agraria ‘Filippo Silvestri’ (Portici), 50: 199–247.

Moharana S. 1990. Cytotaxonomy of Coccids (Homoptera: Coccoidea). In: Koteja J. (Ed.). Proceedings of the Sixth International Symposium of Scale Insect Studies, Part II. Cracow, Poland: August 6–12, 1990: 47–54. Cracow: Agricultural University Press.

Morales C.F. 1991. Margarodidae (Insecta: Hemiptera). Fauna of New Zealand, 21: 123 p.

Mordvilko A.K. 1914. Aphidoidea. Vypusk 1 [Aphidoidea. Part 1]. Fauna Rissii i sopredel’nykh stran preimushchestvenno po kollektsiyam Zoologicheskogo muzeya Imperskoy Akademii Nauk. Nasekomye poluzhestkokrylye (Insecta: Hemiptera). Tom 1 [Fauna of Russia and adjacent countries, mainly based on collections of the Zoologocal Museum of the Imperial Academy of Sciences. Hemipterans. Vol. I]. Petrograd: Imperatorskaya Akademiya Nauk. clxv+236+9 p.

Morrison H. 1925. Classification of scale insects of the subfamily Ortheziinae. Journal of Agricultural Research, 30: 97–154.

Morrison H. 1927. Descriptions of new genera and species belonging to the coccid family Margarodidae. Proceedings of the Biological Society of Washington, 40: 99–109.

Morrison H. 1928. A classification of the higher groups and genera of the coccid family Margarodidae. United States Department of Agriculture Technical Bulletin, 52: 1–239.

Morrison H. 1952. Classification of the Ortheziidae. Supplement to classification of scale insects of the subfamily Ortheziinae. United States Department of Agriculture. Technical Bulletin, 1052: 1–80.

Morrison H. & Morrison E.R. 1922. A redescription of the type species of the genera of Coccidae based on species originally described by Maskell. Proceedings of the United States National Museum (Washington), 60: 1–120. https://doi.org/10.5479/si.00963801.60-2407.1

Narasimham A.U. & Chacko M.J. 1988. Rastrococcus spp. (Hemiptera: Pseudococcidae) and their natural enemies in India as potential biological control agents for R. invadens Williams. Bulletin of Entomological Research, 78: 703–708. https://doi.org/10.1017/S000748530001556X

Nokkala S., Kuznetsova V.G., Maryanska-Nadachowska A. & Nokkala C. 2004. Holocentric chromosomes in meiosis. I. Restriction of the number of chiasmata in bivalents. Chromosome Research, 12: 733–739. https://doi.org/10.1023/B:CHRO.0000045797.74375.70

Nokkala S., Kuznetsova V.G., Maryanska-Nadachowska A. & Nokkala C. 2006. Holocentric chromosomes in meiosis. II. The modes of orientation and segregation of trivalents. Chromosome Research, 14: 559–565. https://doi.org/10.1007/s10577-006-1053-6

Nur U. 1979. Gonoid thelytoky in soft scale insects (Coccidae: Homoptera). Chromosoma (Berlin), 72: 89–104. https://doi.org/10.1007/BF00286431

Nur U. 1980. Evolution of unusual chromosome systems in scale insects (Coccoidea: Homoptera). In: Blackman R.L., Hewitt G.M. & Ashburner M. (Eds.) Insect Cytogenetics. 97–117. Royal Entomological Society, London.

Nur U. 1990. Parthenogenesis. In: Rosen D. (Ed.). Armored scale insects, their tiology, natural enemies and control., 4A: 191–196. Amsterdam–Oxford–New York–Tokyo: Elsevier.

Oguma K. 1919. A new scale-insect, Xylococcus alni, on alder, with special reference to its metamorphosis and anatomy. Journal of the College of Agriculture, Hokkaido Imperial University, 8: 77–109.

Oliveira C.M., Frizzas M.R., Fialho J.F. & Gullan P.J. 2008. Biology of Protortonia navesi (Hemiptera: Monophlebidae), a new Cassava pest in Brazil, with notes on its behavior in the field. Annals of the Entomological Society of America, 101(4): 779–785. https://doi.org/10.1093/aesa/101.4.779

Parida B.B. & Moharana S. 1982. Studies on the chromosome constitution in 42 species of scale insects (Coccoidea: Homoptera) from India. Chromosome Information Service (Tokyo), 32: 18–20.

Pearse A.S. 1936. Zoological names. A list of phyla, classes, and orders. Durham: Duke University Press. 24 p.

Pesson P. 1939. Une glande exsertile ventrale chez les coccides. Bulletin de la Société Zoologique de France, 64: 144–151.

Pesson P. 1951. Ordre des Homoptéres (Homoptera Leach, 1815) In: P.P. Grassé (Ed.). Traité de Zoologie: Anatomie, Systématique, Biologie. 10: 976–1948. Paris: Masson et Cie.

Popov Yu.A. 1980. Superorder Cimicidea Laicharting, 1781. In: Rohdendorf B.B. & Rasnitsyn A.P. (Eds). Istoricheskoe razvitie klassa Nasekomykh [Historical development of the class Insecta]. Trudy Paleontologicheskogo Instituta Academii Nauk SSSR, 175: 58–69.

Popova A.A. 1967. Tipy prisposobleniy tley k pitaniyu na kormovykh rasteniyakh [Types of aphid adaptations to host plants]. Leningrad: Nauka. 291 p.

Rasnytsyn A.P. 1989. Phytospreading from the selectionist’s viewpoint. Zhurnal Obshchey Biologii, 50: 581–583. (In Russian). https://doi.org/10.1111/j.1751-1097.1989.tb05568.x

Rasnytsyn A.P. 2010. Molecular phylogenetics, morphological cladistics and fossils. Entomological Review, 89(1): 85–132.

Razak T.A., Ananthi D.V. & Jayanthi R. 1994. Biology of the sugarcane mealybug, Dysmicoccus carens Williams (Homoptera: Pseudococcidae). Journal of Entomological Research (New Delhi), 18(2): 169–174.

Reyne A. 1954. A redescription of Puto antennatus Sign. (Homoptera, Coccidea). With notes on Ceroputo pilosellae Sŭlc and Macrocerococcus superbus Leon. Zoologische Mededelingen Uitgegeven door het Rizksmuseum van Natuurlijke Historie te Leiden, 32: 291–324.

Reyne A. 1957. Walkeriana tosariensis, a new monophlebine coccid from Casuarina junghununiana Miq. in East Java. Beaufortia (Amsterdam), 6: 115–145.

Reyne, A. 1965. Observations on some Indonesian scale insects. Tijdschrift voor Entomologie, 108: 145–188.

Richard C. 1971. Contribution à l’étude morphologique et biologique des Stictococcinae (Hom. Coccoidea). Annales de la Société Entomologique de France, 7: 571–609.

Richard C. 1986. Carayonemidae famille nouvelle Carayonema orousseti, n. gen., n. sp. de Guyane française (Homoptera, Coccoidea). Annales de la Société Entomologique de France, 22(2): 268–273.

Richards A.M. 1985. Biology and defensive adaptations in Rodatus major (Coleoptera: Coccinellidae) and its prey, Monophlebulus pilosior (Hemiptera: Margarodidae). Journal of Zoology (London) (A), 205: 287–295. https://doi.org/10.1111/j.1469-7998.1985.tb03535.x

Rohdendorf B.B. 1977. The rationalization of names of higher taxa in zoology. Paleontological Journal, 11: 149–155.

Romeis B. 1953. Mikroskopicheskaya tekhnika [Microscopic technics]. Moscow. 718 p. (In Russian).

Royer M. 1966. Développement comparé des individus hermaphrodites et des mâles chez Icerya purchasi (Mask.), Homoptère. Bulletin de la Société Zoologique de France, 91: 341.

Royer M. 1973a. Rôle de la polyspermie dans la formation des lignées cellulaires haploid et diploide chez l’insecte Icerya purchasi. Annales d’Embryologie et de Morphogénèse, 6: 243–252.

Royer M. 1973b. La formation des ébauches gonadiques dans l’embryon de l’insecte hermaphrodite Icerya purchasi Mask. Comptes Rendus de l’Academie des Sciences Paris (Series D), 276: 1605–1608.

Rosas-Pérez T., Rosenblueth M., Rincón-Rosales R., Mora J. & Martinez–Romero E. 2014. Genome Sequence of “Candidatus Walczuchella monophlebidarum” the Flavobacterial Endosymbiont of Llaveia axin axin (Hemiptera: Coccoidea: Monophlebidae). Genome Biology and Evolution, 6(3): 714–726. https://doi.org/10.1093/gbe/evu049

Savage R.E. 1914. The respiratory system of Monophlebus stebbingi, var. octocaudata. Bulletin of Entomological Research, 5: 45–47. https://doi.org/10.1017/S0007485300034520

Shcherbakov D.E. 1990. Extinct four-winged ancestors of scale insects (Homoptera: Sternorrhyncha). In: Koteja J. (Ed.). Proceedings of the Sixth International Symposium of Scale Insect Studies, Part II. Cracow, Poland: August 6–12, 1990: 23–29. Cracow: Agricultural University Press.

Shcherbakov D.E. 2007. Extinct four-winged precoccids and the ancestry of scale insects and aphids (Hemiptera). Russian Entomological Journal, 16(1): 47–62.

Shcherbakov D.E. & Popov Yu.A. 2002. Superorder Cimicidea. In: Rasnitsyn A.P & Quicke D.L.J. (Eds). History of insects. 143–157. New York, Boston, Dordrecht, London, Moscow.

Schrader F. 1930. Observations on the biology of Protortonia primitiva (Coccidae). Annals of the Entomological Society of America, 23: 126–132. https://doi.org/10.1093/aesa/23.1.126

Schrader F. 1931. The chromosome cycle of Protortonia primitiva (Coccidae) and a consideration of the meiotic division apparatus in the male. Zeitschrift für Wissenschaftliche Zoologie, 138: 386–408.

Silvestri F. 1924. A new myrmecophilous genus of Coccidae (Hemiptera) from India. Record of the Indian Museum, 26: 311–315.

Silvestri F. 1926. Descrizione di un novo genere di Coccidae (Hemiptera) mirmecofilo della cina. Bollettino del Laboratorio di Zoologia Generale e Agraria della R. Scuola Superior Agricoltura (Portici), 18: 271–275.

Simpson G.G. 1961. Principles of animal taxonomy. New York: Columbia University Press. 247 p. https://doi.org/10.7312/simp92414

Song N., Liang A.P. & Bu C.P. 2012. A molecular phylogeny of Hemiptera inferred from mitochondrial genome sequences. PLoS ONE, 7 (11): e48778. https://doi.org/10.1371/journal.pone.0048778

Stickney F.S. 1934. The external anatomy of the red date scale Phoenicococcus marlatti Cockerell, and its allies. United States Department of Agriculture Technical Bulletin, 404: 1–162.

Szklarzewicz T. 1997. Structure and development of the telotrophic ovariole in ensign scale insects (Hemiptera, Coccomorpha, Ortheziidae). Tissue and Cell, 29: 31–38. https://doi.org/10.1016/S0040-8166(97)80069-9

Szklarzewicz T. 1998a. Structure of ovaries of scale insects: II. Margarodidae (Insecta, Hemiptera, Coccinea). International Journal of Insect Morphology and Embryology, 27(4): 319–324. https://doi.org/10.1016/S0020-7322(98)00024-5

Szklarzewicz T. 1998b. The ovaries of scale insects (Hemiptera, Coccinea). Morphology and phylogenetic conclusions. Folia Histochemica et Cytobiologica, 36(4): 157–165.

Szklarzewicz T., Kedra K. & Niznik S. 2005. Ultrastructural studies of the ovary of Palaeococcus fuscipennis (Burmeister) (Insecta, Hemiptera, Coccinea: Monophlebidae). Folia Biologica (Krakow), 53(1–2): 45–50. https://doi.org/10.3409/1734916054663483

Szklarzewicz T., Kalandyk-Kolodziejczyk M., Kot M., Michalik A. 2011(2013). Ovary structure and transovarial transmission of endosymbiotic microorganisms in Marchalina hellenica (Insecta, Hemiptera, Coccomorpha: Marchalinidae). Acta Zoologica (Stockholm), 944(2): 184–192. https://doi.org/10.1111/j.1463-6395.2011.00538.x

Szklarzevicz T., Michalik A., Kalandyk-Kolodziejczyk M. et al. 2014. Ovary of Matsucoccus pini (Insecta, Hemiptera, Coccinea: Matsucoccidae): morphology, ultrastructure, and phylogenetic implications. Microscopy Research and Technique, 77: 327–334. https://doi.org/10.1002/jemt.22347

Tang F.T. 1992. The Pseudococcidae of China. Shanxi. 768 p. (In Chinese with partial English translation).

Tait S.M., Dahlsten D.L., Gill R.J., Doyen J.T. 1990. Life history of the incense cedar scale, Xylococculus macrocarpae (Homoptera: Margarodidae), on incense cedar in California with a description of the larvae of one of its common predators Eronyxa expansus Van Dyke (Coleoptera: Trogositidae). Hilgardia, 58(2): 1–19. https://doi.org/10.3733/hilg.v58n02p019

Ter-Grigorian M.A. 1956. Coccids of cultivated fruits in Armenia. In: Avetian A.S. (Ed.). Materialy po izucheniyu fauny Armyanskoy SSR [Material on the examination of the faune Armenian SSR]. Zoologicheskij Sbornik, 9: 33–57. (In Russian).

Ter-Grigorian M.A. 1973. Muchnistye chervetsy (Pseudococcidae) [Mealybugs (Pseudococcidae)]. Fauna of the Armenian SSR. Erevan: Akademiya Nauk Armianskoy SSR. 246 p. (In Russian).

Theron J.G. 1958. Comparative studies on the morphology of male scale insects (Hemiptera: Coccoidea). Annals of the University of Stellenbosch (Section A), 34: 1–71.

Theron J.G. 1962. Structure and relationships of the male of Phenacoleachia zealandica (Maskell) (Hemiptera: Coccoidea). Proceedings of the Royal Entomological Society of London, Series A: General Entomology, 37: 145–153. https://doi.org/10.1111/j.1365-3032.1962.tb00005.x

Trapeznikova I.V. & Gavrilov I.A. 2008. About ovoviviparity in mealybugs (Homoptera: Coccinea: Pseudococcidae). Trudy Zoologicheskogo Instituta RAS, 312: 43–53. (In Russian).

Tremblay E. 1997. Embryonic development; oviparity and viviparity. In: Ben-Dov Y. & Hodgson C.J. Soft scale insects: their biology, natural enemies and control, 7A: 257-260. Amsterdam–Oxford–New York–Tokyo: Elsevier. https://doi.org/10.1016/S1572-4379(97)80055-3

Unruh C.M. & Gullan P.J. 2007. Molecular data reveal convergent reproductive strategies in iceryine scale insects (Hemiptera: Coccoidea: Monophlebidae), allowing the re-interpretation of morphology and a revised generic classification. Systematic Entomology, 33: 8–50. https://doi.org/10.1111/j.1365-3113.2007.00404.x

Unruh C.M. & Gullan P.J. 2008. Identification guide to species in the scale insect tribe Iceryini (Coccoidea: Monophlebidae). Zootaxa, 1803: 1–106. https://doi.org/10.11646/zootaxa.1803.1.1

Vayssière P. 1926. Contribution à l’étude biologique et systématique des Coccidae. Annales des Epiphyties, 12: 187–382.

Vayssière P. 1932 Monophlebines et pseudococcines de l’Afrique du Nord. Hem. Coccidae. Bulletin de la Société Entomologique de France, 37: 24–28.

Vayssière P. & Hughes-Schrader S. 1948. Étude morphologique et biologique Nautococcus schraderae Vayss. (Coccoidea–Margarodidae). Mémoires du Museum National d’Histoire Naturelle (N.S.), Serie A, Zoologie, 26: 57–74.

Vea I.M. 2014. Morphology of the males of seven species of Ortheziidae (Hemiptera: Coccoidea). American Museum Novitates (New York), 3812: 1–36 https://doi.org/10.1206/3812.1

Vea I. & Gimaldi D.A. 2012. Phylogeny of ensign scale insects (Hemiptera:Coccoidea: Ortheziidae) based on the morphology of Recent and Fossil Females. Systematic Entomology, 37: 758–783. https://doi.org/10.1111/j.1365-3113.2012.00638.x

Vershinina A.O. & Kuznetsova V.G. 2016. Parthenogenesis in Hexapoda: Entognatha and non–holometabolous insects. Journal of Zoological Systematics and Evolutional Research, 54(4): 257–268. https://doi.org/10.1111/jzs.12141

von Dohlen C.D. & Moran N.A. 1995. Molecular phylogeny of the Homoptera: a paraphyletic taxon. Journal of Molecular Evolution, 41: 211–223. https://doi.org/10.1007/BF00170675

Wagner N.P. 1862. Samoproizvol’noe razmnozhenie gusenits u nasehomykh. [Spontaneous reproduction in insect larvae]. Kazan’. 50 p. (In Russian).

Walczuch A. 1932. Studien an Coccidensymbioten. Zeitschrift für Morphologie und Ökologie der Tiere, 25: 623–729. https://doi.org/10.1007/BF00419299

Watson W.Y., Underwood G.R. & Reid J. 1960. Notes on Matsucoccus macrocicatrices Richards (Homoptera: Margarodidae) and its association with Septobasidium pinicola Snell in eastern Canada. Canadian Entomologist, 92: 662–667. https://doi.org/10.4039/Ent92662-9

Westwood J.O. 1838. The entomologist’s text book; an introduction to the natural history, structure, physiology, and classification of insects. London: Wm.S. Orr and Co., Amen Corner, Paternoster Row. 432 p. https://doi.org/10.5962/bhl.title.23744

Westwood J.O. 1845. Plate VI. Illustrations of various species of Coccidae, belonging to the genus Monophlebus. Arcana Entomologica, 1: 21–22.

White M.J.D. 1973. Animal Cytology and Evolution. Cambridge. 961 p.

Williams D.J. 1969. The family-group names of the scale insects (Hemiptera: Coccoidea). Bulletin of the British Museum (Natural History), Entomology, 23: 315–341. https://doi.org/10.5962/bhl.part.15136

Williams D.J. 1978. The anomalous ant-attended mealybugs (Homoptera: Pseudococcidae) of South-East Asia. Bulletin of the British Museum (Natural History), Entomology, 37: 1–72.

Williams D.J. 1986(1987). Unusual instars in Xenococcus annandalei Silvestri (Homoptera: Pseudococcidae). Bollettino del Laboratorio di Entomologia Agraria ‘Filippo Silvestri’, 43 (Suppl.): 73–77.

Williams D.J. 1998. Mealybugs of the genera Eumyrmococcus Silvestri and Xenococcus Silvestri associated with the ant genus Acropyga Roger and a review of the subfamily Rhizoecinae (Hemiptera, Coccoidea, Pseudococcidae). Bulletin of the British Museum (Natural History), Entomology, 67: 1–64.

Williams D.J. 2004. Mealybugs of Southern Asia. Kuala Lumpur: Southdene SDN. 896 p.

Williams D.J. & Granara de Willink M.C. 1992. Mealybugs of Central and South America. London: CAB International. 635 p.

Williams D.J. & Gullan P.J. 2008. A revision of the Neotropical scale insect genus Protortonia Townsend (Hemiptera: Coccoidea: Monophlebidae: Llaveiini). Journal of Natural History, 42(1–2): 77–128. https://doi.org/10.1080/00222930701838054

Williams D.J. & Watson G.W. 1990. The Scale Insects of the Tropical South Pacific Region. Pt. 3: The Soft Scales (Coccidae) and Other Families. Wallingford: CAB International. 267 p.

Wojciechowski W. 1992. Studies on the systematic system of aphids (Homoptera, Aphidinea). Katowice. 75 p.

Wu S.A. 2008. Morphology of Kuwania bipora Borchsenius (Hemiptera: Coccoidea: Margarodidae). Entomotaxonomia, 30(3): 207–2014.

Wu S.A. & Nan N. 2012. Neogreenia lonicera sp. nov., a new species of Margarodidae sensu lato (Hemiptera: Coccoidea) from China, with a key to species of Neogreenia MacGillivray and placement of the genus in the family Kuwaniidae. Zootaxa, 3274: 43–54. https://doi.org/10.11646/zootaxa.3274.1.5

Wu S.A., Nan N., Gullan P. & Deng J. 2013. The taxonomy of the Japanese oak red scale insect, Kuwania quercus (Kuwana) (Hemiptera: Coccoidea: Kuwaniidae), with a generic diagnosis, a key to species and description of a new species from California. Zootaxa, 3630(2): 291–307. https://doi.org/10.11646/zootaxa.3630.2.5

Zakhvatkin Yu. A. 1975. Embriologiya nasekomykh [Embryology of insects]. Moscow. 328 p. (In Russian).

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