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Scale insects (Hemiptera: Sternorrhyncha: Coccomorpha) feed on plant sap; some are damaging pests in agriculture and forestry. However, the scale insects found in continental Africa have not been extensively studied and the available means of identification are incomplete. This synoptic work, the second in this series, covers the six archaeococcoid scale insect families found in continental Africa: Kuwaniidae, Margarodidae, Matsucoccidae, Monophlebidae, Ortheziidae and Putoidae. The work provides identification keys to a total of 29 genera and 137 species, and a checklist for each family. Pseudaspidoproctus zimmermanni (Newstead) (Monophlebidae) is transferred to Aspidoproctus, as Aspidoproctus zimmermanni (Newstead), comb. n. Praelongorthezia praelonga (Douglas) (Ortheziidae) is recorded from Gambia for the first time.

Las cochinillas de las raíces del café son una plaga limitante en la caficultura colombiana y Puto barberi (Cockerell, 1895) (Hemiptera: Putoidae) es la especie que prevalece. Con el objetivo de proponer una estrategia para el control de esta plaga, se evaluaron en el laboratorio las cepas de Metarhizium anisopliae Ma 9236 y M. robertsii a concentraciones de 1x107 conidias/mL, con mortalidades entre 80% y 84% sobre P. barberi. También se evaluaron en plántulas de café infestadas con la plaga en tres ensayos. En el primero se emplearon plantas de 4 a 6 meses de edad, infestadas con hembras oviplenas de P. barberi, tratadas con 50 cm3 de: 1. M. anisopliae, 2. M. robertsii, ambos a 2x1010 conidias/L, y 3. agua como control. Con el agua, el porcentaje de infestación fue de 100% y se encontraron 25 cochinillas por planta. La aplicación de M. anisopliae disminuyó en 10% la infestación de la cochinilla y redujo la población en 55%. M. robertsii disminuyó la infestación en 40% y la población del insecto en 86%. En el ensayo 2 se comparó M. robertsii vs. agua con resultados similares al ensayo 1 y con una protección del 50% de las plantas. En el ensayo 3, los tratamientos correspondieron a: 1. M. robertsii (2x1010 conidias/L), 2. azadiractina 6% polvo soluble en agua (3 g/L), 3. clorpirifos 75% WG (3 g/L) y 4. Agua. En el tratamiento control agua el porcentaje de infestación fue del 65%, en los demás tratamientos las infestaciones estuvieron entre 11% y 22% difiriendo del control, pero siendo estadísticamente similares entre sí. M. robertsii es un candidato para ser evaluado en almácigos comerciales y campo.

Family PUTOIDAE Tang Common names: Giant mealybugs or putoids. Background: This family consists of one extant genus containing 47 species (and possibly a second, monotypic genus known only from a fossil) (García Morales et al. 2016). It was regarded as part of the family Pseudococcidae until its elevation to family status by Hodgson and Foldi (2006), based mostly on adult male morphology (the male having a row of 8–15 (usually 14) simple eyes surrounding the head, whereas in Pseudococcidae and Rhizoecidae there are usually only 4). Molecular evidence supports its familial status separate from the Pseudococcidae (Gullan & Cook 2007; Choi & Lee 2021). The morphology of adult female putoids is like that of mealybugs (Pseudococcidae) except for the presence of three or four campaniform sensilla on each surface of each trochanter (Fig. 31) (Pseudococcidae and Rhizoecidae normally have only two), and often a pair of denticles on the base of each claw (absent in Pseudococcidae and Rhizoecidae). The status of the Putoidae remains controversial. Putoids occur in the Nearctic, Neotropical, Oriental and Palaearctic regions; they are absent from the Afrotropical and Australasian regions (García Morales et al. 2016). Appearance in life: Putoids are found on all parts of a wide variety of plants, even occasionally on the subterranean crown of grasses. Adult female quite large, about 5.0 mm long, covered with thick tufts of mealy white wax (often with a central ridge) and broad lateral wax filaments; when the wax is removed, the body has two longitudinal grey-to-green dorsal stripes. Legs and antennae well developed and dark brown. Most species do not produce an ovisac. Identification: As recommended for Pseudococcidae by Sirisena et al. (2013), the best specimens for identification are young adult females just after the final moult, before the body has become distended with developing eggs. Body about 5.0 mm long, broadly oval, with large cerarii, ostioles, swirled trilocular pores and well-developed antennae and legs; trochanters each with three or four placoid sensilla on each surface; and claws often with a basal denticle in addition to a subapical denticle; translucent pores present on hind legs. Circulus usually present (Miller et al. 2014). Economic importance: None in Africa. Biology: Putoids reproduce sexually; there are four instars in the female and five in the male (Miller et al. 2014). Usually there is one generation per year, although in a few species a generation can take several years. Early instars overwinter under bark before migration to twigs and leaf bases in early spring. Adults appear in late spring/ early summer, and males are common. Eggs are laid just after midsummer. Putoids feed on a wide range of hosts, particularly on Pinophyta, Poaceae and some woody shrubs, attacking all plant parts (including the subterranean crown of Poaceae in some Nearctic species) (Miller et al. 2014). Checklist and distributions of Putoidae in continental Africa (1 genus, 3 species) Puto peyerimhoffi (Vayssière, 1923): Algeria Puto subericola (Vayssière, 1927): Morocco Puto superbus (Leonardi, 1907): Algeria, Morocco, Tunisia, Published as part of Watson, Gillian W., 2022, Towards identification of the scale insects (Hemiptera: Coccomorpha) of continental Africa: 2. Checklists and keys to six archaeococcoid families, pp. 301-356 in Zootaxa 5105 (3) on pages 350-352, DOI: 10.11646/zootaxa.5105.3.1, http://zenodo.org/record/6332761, {"references":["Garcia 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 (accessed 12 January 2022) https: // doi. org / 10.1093 / database / bav 118.","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), 1 - 250. https: // doi. org / 10.11646 / zootaxa. 1263.1.1","Gullan, P. J. & Cook, L. G. (2007) Phylogeny and higher classification of the scale insects (Hemiptera: Sternorrhyncha: Coccoidea). Zootaxa, 1668 (1), 413 - 425. https: // doi. org / 10.11646 / zootaxa. 1668.1.22","Danzig, E. M. & Gavrilov-Zimin, I. A. (2014) Palaearctic mealybugs (Homoptera: Coccinea: Pseudococcidae). Part 1. Subfamily Phenacoccinae. Russian Academy of Sciences, Zoological Institute, St. Petersburg, 678 pp.","Sirisena, U. G. A. I., Watson, G. W., Hemachandra, K. S. & Wijayagunasekara, H. N. P. (2013) A modified technique for the preparation of specimens of Sternorrhyncha for taxonomic studies. Tropical Agricultural Research Journal, 24 (2), 139 - 149.","Miller, D., Rung, A., Parikh, G., Venable, G., Redford, A. J., Evans, G. A. & Gill, R. J. (2014) Scale Insects, Edition 2. USDA APHIS Identification Technology Program (ITP), Fort Collins, Colorado. Available from: http: // idtools. org / id / scales / factsheet. php? name = 6726 (accessed 15 February 2022)"]}

Scale insects (Hemiptera: Sternorrhyncha: Coccomorpha) are obligate plant parasites feeding on plant sap; some are damaging pests in agriculture, horticulture and forestry. Despite their economic importance, the scale insects found in continental Africa have not been extensively studied and the keys for identifying them are incomplete and scattered through the literature in several languages. The aim of this study is to improve our understanding of the African scale insect fauna. As a first step towards their identification, we provide a key to the 23 families currently known from continental Africa, based on slide-mounted adult females, covering Aclerdidae, Asterolecaniidae, Cerococcidae, Coccidae, Conchaspididae, Dactylopiidae, Diaspididae, Eriococcidae, Halimococcidae, Kermesidae, Kerriidae, Kuwaniidae, Lecanodiaspididae, Margarodidae, Matsucoccidae, Micrococcidae, Monophlebidae, Ortheziidae, Phoenicococcidae, Pseudococcidae, Putoidae, Rhizoecidae and Stictococcidae.

Puto barberi (Hemiptera: Putoidae) y Toumeyella coffeae (Hemiptera: Coccidae) son las especies de insectos más limitantes en el establecimiento de cafetales en Norte de Santander, Colombia. Con el fin de identificar las plantas hospedantes de estas plagas, se realizó un diagnóstico en fincas seleccionadas aleatoriamente a partir de zonas previamente reportadas con estos problemas fitosanitarios. En cada finca se trazaron transectos, donde se desenterraron las arvenses y se registró el número de plantas por morfoespecie y la presencia de P. barberi y T. coffeae en cada una. Para la identificación de las arvenses se preservaron individuos por morfoespecie botánica y los insectos se depositaron en viales con etanol al 76 %. Se trazaron 85 transectos, se evaluaron 6.880 plantas, pertenecientes a 22 familias, 37 géneros y 38 especies. Puto barberi se registró en 20 de las 38 especies, con mayor presencia en Eirmocephala brachiata, Sida rhombifolia, Conyza bonariensis, Amaranthus hybridus, Leonurus japonicus y Emilia sonchifolia con valores entre 14,0 % y 30,0 %. Toumeyella coffeae se registró en nueve especies, siendo Erato vulcanica (11,43 %), Leucaena leucocephala (11,11 %) y E. sonchifolia (8,87 %) los hospedantes más frecuentes. Al relacionar las especies de plantas más comunes en la caficultura de Norte de Santander, así como las mayores presencias de las dos plagas en campo, se concluye que S. rhombifolia, E. sonchifolia y C. bonariensis albergan las mayores poblaciones de P. barberi y T. coffeae. El presente estudio genera las bases para evaluar y diseñar estrategias de control cultural y agroecológico de P. barberi y T. coffeae.

Coffee is one of the most important Brazilian agricultural commodities exported, and Minas Gerais and Espírito Santo States are the main coffee producers. Scale insects are important coffee pests, and 73 species of Cerococcidae (3), Coccidae (18), Diaspididae (6), Eriococcidae (1), Ortheziidae (3), Pseudococcidae (21), Putoidae (2) and Rhizoecidae (19) have been associated with roots, branches, leaves, flowers and fruits of Arabica coffee in the Neotropics. Eight species were found associated with Arabica coffee in Minas Gerais and Espírito Santo States in this study, and Coccidae was the most frequent family. Coccus alpinus, Cc. celatus, Cc. lizeri, Cc. viridis, and Saissetia coffeae (Coccidae) were found in both states; Alecanochiton marquesi, Pseudaonidia trilobitiformis (Diaspididae), and Dysmicoccus texensis (Pseudococcidae) were only found in Minas Gerais. Alecanochiton marquesi and P. trilobitiformis are first reported in Minas Gerais, and Cc. alpinus in Espírito Santo, on Arabica coffee. All scale insect species were associated with coffee leaves and branches, except D. texensis, associated with coffee roots. Fourty seven scale insect species have been found occurring in Brazilian Arabica coffee, and in Espírito Santo (28) and Minas Gerais (23). Widespread and geographical distribution of each species found are discussed.

The Scale Insects, infraorder Coccomorpha (Hemiptera: Sternorryncha) includes many agricultural pests. The latest check list of scale insects of Sakartvelo (Republic of Georgia) was published in 2008, but it contained species mostly from the city of Tbilisi. In this study an updated check list of scale insects of Sakartvelo with 242 species is presented. Fifteen scale insect families from 30 extant families in the world are presented for Sakartvelo. Most species-rich families are: Diaspididae with 90 species belonging to 38 genera, Pseudococcidae with 72 species belonging to 31 genera and Coccidae with 43 species belonging to 23 genera. Other families are: Acanthococcidae (5 genera, 10 species), Asterolecaniidae (3 genera, 5 species), Cryptococcidae (2 genera, 3 species), Eriococcidae (1 genus, 2 species), Kermesidae (1 genus, 3 species), Kerriidae (1 genus, 1 species), Kuwaniidae (1 genus, 1 species), Marchalinidae (1 genus, 1 species), Margarodidae (2 genera, 3 species), Monophlebidae (2 genera, 2 species), Ortheziidae (3 genera, 5 species), Putoidae (1 genus, 1 species). Anophococcus agropyri was found in the collection of Z. Hadzibeyli and it is herein recorded for the first time from Georgia. Distribution and host plant data for each species is given.

Resumen Objetivos: En esta investigación se estudiaron las chinches harinosas (Hemiptera: Pseudococcidae y Putoidae) presentes en las raíces de café (Coffea arabica L.) y de arvenses asociadas a este cultivo. Alcance: En los municipios de Sandoná, Consacá, La Unión, San Pablo y San José de Albán, departamento de Nariño, Colombia. Metodología: Se recogieron 114 muestras a partir de raíces de café y de 17 especies de plantas arvenses asociadas a este cultivo, pertenecientes a las familias botánicas Asclepidaceae, Asteraceae, Commelinaceae, Cyperaceae, Poaceae, Plantaginaceae, Polygonaceae, Solanaceae. Los insectos se colectaron de forma manual, en raíces de plantas que presentaran alguna sintomatología asociada al ataque de estos insectos tales como clorosis y debilitamiento. La identificación taxonómica se realizó a partir de ejemplares montados en lámina para microscopía después de diferentes procesos, con base en la comparación de la morfología externa de las hembras adultas con las claves establecidas para el grupo. Principales resultados: Se identificaron por primera vez en Nariño en el cultivo de café, dos especies pertenecientes a los géneros Puto Signoret (Putoidae) y Dysmicoccus Ferris (Pseudococcidae). Conclusión: Esta información es importante para incorporarla dentro de planes de manejo integrado de plagas en zonas productoras de café en Nariño y en Colombia. Con este trabajo se aumenta el conocimiento en la distribución de las especies de insectos escama del café en el país. Abstract Objectives: The mealybugs (Hemiptera: Pseudococcidae and Putoidae) present in roots of coffee (Coffea arabica L.) and herbaceous plants associated to this crop were studied in this research. Scope: This research was carried out in Sandoná, Consacá, La Union, San Pablo and San José de Albán, municipalities of the Department of Nariño. Methodology: A total of 114 samples were collected from roots of coffee and 17 species of herbaceous plants associated with this crop, belonging to the botanical families Asclepidaceae, Asteraceae, Commelinaceae, Cyperaceae, Poaceae, Plantaginaceae, Polygonaceae and Solanaceae. The specimens were collected manually in roots of plants that presented some symptomatology such as chlorosis and weakening associated with the attack of these insects. The taxonomic identification was made from specimens placed in microscope slides after different processes based on the comparison of the external morphology of the adult female with de taxonomic codes established for the group. Main results: Two species belonging to the genus Puto Signoret (Putoidae family) and Dysmicoccus Ferris (Pseudococcidae family) were identified for the first time in coffee crops in Nariño. Conclusion: This information is important to be incorporated into integrated pest management plans in coffee producing areas in Nariño and in Colombia. The knowledge in the distribution of the species of mealybugs of coffee in the country is enriched with this work.

In the last decades, several expeditions were organized in Europe by the researchers of the Hungarian Natural History Museum to collect snails, aquatic insects and soil animals (mites, springtails, nematodes, and earthworms). In this study, scale insect (Hemiptera: Coccomorpha) specimens extracted from Hungarian Natural History Museum soil samples (2970 samples in total), all of which were collected using soil and litter sampling devices, and extracted by Berlese funnel, were examined. From these samples, 43 scale insect species (Acanthococcidae 4, Coccidae 2, Micrococcidae 1, Ortheziidae 7, Pseudococcidae 21, Putoidae 1 and Rhizoecidae 7) were found in 16 European countries. In addition, a new species belonging to the family Pseudococcidae, Brevennia larvalis Kaydan, sp. n. and a new species of Ortheziidae, Ortheziola editae Szita & Konczné Benedicty, sp. n. are described and illustrated based on the adult female stage. Revised keys to the adult females of Brevennia and Ortheziola are presented. © Mehmet Bora Kaydan et al.

The scale insect Puto superbus (Putoidae) lives in mutualistic symbiotic association with bacteria. Molecular phylogenetic analyses have revealed that symbionts of P. superbus belong to the gammaproteobacterial genus Sodalis. In the adult females, symbionts occur both in the bacteriocytes constituting compact bacteriomes and in individual bacteriocytes, which are dispersed among ovarioles. The bacteriocytes also house a few small, rod-shaped Wolbachia bacteria in addition to the numerous large, elongated Sodalis-allied bacteria. The symbiotic microorganisms are transovarially transmitted from generation to generation. In adult females which have choriogenic oocytes in the ovarioles, the bacteriocytes gather around the basal part of the tropharium. Next, the entire bacteriocytes pass through the follicular epithelium surrounding the neck region of the ovariole and enter the space between oocyte and follicular epithelium (perivitelline space). In the perivitelline space, the bacteriocytes assemble extracellularly in the deep depression of the oolemma at the anterior pole of the oocyte, forming a “symbiont ball”.

PUTOIDAE ambiguus Puto (Fullaway) 3f, 1i, d. bahiae Phenacoccus Ehrhorn 1f, 1m. Puto yuccae (Coquillett). barberi Puto (Cockerell) 1f. cupressi Puto (Coleman) 1f. sp. A Puto Green 2f, d. Cyprus [grandis]. superbus Ceroputo (Leonardi) 3f, d. Puto superbus (Leonardi). yuccae Puto (Coquillett) 1f, 1m, 1i, d., Published as part of Williams, D. J., 2017, E. E. Green's collection of scale insects (Hemiptera: Sternorrhyncha: Coccomorpha) in The Natural History Museum, London, U. K., pp. 201-253 in Zootaxa 4318 (2) on page 251, DOI: 10.11646/zootaxa.4318.2.1, http://zenodo.org/record/886712

Williams, D. J. (2017): E. E. Green's collection of scale insects (Hemiptera: Sternorrhyncha: Coccomorpha) in The Natural History Museum, London, U. K. Zootaxa 4318 (2): 201-253, DOI: https://doi.org/10.11646/zootaxa.4318.2.1

Fifty-one species in 27 genera of Pseudococcidae, plus one species of Rhizoecidae and one species of Putoidae have been recorded from Iranbased on literature data and on the material in the collection of the Hayk Mirzayans Insect Museum, Iran. Keys to the Iranian genera and species based on adult female morphology are provided and the distribution data of each species and their host-plants are given. Adult females of each species are illustrated in detail. The species Paracoccus burnerae (Brain), Phenacoccus pumilus (Kiritshenko) and Ph . hordei (Lindeman) are recorded for the first time in Iran. Four new species, Exallomochlus balouchestanensis Moghaddam sp. n. , Peliococcus ilamicus Moghaddam sp. n. , Phenacoccus karkasicus Moghaddam sp. n. and Phenacoccus iranica Moghaddam sp. n. are described.

This study aimed to determine the species of mealybugs (Pseudococcidae and Putoidae) found in five provinces (Ağrı, Bitlis, Hakkari, Iğdır and Van) in Eastern Anatolia, (Turkey). Species of the families Pseudococcidae and Putoidae were collected from natural and cultivated plants between 2005 and 2008. New locality and host plant data are given for all species. In total, 60 species of Pseudococcidae in 16 genera and two species of the genus Puto Signoret, 1875 were found, including 25 new species records for the Turkish fauna, namely: Atrococcus arakelianae (Ter-Grigorian, 1964,), A. cracens Williams, 1962, Coccura circumscripta (Kiritchenko, 1963), Dysmicoccus brevipes (Cockerell), Fonscolombia europaea (Newstead, 1897), Heliococcus glacialis (Newstead, 1900), Metadenopus festucaeŠulc, 1931, Mirococcopsis ammophila Bazarov & Nurmamatov, 1975, M. avetianae Ter-Grigorian, 1964, M. subalpina (Danzig, 1985), M. teberdae (Danzig, 1985), Phenacoccus angustatus Borchsenius, 1949, P. eurotiae Danzig, 1975, P. incertus (Kiritshenko, 1940), P. kokandicus Nurmamatov, 1986, P. persimplex Borchsenius, 1949, P. querculus (Borchsenius, 1949), P. prope schmelevi Bazarov, 1980, P. strigosus Borchsenius, 1949, Pseudococcus comstocki (Kuwana, 1902), Puto megriensis (Borchsenius, 1948), Ripersiella parva (Danzig, 1985), R. petiti (Goux, 1941), R. poltavae (Laing, 1929) and Spilococcus vashlovanicus Danzig, 1998. A new mealybug species, Phenacoccus chatakicus sp. nov., is described and illustrated based on adult females.

Mealybugs (Coccoidea: Pseudococcidae) are sap-sucking plant parasites that harbor bacterial endosymbionts within specialized organs. Previous studies have identified two subfamilies, Pseudococcinae and Phenacoccinae, within mealybugs and determined the primary endosymbionts (P-endosymbionts) of the Pseudococcinae to be Betaproteobacteria (“ Candidatus Tremblaya princeps”) containing Gammaproteobacteria secondary symbionts. Here, the P-endosymbionts of phenacoccine mealybugs are characterized based on 16S rRNA from the bacteria of 20 species of phenacoccine mealybugs and four outgroup Puto species (Coccoidea: Putoidae) and aligned to more than 100 published 16S rRNA sequences from symbiotic and free-living bacteria. Phylogenetic analyses recovered three separate lineages of bacteria from the Phenacoccinae, and these are considered to be the P-endosymbionts of their respective mealybug hosts, with those from (i) the mealybug genus Rastrococcus belonging to the Bacteroidetes , (ii) the subterranean mealybugs, tribe Rhizoecini, also within Bacteroidetes , in a clade sister to cockroach endosymbionts ( Blattabacterium ), and (iii) the remaining Phenacoccinae within the Betaproteobacteria , forming a well-supported sister group to “ Candidatus Tremblaya princeps.” Names are proposed for two strongly supported lineages: “ Candidatus Brownia rhizoecola” for P-endosymbionts of Rhizoecini and “ Candidatus Tremblaya phenacola” for P-endosymbionts of Phenacoccinae excluding Rastrococcus and Rhizoecini. Rates of nucleotide substitution among lineages of Tremblaya were inferred to be significantly faster than those of free-living Betaproteobacteria . Analyses also recovered a clade of Gammaproteobacteria , sister to the P-endosymbiont lineage of aphids (“ Candidatus Buchnera aphidicola”), containing the endosymbionts of Putoidae, the secondary endosymbionts of pseudococcine mealybugs, and the endosymbionts of several other insect groups.

The mealybugs (Hemiptera: Coccoidea: Pseudococcidae) are a speciose and ubiquitous group of sap-sucking plant parasites, many of which are very serious agricultural pests. There has been much work on the alpha-level taxonomy, amounting to the description of more than 2000 species, but suprageneric relationships remain poorly known. Downie & Gullan reviewed proposed schemes for a mealybug subfamily-level classification and used DNA sequence data from three nuclear genes to infer the mealybug phylogeny. They recognized three subfamilies: the Rhizoecinae, the Phenacoccinae, and the Pseudococcinae; and within the Pseudococcinae, recognized the tribes Pseudococcini, Trabutinini, and Planococcini. Excepting the Trabutinini, none of these groupings was well-supported. We improve our estimation of the mealybug phylogeny by: (i) increasing the taxon sampling by 50%; (ii) adding a morphological character matrix; and (iii) performing mixed model Bayesian and maximum likelihood inference procedures. We recover two primary clades within the Pseudococcidae, to which we apply the subfamily names Phenacoccinae and Pseudococcinae. Within the Phenacoccinae, we recover support for the monophyly of the hypogaeic mealybugs (Rhizoecini), and within the Pseudococcinae, we find support for the tribes Pseudococcini, Trabutinini, and Planococcini. Our results suggest a clear sequence of (i) β-Proteobacteria primary endosymbionts infecting the mycetome of the most recent common ancestor of the Pseudococcinae, followed by (ii) γ-Proteobacteria secondary endosymbionts infecting the primary β-Proteobacteria endosymbionts. For each subfamily, we provide a morphological diagnosis and a comprehensive list of included genera. We resurrect the genus Ceroputo Sulc, currently under synonymy with the genus Puto Signoret (Putoidae), and place it in the mealybug subfamily Phenacoccinae. Greenoripersia kaiseri Bodenheimer is transferred from the Pseudococcidae to the Eriococcidae.

Ben-Dov Y (1994) A systematic catalogue of the mealybugs of the world (Insecta: Homoptera: Coccoidea: Pseudococcidae and Putoidae) with data on geographical distribution, host plants, biology and economic importance. Intercept Limited, Andover, 686 p Cox JM (1989) The mealybug genus Planococcus (Homoptera: Pseudococcidae). Bull Br Museum (Nat Hist) Entomol 58(1):1–78 CIE (1975) Distribution maps of plant pests. Pseudococcus comstocki [Distribution map]. Map 338. CIE, London Williams DJ (2004) Mealybugs of southern Asia. The Natural History Museum/Southdene SDN. BHD, London/Kuala Lumpur, 896 p

This publication includes general discussions on the Conchaspididae, Diaspididae, Eriococcidae, Ortheziidae, Pseudococcidae, and Putoidae. Keys are presented for genera in the families Eriococcidae, Ortheziidae, and Pseudococcidae. Material for each family include introduction, field appearance, diagnosis, life history, important references, illustration of a slide-mounted adult female, and a checklist of the species occurring in the Southern Region of the United States and their distribution by state.

Mealybugs (Hemiptera: Pseudococcidae) are small, plant-sucking insects which comprise the second largest family of scale insects (Coccoidea). Relationships among many pseudococcid genera are poorly known and there is no stable higher level classification. Here we review previous hypotheses on relationships and classification and present the first comprehensive phylogenetic study of the Pseudococcidae based on analysis of nucleotide sequence data. We used three nuclear genes, comprising two noncontiguous fragments of elongation factor 1α (EF-1α 5′ and EF-1α 3′), fragments of the D2 and D10 expansion regions of the large subunit ribosomal DNA gene (28S), and a region of the small subunit ribosomal DNA gene (18S). We sampled sixty-four species of mealybug belonging to thirty-five genera and representing each of the five subfamilies which had been recognized previously, and included four species of Puto (Putoidae) and one species each of Aclerda (Aclerdidae) and Icerya (Margarodidae), using Icerya as the most distant outgroup. A combined analysis of all data found three major clades of mealybugs which we equate to the subfamilies Pseudococcinae, Phenacoccinae and Rhizoecinae. Within Pseudococcinae, we recognize the tribes Pseudococcini (for Pseudococcus, Dysmicoccus, Trionymus and a few smaller genera), Planococcini (for Planococcus and possibly Planococcoides) and Trabutinini (represented by a diverse range of genera, including Amonostherium, Antonina, Balanococcus, Nipaecoccus and non-African Paracoccus), as well as the Ferrisia group (for Ferrisia and Anisococcus), some ungrouped African taxa (Grewiacoccus, Paracoccus, Paraputo and Vryburgia), Chaetococcus bambusae and Maconellicoccus. The ‘legless’ mealybugs Antonina and Chaetococcus were not closely related and thus we confirmed that the Sphaerococcinae as presently constituted is polyphyletic. In our analyses, the subfamily Phenacoccinae was represented by just Phenacoccus and Heliococcus. The hypogeic mealybugs of the Rhizoecinae usually formed a monophyletic group sister to all other taxa. Our molecular data also suggest that the genera Pseudococcus, Dysmicoccus, Nipaecoccus and Paracoccus are not monophyletic (probably polyphyletic) and that Phenacoccus may be paraphyletic, but further sampling of species and genes is required. We compare our phylogenetic results with published information on the intracellular endosymbionts of mealybugs and hypothesize that the subfamily Pseudococcinae may be characterized by the possession of β-Proteobacteria (primary endosymbionts) capable of intracellular symbiosis with γ-Proteobacteria (secondary endosymbionts). Furthermore, our data suggest that the identities of the secondary endosymbionts may be useful in inferring mealybug relationships. Finally, cloning polymerase chain reaction products showed that paralogous copies of EF-1α were present in at least three taxa. Unlike the situation in Apis and Drosophila, the paralogues could not be distinguished by either the presence/absence or position of an intron.

Scale insects (Hemiptera: Sternorrhyncha: Coccoidea) are a speciose and morphologically specialized group of plant-feeding bugs in which evolutionary relationships and thus higher classification are controversial. Sequences derived from nuclear small-subunit ribosomal DNA were used to generate a preliminary molecular phylogeny for the Coccoidea based on 39 species representing 14 putative families. Monophyly of the archaeococcoids (comprising Ortheziidae, Margarodidae sensu lato, and Phenacoleachia) was equivocal, whereas monophyly of the neococcoids was supported. Putoidae, represented by Puto yuccae, was found to be outside the remainder of the neococcoid clade. These data are consistent with a single origin (in the ancestor of the neococcoid clade) of a chromosome system involving paternal genome elimination in males. Pseudococcidae (mealybugs) appear to be sister to the rest of the neococcoids and there are indications that Coccidae (soft scales) and Kerriidae (lac scales) are sister taxa. The Eriococcidae (felt scales) was not recovered as a monophyletic group and the eriococcid genus Eriococcus sensu lato was polyphyletic.

Past works on scale insects (Hemiptera: Coccoidea) from the Maltese Archipelago are reviewed. Based on the literature and contemporary collections, a total of 93 species of scale insects belonging to 12 scale insect families are here reported (Aclerdidae 1 species; Asterolecaniidae 4; Coccidae 17; Diaspididae 46; Eriococcidae 5; Kermesidae 1; Margarodidae 1; Micrococcidae 1; Monophlebidae 2; Pseudoccocidae 11; Putoidae 2 and Rhizoecidae 2). Of these, 17 species represent new distribution records. Ten species are excluded from the scale insect fauna of the Maltese Islands. Of the 93 species present, only 29 (31.18%) are probably indigenous and the rest (68.82%) represent established introductions from elsewhere. More than 65% of the indigenous species are typical Mediterranean in distribution, with a few species having a mainly European chorotype. A quarter of the established aliens originate from Eurasia, followed by an East Asian/ Oriental component (20.31%); European (14.06%); Neotropical (14.06%); cryptogenic (14.06%); African (7.81%) and Australasian (4.70%). Movement of live fruit trees and ornamental plants into the Maltese Archipelago from nearby countries is probably the main route for entry of alien scale insects into the country. Some possible future introductions are discussed.

In the past, the morphology of adult males of Coccoidea has provided strong support for diagnosing the higher taxonstatus of scale insects (Coccoidea). In particular, studies on adult male morphology have produced some of the stron-gest evidence for considering the Putoidae and Eriococcidae (as then defined) as separate families from the Pseudo-coccidae. This paper uses adult male morphology to assess the relationships of the Pseudococcidae and the hypogaeicand myrmecophilous mealybugs. The latter most often are classified as a subfamily (Rhizoecinae) of the Pseudococ-cidae. In order to diagnose the latter taxa, the adult males of fifteen named species of hypogaeic rhizoecine mealybugs(Kissrhizoecus hungaricus Kozar & Konczne Benedicty, Rhizoecus cacticans (Hambleton), Rh. coffeae Laing, Rh. dianthi Green, Rh. falcifer Kunckel d’Herculais, Rh. kazachstanus Matesova, Ripersiella cryphia (Williams), Ri. hibisci (Kawai & Takagi), Ri. kondonis (Kuwana), Ri. malschae (Williams), Ri. puhiensis (Hambleton), Capitisetella migrans (Green) and Pseudorhizoecus proximus Green) plus two unidentified Ripersiella species are described. Inaddition, the adult males of a Xenococcus sp., three Eumyrmococcus spp. and two Neochavesia spp. are illustratedfrom previously published papers and the adult male of another Neochavesia sp. is described and illustrated. In orderto compare the diagnoses of the above taxa with that of adult males of Pseudococcidae (minus the Rhizoecinae), theadult males of two apterous pseudococcid mealybugs are described or redescribed: Asaphococcus agninus Cox andthe myrmecophilous Promyrmococcus dilli Williams, both belonging to the Pseudococcinae. In addition, threemacropterous Pseudococcidae, namely Phenacoccus solenopsis Tinsley (Phenacoccinae), Planococcus glaucus (Maskell) and Maconellicoccus hirsutus (Green) (Pseudococcinae) are also described and/or illustrated. Prior to thisstudy, the hypogaeic and myrmecophilous mealybugs generally were included in the subfamily Rhizoecinae of thePseudococcidae, with the hypogaeic mealybugs in tribe Rhizoecini and the myrmecophilous mealybugs in Xenococ-cini. Based on the present study and on phylogenetic data, it is concluded that the rhizoecine mealybugs form a sepa-rate family from the Pseudococcidae — Rhizoecidae Williams. This family is considered here to include twosubfamilies, Rhizoecinae Williams and Xenococcinae Tang. Based on adult male characters, there is little support forthe present generic divisions of the Rhizoecinae. Keys are given for separating the adult males of Rhizoecidae from those of Pseudococcidae, and for separating the known adult males within each subfamily.

For almost a century, the scale insect genus Puto Signoret (Hemiptera: Sternorrhyncha: Coccoidea) was considered to belong to the family Pseudococcidae (the mealybugs), but recent consensus accords Puto its own family, the Putoidae. This paper reviews the taxonomic history of Puto and family Putoidae, compares the morphology of Puto to that of Ceroputo Šulc and Phenacoccus Cockerell, and reassesses the status of all species that have been placed in Puto to determine whether they belong to the Putoidae or to the Pseudococcidae. For 49 of 57 species that have been placed in Puto, as listed in the online database ScaleNet, we score and tabulate features that are diagnostic for Putoidae and then list all species in their correct family placement. For comparison, we include a few species of Pseudococcidae, namely five species of Phenacoccus, including the type species Phenacoccus aceris (Signoret), and the type species of Ceroputo, C. pilosellae Šulc. We provide revised synonymy lists for Puto and Ceroputo, a brief diagnosis of each genus, synonymy lists and notes for several species for which we suggest recombinations or additional synonyms, or for which we have additional data on morphology. We provide a brief diagnosis of Phenacoccus for comparison with Ceroputo and Puto. As a result of our study, we recognise 47 extant and two fossil species of Puto, and six species of Ceroputo. The New World species Puto mimicus McKenzie and Puto nulliporus McKenzie are transferred to the mealybug genus Ceroputo as Ceroputo mimicus (McKenzie) comb. nov. and Ceroputo nulliporus (McKenzie) comb. nov., respectively, and the Old World species Puto pini Danzig and Puto vaccinii Danzig are recognised as Ceroputo pini (Danzig) comb. rev. and Ceroputo vaccinii (Danzig) comb. rev., respectively, in agreement with Tang (1992). The Old World species Puto graminis Danzig is transferred to Ceroputo as Ceroputo graminis (Danzig) comb. nov. Based only on a study of the literature, the following two names are treated here as junior subjective synonyms of Ceroputo pilosellae: Phenacoccus asteri Takahashi syn. nov. and Puto jarudensis Tang syn. nov. We agree with Tang (1992) that Leococcus erigeroneus Kanda should be treated as a junior subjective synonym of C. pilosellae and thus the genus name Leococcus Kanda, erected for L. erigeroneus and formerly treated as a junior synonym of Puto, is a junior synonym of Ceroputo.