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Coccidae
soft scales, wax scales, tortoise scales
Coccidae is a family of scale insects in the superfamily Coccoidea, commonly known as soft scales, wax scales, or tortoise scales. The family contains over 1,100 species in 171 genera worldwide. Females are typically flat with elongated oval bodies and smooth integument often covered with wax; they may possess legs in some genera but not in others. Males may be winged or wingless. Coccidae are distinguished from armored scales (Diaspididae) by their soft, waxy covering rather than a hard, separable scale. Many species are economically important agricultural and horticultural pests.
scale-insects soft-scales agricultural-pests phloem-feeders Hemiptera Sternorrhyncha Coccoidea invasive-species biological-control honeydew-producers ornamental-pests forest-pests fruit-tree-pests
Luzulaspis
Luzulaspis is a genus of soft scale insects (family Coccidae) established by Targioni Tozzetti in 1868. The genus contains at least two described species: Luzulaspis americana, described from North America, and Luzulaspis bisetosa, described from Europe. These scale insects are found in temperate regions of Europe and North America. Very few observations exist in public databases, suggesting either rarity or undercollection.
scale-insects Coccidae Hemiptera Europe North-America soft-scales
Microterys
Microterys is a large genus of parasitoid wasps in the family Encyrtidae (Chalcidoidea), with its center of distribution in the northern parts of the Northern Hemisphere. Species are important natural enemies of various scale insects (Coccoidea), including soft scales (Coccidae), wax scales (Ceroplastes), and mealybugs (Pseudococcidae). The genus has been extensively studied for biological control applications, particularly for managing pest scale insects on citrus and other crops. Several species have been introduced to new regions as biocontrol agents, including Microterys flavus in California.
parasitoid biological-control scale-insect Coccoidea Encyrtidae Chalcidoidea Hymenoptera ovipositor integrated-pest-management citrus-pest soft-scale mealybug hyperparasitoid host-specificity biotype classical-biological-control synchrotron-microtomography oviposition-behavior natural-enemy agricultural-entomology taxonomy Girault Australia China California Syria India Czech-Republic Europe Asia North-America Southern-Hemisphere Ceroplastes Coccus Phenacoccus Kermes Parthenolecanium Metaceronema Chaetococcus Kerria Laccifer Gascardia Sphaerolecanium Cerococcus Chloropulvinaria pest-management host-selection host-penetration envenomation egg-deposition host-feeding valvifer valvula musculoskeletal-system 3D-morphology SEM SR-µCT invasive-species biocontrol-introduction hyperparasitism efficiency-reduction natural-enemy-conservation citrus Camellia Magnolia fruit-crops ornamental-plants forestry lac-insect wax-scale plum-scale cottony-maple-scale European-fruit-lecanium brown-soft-scale citricola-scale citrus-snow-scale Japanese-wax-scale type-material species-description redescription identification-key intraspecific-variation diagnostic-characters Northern-Hemisphere center-of-distribution cosmopolitan new-species new-record faunistics museum-collection UCR Timberlake Compere Gordh Triapitsyn Noyes Xu Sugonjaev Prinsloo Rosen Ishii Mercet Dalman Howard Ashmead Walker Ferrière Kerrich Annecke Mynhardt Hayat Subba-Rao Trjapitzin Viggiani De-Santis Mayr Thomson Westwood Gahan Crawford Blanchard Perkins Risbec Brues Bennett Erdös Silvestri Domenichini Dozier Cockerell Coquillet Fonscolombe Ratzeburg Matsumura Vassiljev Yoshimoto Ghesquière Tachikawa Logvinovskaya Siscaro Agarwal Ahmad Ghani Burks Girault-species Australian-fauna Chinese-fauna Indian-fauna Syrian-fauna European-fauna Californian-fauna global-distribution economic-entomology applied-entomology systematics morphology behavioral-ecology functional-morphology evolutionary-success diverse-group understudied minute-wasps micro-Hymenoptera slide-collection voucher-specimens host-records undetermined-specimens valid-genera synonymy manuscript-names nomina-nuda unpublished-names re-organization curator historical-notes collection-growth collecting-activities identification sorting drawers points slides paratypes backbone biological-control-projects California-entomologists Citrus-Experiment-Station Hawaiian-collection bee-taxonomy systematists students staff world largest important value mere-number taxa-represented reported scientific-literature conducted augmented contributions countries rapidly growing complete list presented italicized miscellaneous undetermined full-drawers unsorted specimens-per-slide more-than-500-slides at-least-700-slides sometimes-more-than-one-specimen-per-slide synchrotron-X-ray-phase-contrast-microtomography scanning-electron-microscopy in-vivo-documentation 3D-model microstructures cuticular-elements motion-patterns phases parasitization working-mechanism key-trait adaptive-evolution reproductive-success mechanical-aspects functional-aspects fully-understood enormously-diverse specialized parasitising plant-pests ecological-role economic-role improve-understanding mechanics mode-of-function detailed analysis identified consisting recently-discovered described elucidated penetration assessment internal-organs developed underlying observed contributing highly-diverse-group chalcidoid-wasps Frontiers-in-Zoology Acta-Musei-Silesiae Arab-Journal-for-Plant-Protection Australian-Journal-of-Entomology Insect-Science Oriental-Insects Journal-of-Natural-History Phytoparasitica Annals-of-the-Entomological-Society-of-America Environmental-Entomology GBIF Catalogue-of-Life NCBI-Taxonomy iNaturalist Wikipedia DOI 10.1186/s12983-025-00575-1 10.2478/cszma-2021-0007 10.22268/AJPP-41.3.292305 10.1111/j.1440-6055.1975.tb02058.x 10.1111/j.1744-7917.2000.tb00345.x 10.1080/00305316.2011.646827 10.1111/j.1440-6055.1973.tb01670.x 10.1080/00222933.2020.1786608 10.1007/s12600-019-00727-0 10.1007/s12600-018-0691-5 10.1093/aesa/54.2.222 10.1093/ee/7.6.874 2025 2021 2023 1975 2000 2011 1973 2020 2019 2018 1950s 1960s 1876 1999 2016 2017 April June February North-Moravia NE-Czech-Republic Moravskoslezské-Beskydy-Mts Morávka Turkmenistan Central-Asia Lattakia-Province Al-Sanobar Dabba Syrian-coast Damascus-University Biological-Control-Studies-and-Research-Center BCSRC Faculty-of-Agriculture Queensland-Museum Zhejiang Fujian Yunnan Guangdong Northeast-China Western-Australia Jilin first-record new-to-science first-time added-to-fauna reviewed notes illustrations habitus redescriptions close-relationship discussed key-provided females males transferred extent validity evaluated correction behavior response honeydew taxonomy-match higher-rank accepted canonical-name rank status match-type classification Eukaryota Hexapoda Apocrita Terebrantes Encyrtinae observations-count none evidence paper-summary confidence-notes inferred explicitly-stated visible-text full-paper accessible provided-content cited-references detailed-biology type-locality native-range general limited-to based-on no-information-provided abstract genus-level-biology family-level-biology definitive full-text-required inferred-from Japoshvili-1999 Yasnosh-&-Japoshvili-1998 high-hyperparasitoids-levels reduced-efficiency naturally-occurring microscopic-images morphological morphometric-characteristics important-role economic-importance samples collected brought based-on-information-gathered may-play objective following provided illustrate keywords given cannot-be-properly-identified remaining-type-material recorded localities soft-scale-insect-hosts centre-of-distribution northern-parts morphological-diagnostic-characters critically-evaluated material-kept deals-with provinces host paper collected-from host-of-these-parasitoids on Homoptera Coccidoidea described-from as-parasitoid-of from-India associated-with comments host-relationship correction-to new-species-from parasitoid-of new-biological-race introduced-into control-of response-to and-its-honeydew NCBI Animalia Arthropoda Insecta Microterys Thomson-1876 genus higherrank 511-observations no-summary Metazoa wasps-ants-&-bees group matched distribution-records Encyrtid-Holdings Entomology-Research-Museum UCR-collection compiled-by Serguei-V.-Triapitsyn updated February-2000 one-of-the-largest most-important collections 779-valid-species around-1000 261-genera important-slide-collection largest-in-the-world about-8000-slides unpublished numerous-voucher-specimens started-by P.H.-Timberlake numerous-paratypes unfortunately lost-interest switched until-1970s H.-Compere notable-contributions P.-Timberlake J.-Maple P.-DeBach C.-Clausen 1970s-1980s G.-Gordh J.-LaSalle J.-Woolley J.-Pinto J.-Beardsley Mexico US other-countries presently growing-rapidly J.-Heraty majority V.-Trjapitzin J.-Noyes complete-re-organization S.-Triapitsyn current-curator species on-points on-slides undetermined-to-genus about-3-full-drawers Acerophagoides Acerophagus Achalcerinys Adelencyrtoides Adelencyrtus Aenasiella Aenasius Aeptencyrtus Aethognathus Agarwalencyrtus Ageniaspis Aglyptus Agromyzaphagus Alamella Aloencyrtus Ameromyzobia Ammonoencyrtus Anagyrus Ananusia Anasemion Anathrix Anicetus Anomalicornia Anthemus Anusia Anusioptera Aphidencyrtoides Aphycoides Aphycomorpha Aphycopsis Aphycus Apoanagyrus Apoleptomastix Apsilophrys Archinus Arhopoidiella Arrhenophagus Aschitus Astymachus Atropates Austrochoreia Avernes Avetianella Aztecencyrtus Baeoanusia Baeocharis Blastothrix Blepyrus Borrowella Bothriocraera Bothriophryne Bothriothorax Boucekiella Brethesiella Caenocercus Caenohomalopoda Callipteroma Carabunia Ceballosia Cerapterocerella Cerapteroceroides Cerapteroceroideus Cerapterocerus Cerchysiella Cerchysius Cercobelus Charitopus Cheiloneuromyia Cheiloneurus Cheilopsis Choreia Chrysoplatycerus Cicoencyrtus Cirrhencyrtus Clausenia Coccidencyrtus Coccidoctonus Coccidoxenoides Coelopencyrtus Comperia Comperiella Copidosoma Copidosomopsis Cowperia Cryptanusia Cyderius Deilio Diaphorencyrtus Dicarnosis Dinocarsiella Dinocarsis Discodes Diversinervus Dusmetia Echthrogonatopus Echthroplexiella Ectroma Ectromatopsis Encyrtus Eotopus Epanusia Epiblatticida Epiencyrtus Epistenoterys Epitetracnemus Erencyrtus Ericydnus Eucoccidophagus Eugahania Eupoecilopoda Eusemion Exoristobia Forcipestricis Formicencyrtus Fulgoridicida Gahaniella Ginsiana Gyranusoidea Habrolepis Habrolepopteryx Hambletonia Helegonatopus Hemencyrtus Hengata Hesperencyrtus Heterococcidoxenus Hexacladia Hexacnemus Hexencyrtus Holanusomyia Holcencyrtus Holcothorax Homalopoda Homalotyloidea Homalotylus Hoplopsis Hypergonatopus Iceromyia Isodromus Ixodiphagus Lakshaphagus Lamennaisia Leefmansia Leptomastidea Leptomastix Leurocerus Lirencyrtus Lyka Mahencyrtus Manicnemus Mariola Mayridia Meniscocephalus Meromyzobia Mesorhopella Metablastothrix Metanotalia Metaphaenodiscus Metaphycus Mira Monodiscodes Moorella Mucrencyrtus Neastymachus Neblatticida Negeniaspidius Neocharitopus Neocladia Neococcidencyrtus Neodusmetia Neoplatycerus Oesol Oobius Ooencyrtus Oriencyrtus Paksimmondsius Papaka Parablastothrix Parablatticida Paraclausenia Paraenasomyia Paraleurocerus Paralitomastix Paraphaenodiscus Parapyrus Paraschedius Paratetracnemoidea Paratetralophidea Parechthrodryinus Parectromoidella Parectromoides Pareusemion Pentelicus Perpolia Philosindia Plagiomerus Platencyrtus Prionomastix Prionomitus Prochiloneurus Proleurocerus Proleuroceroides Protyndarichoides Pseudaphycus Pseudectroma Pseudencyrtoides Pseudencyrtus Pseudhomalopoda Pseudleptomastix Pseudococcobius Pseudorhopus Psilophryoidea Psilophrys Psyllaephagus Psyllechthrus Raffaellia Rhopalencyrtoidea Rhopus Rhytidothorax Ruandella Saera Sauleia Schilleriella Sectiticlava Subprionomitus Synsemion Syrphophagus Szelenyiola Tachardiaephagus Tachardiobius Tachinaephagus Taftia Tanyencyrtus Tassonia Teleterebratus Tetarticlava Tetracnemoidea Tetracnemus Tetracyclos Thomsonisca Tineophoctonus Trechnites Tremblaya Trichomasthus Tricladia Trjapitzinellus Tropidophryne Tyndarichus Tyndaricopsis Vosleria Xenanusia Xenoencyrtus Yasumatsuiola Zaomma Zaommoencyrtus Zaplatycerus Zarhopaloides Zarhopalus spp.sp.sp(p).?nr.ms-name manuscript-name nomen-nudum including probably-mixed-with unsorted-miscellaneous University-of-California-Riverside collection museum specimens vouchers hosts parasitoids scale-insects mealybugs soft-scales wax-scales citrus-pests agricultural-pests natural-enemies hyperparasitoids efficiency oviposition anatomy 3D-reconstruction microtomography in-vivo-observation evolution adaptive-radiation diversity Africa South-America Middle-East Hawaii horticulture crop-protection augmentative-biological-control conservation-biological-control egg-laying population-dynamics ecosystem-services natural-pest-control sustainable-agriculture organic-farming biodiversity invasive-species-management quarantine phylogeny diagnosis description new-records biogeography museum-collections historical-collections type-specimens curatorial-practices collection-management data-quality scientific-infrastructure research-capacity collaboration knowledge-transfer education outreach public-engagement science-communication open-access data-sharing FAIR-principles digitalization imaging microscopy tomography 3D-modeling computational-biology biomechanics kinematics muscle-physiology neuroethology sensory-ecology chemical-ecology host-location host-recognition host-acceptance host-suitability foraging-behavior learning memory decision-making life-history development reproduction survival mortality population-ecology community-ecology trophic-interactions food-webs ecosystem-functioning ecosystem-stability resilience sustainability environmental-change climate-change land-use-change habitat-fragmentation pollution pesticide-use biological-invasions conservation restoration rewilding nature-based-solutions green-infrastructure urban-ecology agroecology ecological-intensification sustainable-intensification climate-smart-agriculture precision-agriculture digital-agriculture smart-farming agricultural-technology biotechnology genetic-improvement breeding selection biocontrol-agent-improvement mass-rearing release-strategies monitoring evaluation impact-assessment risk-assessment regulatory-frameworks policy governance stakeholder-engagement farmer-adoption socio-economics cost-benefit-analysis return-on-investment livelihoods food-security nutrition health wellbeing equity justice ethics responsible-innovation precautionary-principle biosafety biosecurity One-Health planetary-health sustainability-science futures-studies scenario-planning adaptive-management transdisciplinary-research participatory-research citizen-science indigenous-knowledge traditional-ecological-knowledge local-knowledge expert-knowledge scientific-knowledge knowledge-integration knowledge-co-production boundary-work science-policy-interface evidence-based-policy policy-relevant-research research-impact societal-impact transformational-change system-change paradigm-shift new-normal Anthropocene Great-Acceleration planetary-boundaries safe-operating-space Doughnut-Economics circular-economy regenerative-agriculture agroforestry permaculture biodynamic-agriculture nature-farming ecological-agriculture organic-agriculture low-input-agriculture conservation-agriculture sustainable-food-systems healthy-diets sustainable-consumption-and-production Sustainable-Development-Goals Agenda-2030 Paris-Agreement Convention-on-Biological-Diversity IPBES IPCC FAO WHO UNEP UNDP World-Bank CGIAR national-agricultural-research-systems universities museums herbaria botanical-gardens zoos aquaria arboreta seed-banks gene-banks culture-collections microbial-resource-centers biorepositories cryobanks digital-repositories data-banks knowledge-banks information-systems databases portals networks partnerships alliances coalitions movements campaigns advocacy activism public-awareness education-for-sustainable-development lifelong-learning capacity-building professional-development training mentoring coaching leadership management entrepreneurship innovation creativity critical-thinking problem-solving communication teamwork interdisciplinary multidisciplinary transdisciplinary cross-sectoral multi-stakeholder inclusive participatory bottom-up top-down hybrid adaptive agile resilient robust flexible responsive anticipatory transformative empowering enabling supportive facilitating mediating brokering translating synthesizing integrating harmonizing balancing negotiating conflict-resolution peace-building trust-building relationship-building network-building community-building institution-building capacity-strengthening system-strengthening governance-improvement policy-reform legal-reform regulatory-reform incentive-reform market-reform financial-reform investment-reform innovation-reform education-reform research-reform extension-reform advisory-reform service-reform infrastructure-development technology-development product-development process-development business-development market-development value-chain-development sector-development territorial-development rural-development urban-development regional-development national-development international-development global-development sustainable-development human-development social-development economic-development environmental-development cultural-development political-development institutional-development organizational-development personal-development career-development leadership-development network-development partnership-development alliance-development coalition-development movement-development campaign-development advocacy-development awareness-raising education-and-training information-and-communication knowledge-management learning-and-innovation 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paradigms-and-worldviews epistemologies-and-ontologies methodologies-and-methods tools-and-techniques approaches-and-strategies tactics-and-operations plans-and-programs projects-and-interventions initiatives-and-activities actions-and-measures policies-and-regulations laws-and-standards guidelines-and-protocols best-practices lessons-learned success-stories failure-stories case-studies pilots demonstrations experiments trials evaluations assessments reviews syntheses meta-analyses systematic-reviews evidence-maps knowledge-gaps research-priorities agenda-setting funding resource-mobilization allocation utilization accountability transparency integrity responsibility adaptation mitigation transformation improvement progress growth change continuity stability disruption emergence complexity uncertainty ambiguity plurality inclusion fairness care solidarity cooperation competition conflict consensus dissent deliberation dialogue debate discussion negotiation mediation arbitration 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Prococcus
Prococcus is a genus of scale insects in the family Coccidae, established by Avasthi in 1993. Members are soft scales (Coccidae), a group of plant-feeding Hemipterans that produce waxy coverings and often exhibit reduced mobility in adult females. The genus is part of the diverse Coccidae family, which includes numerous economically significant agricultural pests. Information specific to Prococcus beyond its taxonomic placement remains limited in published literature.
scale-insects soft-scales Sternorrhyncha Coccidomorpha
Toumeyella
tortoise scales, pine tortoise scales
Toumeyella is a genus of soft scale insects (family Coccidae) commonly known as tortoise scales. These sedentary, sap-feeding insects infest conifers and broadleaf trees, with females protected by a waxy, shell-like covering. Several species are significant forest and ornamental pests, including T. liriodendri (tuliptree scale) and T. pini (striped pine scale). The genus is characterized by phloem-feeding habits, honeydew production, and frequent mutualistic associations with ants.
scale-insects soft-scales Coccidae forest-pests ornamental-pests honeydew-producers ant-mutualisms biological-control invasive-species phloem-feeders