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Phrynocaria funebris (Crotch, 1874) (Fig. 4) Coelophora funebris Crotch, 1874: 154.— Korschefsky 1932: 292. Phrynocaria funebris: Poorani 2002: 338. Diagnosis: Phrynocaria funebris (Figs. 4a, b) is superficially similar to the fully melanic form of P. perrotetti in having black elytra, but the pronotum is yellowish-ochreous with a pair of black maculae on basal margin and a pair of much smaller, obliquely transverse discal markings and the lateral margins of elytra are narrowly ochreous and the male genitalia are diagnostic. It also resembles Synona spp. in having fully black elytra, but the latter can be distinguished by the semicircularly emarginate clypeal margin and the more convex and broader body outline with a distinct marginal bead. The male genitalia (Figs. 4 c–e) are, however, diagnostic. Brief redescription: Form (Figs. 4a, b) broadly rounded, dorsum strongly convex and glabrous. Head and pronotum ochreous, pronotum with a pair of characteristic black maculae on basal margin, not connected, together forming a semicircular median trough, above these a pair of much smaller, obliquely transverse markings; scutellar shield black; elytra dark brown to black, lateral margins narrowly yellowish-ochreous for up to ¾ of elytral length, apically darker. Ventral side yellowish. Head with eyes posteriorly strongly divergent, interocular distance subequal to eye width. Punctures on head irregular, closer and denser next to ocular margins than in middle, interspaces between punctures reticulately sculptured. Pronotum finely punctate, punctures shallowly impressed, separated by 2–4 diameters, interspaces with weak, reticulate sculpture. Elytral disc finely punctate, punctures slightly finer than those on pronotum, separated by 4–6 diameters, interspaces apparently smooth; lateral borders with many coarser punctures mixed with smaller ones. Prothoracic hypomera with prominent foveae on anterolateral corners. Prosternal carinae present. Legs without tibial spurs, tarsal claws appendiculate. Abdominal postcoxal line incomplete. Male genitalia (Figs. 4 c–e) as illustrated, penis guide (Figs. 4c, d) shorter than parameres, broadest at 2/4 th length, narrower at 3/4 th length, apical 1/4 th still narrower, apex broadly arcuate with a short blunt projection, outer and apical margins of parameres densely setose. Penis capsule (Fig. 4e) well developed, outer process wider and longer than inner one, robust in the middle, penis apex as illustrated. Material examined: “ Type (red bordered circular label) / Bengalore / Bowring 63.47* / Coelophora funebris, Crotch, type / NHMUK014016390 ” (BMNH). Distribution: India (Karnataka). This species was originally described from Bangalore and appears to be rare as it is known only from the type specimen. We have not seen any corresponding examples in any of the major Indian collections., Published as part of Poorani, J., Sankararaman, H. & Anusree, S. S., 2021, A review of Indian species of Phrynocaria Timberlake (Coleoptera, Coccinellidae) with notes on synonymy and a new species from South India, hitherto misidentified as Coelophora circumusta (Mulsant), pp. 117-133 in Zootaxa 4926 (1) on page 124, DOI: 10.11646/zootaxa.4926.1.8, http://zenodo.org/record/4500738, {"references":["Crotch, G. R. (1874) A revision of the coleopterous family Coccinellidae. E. W. Janson, London, 311 pp. https: // doi. org / 10.5962 / bhl. title. 8975","Korschefsky, R. (1932) Coleopterorum Catalogus. Pars 120. Coccinellidae II. W. Junk, Berlin, 435 pp.","Poorani, J. (2002) An annotated checklist of the Coccinellidae (Coleoptera) (excluding Epilachninae) of the Indian Subregion. Oriental Insects, 36, 307 - 383. https: // doi. org / 10.1080 / 00305316.2002.10417335"]}

Phrynocaria circumusta (Mulsant, 1850), comb. nov. (Fig. 1) Artemis circumusta Mulsant, 1850: 374, 388; 1866: 256.—Preoccupied in Lepidoptera (not Artemis Kirby et Spence, 1828 (Insecta: Lepidoptera: Saturniidae) (Ukrainsky 2006: 400)). Coelophora circumusta: Crotch 1874: 150.— Korschefsky 1932: 291.— Poorani 2002: 328.— Kovář 2007: 613. Lemnia (Artemis) circumusta: Iablokoff-Khnzorian 1982: 233. Lemnia (Neoartemis) circumusta: Ukrainsky 2006: 400. Lemnia circumusta: Yu 2010: 76–77. Coelophora moseri Weise, 1902: 500. New synonym. Lemnia moseri: Iablokoff-Khnzorian 1982: 259. Phrynocaria moseri: Ślipiński et al. 2020: 105. Coelophora westermanni Mulsant, 1850: 391; 1866: 257.— Crotch 1874: 151.—Synonymized by Iablokoff-Khnzorian 1982: 233. Artemis mandarina Mulsant, 1850: 389.— Crotch 1874: 150 (as synonym of C. circumusta). Artemis rufula Mulsant, 1850: 389.— Crotch 1874: 150 (as synonym of C. circumusta). Diagnosis: Phrynocaria circumusta can be identified by its reddish / yellowish head and pronotum and black elytra, each elytron bearing a red macula on the basal margin adjacent to the scutellar shield (Fig. 1a) that is often much enlarged and comma shaped (Fig. 1b). Variants with reddish elytra bearing narrow black external margins and with immaculate reddish elytra are known from the type locality (Hong Kong) and also from northern India. Rarely, forms with yellowish pronotum and fully black elytra are found that are externally similar to Synona spp. The male genitalia (Figs. 1i, j) are, however, diagnostic. Material examined: INDIA: “Madras, India / G. Bryant coll. 1919-147/ Artemis circumusta / male genitalia in glass vial” (1, BMNH); “ Assam: Monkhooshi T.E. Assam / Assoc. with tea / Coelophora circumusta (Muls.), det. R.G. Booth” (1, BMNH); “Chabua, Assam ” (1, BMNH). Distribution: India: Andhra Pradesh; Assam; Kerala; Jammu & Kashmir; Uttarakhand. Nepal. Thailand. China Hong Kong (Lee & Winney 1981). Taiwan. . It was ‘purposely introduced, not established’ in Hawaii (Leeper 2015). Timberlake (1943) and Nishida (2002) also mentioned its introduction in Hawaii and the intended target pest was ‘Psylla’. Notes: Mulsant (1850) originally described the genus Artemis with three species: A. circumusta (with reddish elytra bearing narrow black external margins), A. rufula (with immaculate reddish elytra) and A. mandarina (with black elytra, each bearing a red macula on the anterior margin adjacent to the scutellar shield (Fig. 1a) that is often much enlarged and comma shaped (Fig. 1b)). All three newly described species were from Hong Kong (China). Mulsant opined that they were much alike, differing only in the elytral colour pattern. Crotch (1874) synonymized the three species noting that ‘these three forms differed, as Mulsant supposed, only in colour’. Crotch (1874) recognized the combination Coelophora circumusta and treated Lemnia and Artemis as congeners of Coelophora stating that they ‘hardly differed in any points of importance’. The generic placement of this species has been contentious. Iablokoff-Khnzorian (1982) treated Coelophora and Lemnia as distinct genera and Artemis as a subgenus of Lemnia, using the name Lemnia (Artemis) circumusta. Ukrainsky (2006) gave a replacement name, Neoartemis, for this subgenus because Artemis was preoccupied in Lepidoptera (Saturniidae). Ślipiński (2007) treated Lemnia as a synonym of Coelophora, but Ren et al. (2009) and Yu (2010) considered Lemnia and Coelophora as distinct genera, and used the name ‘ L. circumusta ’. Poorani (2002) included C. circumusta in the checklist of Indian Coccinellidae with a note that ‘it may not be a true Coelophora as it lacks tibial spurs.’. It is transferred here to Phrynocaria in view of the typical generic characters (new combination). The male genitalia of P. moseri (Weise, 1902), illustrated in the revision of Australo-Pacific Coccinellini by Ślipiński et al. (2020), look extremely close to those of P. circumusta and the two species are treated here as synonyms (new synonym)., Published as part of Poorani, J., Sankararaman, H. & Anusree, S. S., 2021, A review of Indian species of Phrynocaria Timberlake (Coleoptera, Coccinellidae) with notes on synonymy and a new species from South India, hitherto misidentified as Coelophora circumusta (Mulsant), pp. 117-133 in Zootaxa 4926 (1) on page 119, DOI: 10.11646/zootaxa.4926.1.8, http://zenodo.org/record/4500738, {"references":["Mulsant, E. (1850) Species des Coleopteres Trimeres Securipalpes. Annales des Sciences Physiques et Naturelles, d'Agriculture et d'Industrie, publiees par la Societe nationale d'Agriculture, etc., de Lyon, Deuxieme Serie, 2, 1 - 1104. https: // doi. org / 10.5962 / bhl. title. 8953","Kirby, W. & Spence, W. (1828) An Introduction to Entomology: or, Elements of the Natural History of Insects. Vol. III. 5 th Edition. Longman, Rees, Orme, Brown & Green, London, 732 pp. https: // doi. org / 10.5962 / bhl. title. 152792","Ukrainsky, A. S. (2006) Five new replacement ladybird (Coleoptera: Coccinellidae) generic names. Russian Entomological Journal, 15 (4), 399 - 400.","Crotch, G. R. (1874) A revision of the coleopterous family Coccinellidae. E. W. Janson, London, 311 pp. https: // doi. org / 10.5962 / bhl. title. 8975","Korschefsky, R. (1932) Coleopterorum Catalogus. Pars 120. Coccinellidae II. W. Junk, Berlin, 435 pp.","Poorani, J. (2002) An annotated checklist of the Coccinellidae (Coleoptera) (excluding Epilachninae) of the Indian Subregion. Oriental Insects, 36, 307 - 383. https: // doi. org / 10.1080 / 00305316.2002.10417335","Iablokoff-Khnzorian, S. M. (1982) Les Coccinelles Coleopteres-Coccinellidae. Tribu Coccinellini des regions Palearctique et Orientale. Societe nouvelle des editions Boubee, Paris, 568 pp.","Yu, G. (2010) Chinese Lady Beetles (The Subfamily Coccinellinae). China Scientific Press, Beijing, 180 pp. [in Chinese]","Weise, J. (1902) Coccinelliden aus der Sammlung des Ungarischen National-Museums. Termeszetrajzi Fuzetek, 25, 489 - 520.","Slipinski, A., Li, J. & Pang, H. (2020) Ladybird beetles of the Australo-Pacific region. Coleoptera: Coccinellidae: Coccinellini. CSIRO Publishing, Clayton South, 231 pp. https: // doi. org / 10.1071 / 9781486303885","Mulsant, E. (1866) Monographie des Coccinellides. 1 re Partie Coccinelliens. Savy et Deyrolle, Paris, 294 pp.","Lee, L. H. Y. & Winney, R. (1981) Check List of agricultural insects of Hong Kong 1981. Agriculture and Fisheries Department Bulletin, Hong Kong, 2, 162.","Leeper, J. R. (2015) An annotated checklist of the coccinellid (Coleoptera Coccinellidae) introductions and establishments in Ha- waii: 1885 to 2015. Available from: https: // evols. library. manoa. hawaii. edu / handle / 10524 / 48742 (accessed 5 May 2020)","Timberlake, P. H. (1943) The Coccinellidae or Ladybeetles of the Koebele Collection-Part 1. Hawaiian Planters' Record, 47 (1), 1 - 67.","Nishida, G. M. (2002) Hawaiian terrestrial arthropod checklist. Fourth Edition. Bishop Museum Technical Report, 22, 1 - 313.","Slipinski, A. (2007) Australian ladybird beetles (Coleoptera: Coccinellidae), their biology and classification. Australian Biological Resources Study, Canberra, 286 pp.","Ren, S. X., Wang, X. M., Pang, H., Peng, Z. Q. & Zeng, T. (2009) Colored pictorial handbook of ladybird beetles in China. Science Press, Beijing, 336 pp. [in Chinese]"]}

Phrynocaria perrotteti (Mulsant, 1850) (Figs. 5, 6) Coelophora perrotteti Mulsant, 1850: 409. Coelophora perrotteti: Crotch 1874: 54.— Korschefsky 1932: 296.— Gordon 1987: 19. Anegleis (Pseudanegleis) perrotteti: Iablokoff-Khnzorian 1982: 296. Phrynocaria perrotteti: Poorani 2002: 339. Diagnosis: The nominate form of Phrynocaria perrotteti (Figs. 5 a–c) is superficially strikingly similar (both as adults and larvae) to Anegleis cardoni (Weise) and both have surprisingly similar host affinities too and are frequently associated with whiteflies. Both species coexist in South India and the propensity for feeding on whitefly prey is much more pronounced in the case of P. perrotteti (Figs. 5j, k). Phrynocaria perrotetti is highly variable (Fig. 5 a–i) and can be identified only by a combination of the generic characters. Anegleis cardoni can be differentiated from the nominate form of P. perrotteti by the elytral pattern, widely separated and smaller eyes, much smaller scutellar shield, prothoracic hypomeron without foveae, and the male genitalia. Brief redescription: Length 4.00– 4.70 mm, width 3.60–4.20 mm. Form (Figs. 5 a–i) variable from round to slightly broadly oval, strongly convex. Nominate form (Figs. 5 a–c) pale creamy yellow to lemon yellow, with black markings on pronotum and elytra as follows: pronotum with a pair of triangular markings on posterior margin, a circular median spot and two smaller lateral spots; scutellar shield broadly triangular; elytra with three stripes, one along suture, two on each elytron–inner stripe short and straight, outer stripe longer, curved, posteriorly clubbed. Highly variable with the following common forms: completely black except anterior and lateral margins of pronotum narrowly pale yellow (Fig. 5i) / completely orange to yellowish brown (Figs. 5g, h) / orange yellow with four small black spots (Fig. 5f) / orange yellow with a large black patch occupying most of the discal area except proximal one-fourth (Figs. 5d, e). Head (Fig. 6a) with large, posteriorly divergent eyes, antenna (Fig. 6b) with terminal antennomere elongate oval. Prosternal carinae present (Fig. 6c), reaching a little beyond middle of prosternum, anterior margin of mesoventrite deeply emarginate. Abdominal postcoxal line incomplete (Fig. 6d). Male genitalia (Figs. 6 e–g) with penis guide of tegmen in ventral view elongate, lanceolate in outline, slightly longer than parameres; penis (Fig. 6g) as illustrated. Female genitalia with spermatheca (Figs. 6h, i) strongly curved, infundibulum present. Larva (Figs. 5j, k) black with white or yellow spots. Distribution: India: Delhi; Karnataka; Tamil Nadu; Pondicherry; Orissa; Punjab; West Bengal. Pakistan. Notes: Poorani (2002) transferred Anegleis (Pseudanegleis) perrotteti to Phrynocaria based on the large, posteriorly distinctly divergent eyes, absence of meso- and metatibial spurs, incomplete abdominal postcoxal lines and deep foveae on anterolateral corners of prothoracic hypomeron. Anegleis Iablokoff-Khnzorian, 1982 is an Oriental genus of Coccinellini established with two anomalous subgenera, namely, Anegleis (Anegleis) and Anegleis (Pseudanegleis). The type species of these subgenera were Micraspis cardoni (Weise, 1892) and Coelophora perrotteti Mulsant, 1850, respectively. These two species are externally strikingly similar and mimic each other, but otherwise unrelated and both are distributed in the Indian Subcontinent with A. cardoni having a relatively wider geographic distribution in South Asia. The name Anegleis was first mentioned and included in Iablokoff-Khnzorian’s key to the Palaearctic genera of Coccinellini (Iablokoff-Khnzorian 1979) without a formal, detailed description and illustrations. Later, Iablokoff-Khnzorian (1982) in his conspectus on Palaearctic and Oriental Coccinellini established Anegleis as a new genus with these two subgenera. Iablokoff-Khnzorian (1984: 217) published an erratum and further list of amendments to his 1982 volume, in which he mentioned that Pseudanegleis was congeneric with Egleis adjuncta Mulsant, 1850 from South America and hence Pseudanegleis fell in synonymy with Egleis and Anegleis became its subgenus. Fürsch (1990) also included Egleis (Anegleis) Iablokoff-Khnzorian in his list of valid genera and subgenera of Coccinellidae. But the name Egleis (Anegleis) has not been used much after it was proposed though Anegleis has been widely used in most of the published literature. Anegleis (Pseudanegleis) should be removed from synonymy with Egleis and treated as synonymous with Phrynocaria following Poorani (2002)., Published as part of Poorani, J., Sankararaman, H. & Anusree, S. S., 2021, A review of Indian species of Phrynocaria Timberlake (Coleoptera, Coccinellidae) with notes on synonymy and a new species from South India, hitherto misidentified as Coelophora circumusta (Mulsant), pp. 117-133 in Zootaxa 4926 (1) on pages 124-128, DOI: 10.11646/zootaxa.4926.1.8, http://zenodo.org/record/4500738, {"references":["Mulsant, E. (1850) Species des Coleopteres Trimeres Securipalpes. Annales des Sciences Physiques et Naturelles, d'Agriculture et d'Industrie, publiees par la Societe nationale d'Agriculture, etc., de Lyon, Deuxieme Serie, 2, 1 - 1104. https: // doi. org / 10.5962 / bhl. title. 8953","Crotch, G. R. (1874) A revision of the coleopterous family Coccinellidae. E. W. Janson, London, 311 pp. https: // doi. org / 10.5962 / bhl. title. 8975","Korschefsky, R. (1932) Coleopterorum Catalogus. Pars 120. Coccinellidae II. W. Junk, Berlin, 435 pp.","Gordon, R. D. (1987) A catalogue of the Crotch collection of Coccinellidae (Coleoptera). Occasional Papers on Systematic Entomology, 3, 1 - 46.","Iablokoff-Khnzorian, S. M. (1982) Les Coccinelles Coleopteres-Coccinellidae. Tribu Coccinellini des regions Palearctique et Orientale. Societe nouvelle des editions Boubee, Paris, 568 pp.","Poorani, J. (2002) An annotated checklist of the Coccinellidae (Coleoptera) (excluding Epilachninae) of the Indian Subregion. Oriental Insects, 36, 307 - 383. https: // doi. org / 10.1080 / 00305316.2002.10417335","Weise, J. (1892) Les Coccinellides du Chota-Nagpore. Annales de la Societe Entomologique du Belgique, 36, 16 - 30.","Iablokoff-Khnzorian, S. M. (1979) Genera der Palaarktischen Coccinellini (Coleoptera: Coccinellidae). Entomologische Blatter fur Biologie und Systematik der Kafer, 75 (1 - 2), 37 - 75.","Iablokoff-Khnzorian, S. M. (1984) Notes sur la tribu des Coccinellini (Coleoptera, Coccinellidae). Nouvelle Revue d'Entomologie, New Series, 1 (2), 203 - 220.","Fursch, H. (1990) Valid genera and subgenera of Coccinellidae. Coccinella, 2, 7 - 18."]}

Phrynocaria Timberlake, 1943 Phrynocaria Timberlake, 1943: 34. Type species by original designation, Coccinella congener Billberg in Schönherr 1808: 179 (= Coccinella unicolor Fabricius, 1792). Phrynolemnia Iablokoff-Khnzorian, 1984: 204. Type species by original designation, Coccinella congener Billberg in Schönherr, 1808. Synonymized by Ślipiński 2007: 176. Lemnia (Phrynocaria): Iablokoff-Khnzorian 1984: 205, 210–214. Artemis Mulsant, 1850: 374, 388; 1866: 256.—Preoccupied in Lepidoptera [not Artemis Kirby & Spence, 1828 (Insecta: Lepidoptera: Saturniidae) (Ukrainsky 2006: 400)]. Lemnia (Neoartemis): Ukrainsky 2006: 400 (replacement name for Artemis Mulsant). Anegleis (Pseudanegleis) Iablokoff-Khnzorian, 1982: 296. Type species by original designation, Coelophora perrotteti Mulsant, 1850: 409. —Synonymized by Poorani 2002: 339. Diagnosis: Medium-sized beetles, form broadly oval to almost circular, dorsum moderately to strongly convex, glabrous. Anterior clypeal margin of head straight between lateral projections. Antenna with 11 antennomeres, last four forming a terminal club; eyes large, prominent, eye canthus deep and narrow; frons narrow between eyes, markedly divergent and broadened anteriorly toward antennal insertions. Prothoracic hypomeron with distinct foveae near anterolateral corners. Mesoventrite anteriorly deeply, triangularly emarginate. Abdominal postcoxal lines incomplete, without an oblique dividing line. Meso- and metatibial spurs absent. Phrynocaria may be confused with related genera such as Coelophora Mulsant, 1850 (including Lemnia Mulsant, 1850), Microcaria Crotch, 1871 (formerly a synonym of Coelophora), and Synona Pope, 1989 that share a roughly hemispherical body shape with a narrowly explanate elytral margin; however, it is easily distinguished from these genera by the large eyes, much narrower frons (interocular distance at narrowest point equal to from 0.8 to 1.3 times the width of an eye) and lack of meso- and metatibial spurs. Notes: Timberlake (1943) established Phrynocaria and provided a key to Coelophorine genera, including Phrynocaria, based on the male genitalia. Iablokoff-Khnzorian (1984) treated it as a subgenus of Lemnia. He also established a subgenus Lemnia (Microlemnia) which had Coelophora vidua Mulsant (= Coelophora inaequalis (F.)) as the type species, but also included species that truly belonged to Phrynocaria (e.g. P. unicolor (F.), which is synonymous with P. congener, the type species of Phrynocaria). Lemnia (Microlemnia) was listed as a synonym of Coelophora by Ślipiński et al. (2020) with a note that ‘it was an invalid taxon not accompanied by a description’. Pope (1989) stated that Iablokoff-Khnzorian’s interpretation of Phrynocaria probably includes ‘species related to Lemnia (Artemis) circumusta (Mulsant) ’. Ślipiński et al. (2020) revised the Australo-Pacific species of Phrynocaria and stated that the composition of Phrynocaria was uncertain and its relationships with Coelophora and Lemnia needed further research. He noted that preliminary unpublished research supported the separation of Coelophora and Phrynocaria, but the position of Lemnia remained unresolved. Anegleis (Pseudanegleis) Iablokoff-Khnzorian (1982) is a synonym of Phrynocaria Mulsant as its type species, Anegleis (Pseudanegleis) perrotteti was transferred to Phrynocaria by Poorani (2002). Biology / hosts: Aphids, coccids and psyllids have been recorded as prey of Phrynocaria spp. from Australia (Pope 1989). Australo-Pacific Phrynocaria spp. are known to be associated with soft scales (Hemiptera: Sternorrhyncha: Coccidae) and aphids (Houston 1988; Ślipiński et al. 2020). However, some Indian species of Phrynocaria seem to have a proclivity to feed on whiteflies (Hemiptera: Sternorrhyncha: Aleyrodidae). At least two species, Phrynocaria unicolor (= P. congener) and P. perrotteti (Mulsant), are known to feed on whiteflies in India (Kapur 1940). Phrynocaria perrotteti is predatory on spiralling whitefly (Aleurodicus dispersus Russell) on many host plants in Karnataka (Ramani et al. 2002), Dialeuropora decempuncta Quaintance on mulberry (Narendra Kumar et al. 2013) and unidentified whiteflies on other hosts (personal observation / label data). Phrynocaria unicolor is also known to be a whitefly feeder and is collected in association with Dialeuropora sp. on Persea bombycina, a host plant of muga silkworm, in northeastern India. Wu et al. (2010) studied its comparative biology (under the name P. congener) on a whitefly (Bemisia tabaci (Gennadius)) and an aphid (Myzus persicae (Sulzer)) and found that the developmental period of P. unicolor was shorter and survival rate greater on B. tabaci than M. persicae. Escalona et al. (2017) observed ‘phylogenetically independent food preference transitions from aphidophagy to other food sources in many Coccinellini’, but none of the genera / species of Coccinellini studied by them is known to feed on whiteflies., Published as part of Poorani, J., Sankararaman, H. & Anusree, S. S., 2021, A review of Indian species of Phrynocaria Timberlake (Coleoptera, Coccinellidae) with notes on synonymy and a new species from South India, hitherto misidentified as Coelophora circumusta (Mulsant), pp. 117-133 in Zootaxa 4926 (1) on pages 118-119, DOI: 10.11646/zootaxa.4926.1.8, http://zenodo.org/record/4500738, {"references":["Timberlake, P. H. (1943) The Coccinellidae or Ladybeetles of the Koebele Collection-Part 1. Hawaiian Planters' Record, 47 (1), 1 - 67.","Schonherr, C. J. (1808) Synonymia Insectorum, oder Versuch einer Synonymie Aller bischer bekkanten Insecten nach Fabricii Systema Eleutheratorum etc. geordnet. Erster Band. Eleutherata oder Kafer. Zweuter Theil. Spercheus-Cryptocephalus. Carl Friedr. Marquard, Stockholm, 424 pp.","Fabricius, J. C. (1792) Entomologia Systematica Emendata et Aucta. Secundum Classes, Ordines, Genera, Species, Adjectis, Synonimis, Locis, Observationibus, Descriptionibus. Vol. I. Part 1. Impensis Christ. Gottl. Proft., Hafniae, 330 pp. https: // doi. org / 10.5962 / bhl. title. 122153","Iablokoff-Khnzorian, S. M. (1984) Notes sur la tribu des Coccinellini (Coleoptera, Coccinellidae). Nouvelle Revue d'Entomologie, New Series, 1 (2), 203 - 220.","Slipinski, A. (2007) Australian ladybird beetles (Coleoptera: Coccinellidae), their biology and classification. Australian Biological Resources Study, Canberra, 286 pp.","Mulsant, E. (1850) Species des Coleopteres Trimeres Securipalpes. Annales des Sciences Physiques et Naturelles, d'Agriculture et d'Industrie, publiees par la Societe nationale d'Agriculture, etc., de Lyon, Deuxieme Serie, 2, 1 - 1104. https: // doi. org / 10.5962 / bhl. title. 8953","Kirby, W. & Spence, W. (1828) An Introduction to Entomology: or, Elements of the Natural History of Insects. Vol. III. 5 th Edition. Longman, Rees, Orme, Brown & Green, London, 732 pp. https: // doi. org / 10.5962 / bhl. title. 152792","Ukrainsky, A. S. (2006) Five new replacement ladybird (Coleoptera: Coccinellidae) generic names. Russian Entomological Journal, 15 (4), 399 - 400.","Iablokoff-Khnzorian, S. M. (1982) Les Coccinelles Coleopteres-Coccinellidae. Tribu Coccinellini des regions Palearctique et Orientale. Societe nouvelle des editions Boubee, Paris, 568 pp.","Poorani, J. (2002) An annotated checklist of the Coccinellidae (Coleoptera) (excluding Epilachninae) of the Indian Subregion. Oriental Insects, 36, 307 - 383. https: // doi. org / 10.1080 / 00305316.2002.10417335","Crotch, G. R. (1871) List of Coccinellidae. Self-published by the author, Cambridge, UK.","Pope, R. D. (1989) A revision of the Australian Coccinellidae (Coleoptera). Part 1. Subfamily Coccinellinae. Invertebrate Taxonomy, 2, 633 - 735. https: // doi. org / 10.1071 / IT 9880633","Slipinski, A., Li, J. & Pang, H. (2020) Ladybird beetles of the Australo-Pacific region. Coleoptera: Coccinellidae: Coccinellini. CSIRO Publishing, Clayton South, 231 pp. https: // doi. org / 10.1071 / 9781486303885","Houston, K. J. (1988) Larvae of Coelophora inaequalis (F.), Phrynocaria gratiosa (Mulsant) and P. astrolabiana (Weise) (Coleoptera: Coccinellidae) with notes on their relationships and host records. Journal of the Australian Entomological Society, 27, 199 - 211. https: // doi. org / 10.1111 / j. 1440 - 6055.1988. tb 01525. x","Kapur, A. P. (1940) A note on the ladybird beetles (Coccinellidae) predating upon the cane whitefly, Aleurolobus barodensis Maskell. Current Science, 9, 134.","Ramani, S., Poorani, J. & Bhumannavar, B. S. (2002) Spiralling whitefly, Aleurodicus dispersus, in India. Biocontrol News and Information, 23 (2), 55 N- 62 N.","Narendra Kumar, J. B., Sreenivas, B. T., Divya, S. H. & Shekhar, M. A. (2013) New whitefly species Dialeuropora decempuncta infesting mulberry in Karnataka. Insect Environment, 19 (2), 108 - 109.","Wu, J., Yang, J. I. & Ren, S. X. (2010) Influence of Bemisia tabaci (Gennadius) and Myzus persicae (Sulzer) as preys on biological characteristics of Phrynocaria congener (Billberg). Chinese Journal of Biological Control, 26 (3), 260 - 266.","Escalona, H. E., Zwick, A., Li, H., Li, J., Wang, X., Pang, H., Hartley, D., Jermiin, L. S., Nedved, O., Misof, B., Niehuis, O., Slipinski, A. & Tomaszewska, W. (2017) Molecular phylogeny reveals food plasticity in the evolution of true ladybird beetles (Coleoptera: Coccinellidae: Coccinellini). BMC Evolutionary Biology, 17, 151. https: // doi. org / 10.1186 / s 12862 - 017 - 1002 - 3"]}

Phrynocaria unicolor (Fabricius, 1792) (Figs. 7, 8b) Coccinella unicolor Fabricius, 1792: 267. Coelophora unicolor: Mulsant 1850: 413; 1866: 275.— Crotch 1874: 54.— Korschefsky 1932: 297. Lemnia (Microlemnia) unicolor: Iablokoff-Khnzorian 1982: 262. Phrynocaria unicolor: Yu 2010: 80–82. Coccinella congener Billberg, in Schönherr 1808: 179.—Synonymized by Yu 2010: 80. Coelophora congener: Crotch 1874: 151. Phrynocaria congener: Ren et al. 2009: 226. Coelophora caliginosa Mulsant, 1850: 414.— Crotch 1874: 151.— Weise 1892: 414.— Korschefsky 1932: 297. Coelophora dumortieri Mulsant, 1866: 260.— Crotch 1874: 151.— Weise 1902: 21.— Korschefsky 1932: 297. Coelophora dupasquieri Mulsant, 1866: 261.— Crotch 1874: 151.— Weise 1902: 21.— Korschefsky 1932: 297. Coelophora romani Mulsant, 1866: 273.— Crotch 1874: 160 (as Dysis).— Weise 1892: 22, 24.— Korschefsky 1932: 297. Coelophora petrequini Mulsant, 1866: 276.— Crotch 1874: 154.— Weise 1892: 22, 24.— Korschefsky 1932: 297. Coelophora versipellis Crotch, 1874: 151.— Weise 1892: 20.— Korschefsky 1932: 297. Dysis saundersii Crotch, 1874: 160.— Weise 1892: 22.— Korschefsky 1932: 297. Phrynocaria nigrilimbata Jing, 1986: 307.—Synonymized by Yu 2010: 80. Diagnosis: Length 3.42–4.62 mm, width 3.12–4.20 mm. Form (Figs. 7 a–e) round, hemispherical. Highly variable with numerous variations in elytral pattern and several intermediate forms. Typical form uniformly reddish brown on both sides; at least eight variants or varieties named, commonly found variants (Figs. 7 a–e, 8b) as illustrated. Male genitalia (Figs. 7 i–k) are diagnostic. Distribution: India: Northern and eastern regions (Assam; Bihar; Delhi; Himachal Pradesh; Pondicherry; Jharkhand; Odisha; Uttar Pradesh; Andamans). China. Thailand. Biology / hosts: Associated with whiteflies (Hemiptera: Aleyrodidae) such as Aleurolobus barodensis (Maskell) on sugarcane (Kapur 1940); Dialeuropora decempuncta (Quaintance & Baker) infesting mulberry. Abundant in the oak forests of northeastern region, feeds on Tuberculatus indicus Ghosh (Aphididae) on Quercus acutissima (Shantibala & Singh 1991). It is known to be predatory on the nymphs and adults of the aphid, Schizoneuraphis himalayensis on Persea bombycina in Assam, northeastern India (Easwarareddy & Rajan 2011). Taxonomic notes: Phrynocaria unicolor is a highly variable species with many colour forms and Weise (1892) listed the synonyms, varieties, colour morphs and variants of this species and Korschefsky (1932) also followed this in his catalogue. One female of D. saundersii Crotch (Fig. 8b) (BMNH, London; examined) identified by R. G. Booth through comparison with Crotch’s lectotype (labelled as Coelophora saundersii) was found to be conspecific with P. unicolor as stated by Weise (1892), Korschefsky (1932) and Mader (1937). The type locality of D. saundersii is ‘Lokardaga’ (Type label), located in the present-day Jharkhand state in eastern India. In D. saundersii, the head and pronotum are yellow and the elytra are brownish except lateral borders and apices yellow, with four yellow spots arranged in a 2-2 pattern, first pair touching anterior margin, second pair below midline. Lemnia circumvelata Mulsant, 1850, a replacement name for Coccinella cincta Hope, 1831 described from Nepal, is most likely a synonym of P. unicolor because it is similar to var. cinctipennis Weise, 1892 of P. unicolor. Booth & Pope (1989) designated a lectotype for C. circumvelata and Hope’s type specimen of C. circumvelata (Fig. 8a) (BMNH, examined) looks like a Phrynocaria. It has yellowish head and pronotum with a large black macula on pronotum occupying most of the discal area and yellowish-reddish elytra with black lateral borders about ¼ th as wide as each elytron. Similar variants of P. unicolor from eastern India have been examined (Fig. 7d). There is also a rare variant of P. circumusta with black lateral borders of elytra as in C. circumvelata. However, C. circumvelata is not synonymized because the type could not be dissected for confirmation. Common colour variants of P. unicolor were illustrated by Mader (1937), Kapur (1967), Yu (2010) and Ren et al. (2009)., Published as part of Poorani, J., Sankararaman, H. & Anusree, S. S., 2021, A review of Indian species of Phrynocaria Timberlake (Coleoptera, Coccinellidae) with notes on synonymy and a new species from South India, hitherto misidentified as Coelophora circumusta (Mulsant), pp. 117-133 in Zootaxa 4926 (1) on pages 128-131, DOI: 10.11646/zootaxa.4926.1.8, http://zenodo.org/record/4500738, {"references":["Fabricius, J. C. (1792) Entomologia Systematica Emendata et Aucta. Secundum Classes, Ordines, Genera, Species, Adjectis, Synonimis, Locis, Observationibus, Descriptionibus. Vol. I. Part 1. Impensis Christ. Gottl. Proft., Hafniae, 330 pp. https: // doi. org / 10.5962 / bhl. title. 122153","Mulsant, E. (1850) Species des Coleopteres Trimeres Securipalpes. Annales des Sciences Physiques et Naturelles, d'Agriculture et d'Industrie, publiees par la Societe nationale d'Agriculture, etc., de Lyon, Deuxieme Serie, 2, 1 - 1104. https: // doi. org / 10.5962 / bhl. title. 8953","Mulsant, E. (1866) Monographie des Coccinellides. 1 re Partie Coccinelliens. Savy et Deyrolle, Paris, 294 pp.","Crotch, G. R. (1874) A revision of the coleopterous family Coccinellidae. E. W. Janson, London, 311 pp. https: // doi. org / 10.5962 / bhl. title. 8975","Korschefsky, R. (1932) Coleopterorum Catalogus. Pars 120. Coccinellidae II. W. Junk, Berlin, 435 pp.","Iablokoff-Khnzorian, S. M. (1982) Les Coccinelles Coleopteres-Coccinellidae. Tribu Coccinellini des regions Palearctique et Orientale. Societe nouvelle des editions Boubee, Paris, 568 pp.","Yu, G. (2010) Chinese Lady Beetles (The Subfamily Coccinellinae). China Scientific Press, Beijing, 180 pp. [in Chinese]","Schonherr, C. J. (1808) Synonymia Insectorum, oder Versuch einer Synonymie Aller bischer bekkanten Insecten nach Fabricii Systema Eleutheratorum etc. geordnet. Erster Band. Eleutherata oder Kafer. Zweuter Theil. Spercheus-Cryptocephalus. Carl Friedr. Marquard, Stockholm, 424 pp.","Ren, S. X., Wang, X. M., Pang, H., Peng, Z. Q. & Zeng, T. (2009) Colored pictorial handbook of ladybird beetles in China. Science Press, Beijing, 336 pp. [in Chinese]","Weise, J. (1892) Les Coccinellides du Chota-Nagpore. Annales de la Societe Entomologique du Belgique, 36, 16 - 30.","Weise, J. (1902) Coccinelliden aus der Sammlung des Ungarischen National-Museums. Termeszetrajzi Fuzetek, 25, 489 - 520.","Jing, X. I. (1986) A new species of Phrynocaria Timberlake (Coleoptera: Coccinellidae). Acta Zootaxonomica Sinica, 11 (3), 307 - 308.","Kapur, A. P. (1940) A note on the ladybird beetles (Coccinellidae) predating upon the cane whitefly, Aleurolobus barodensis Maskell. Current Science, 9, 134.","Shantibala, K. & Singh, T. K. (1991) Studies on the vertical distribution of aphidophagous coccinellids (Coleoptera: Coccinellidae) of Manipur and Nagaland. Journal of Aphidology, 5, 39 - 43.","Mader, L. (1937) Evidenz der palaarktischen Coccinelliden und ihrer Aberrationen in Wort und Bild. Entomologische Nachrichten, Troppau, 11, 41 - 66.","Hope, F. W. (1831) Synopsis of the new species of Nepal insects in the collection of Major General Hardwicke. In: Gray, J. E. (Ed.), The Zoological Miscellany. Treutel, Wurtz and Co., London, pp. 21 - 32.","Booth, R. G. & Pope, R. D. (1989) A review of the type material of Coccinellidae (Coleoptera) described by F. W. Pope and by E. Mulsant in the Hope Entomological Collections, Oxford. Entomologica Scandinavica, 20 (3), 343 - 370. https: // doi. org / 10.1163 / 187631289 X 00366","Kapur, A. P. (1967) The Coccinellidae (Coleoptera) of the Andamans. Proceedings of the National Institute of Sciences of India, 32 (B), 148 - 189. [1966]"]}

Phrynocaria perfida Poorani, sp. n. (Figs. 2, 3) Coelophora circumusta sensu Gorham 1903: 344 (not Artemis circumusta Mulsant, 1850; misidentification). Coelophora circumusta var. rufula: Gorham 1903: 344 (not Artemis rufula Mulsant, 1850; misidentification). Diagnosis: The male genitalia of the new species will distinguish it from all other known examples of the genus Phrynocaria, especially the shape of the penis guide which is basally broad and abruptly narrowed around middle to form an elongate lanceolate process with a small rhomboidal expansion in the apical 1/8 bearing minute lateral denticles. This highly variable species (Figs. 2 a–g) has some colour morphs similar to those found in P. circumusta and, given their overlapping distributions, it may not be possible to differentiate the two species based on the elytral colour pattern alone. However, variants of P. perfida with a 2-2-1/2 elytral pattern (Figs. 2c, g) are distinct and easily distinguished from P. circumusta. There are also some minute morphological differences between the two species. If an identifier is fortunate enough to have comparative material of both species at hand it can be seen that P. perfida has a slightly broader body outline and comparatively more widely explanate lateral elytral borders than C. circumusta. Also, the latter appears to be slightly more strongly convex than P. perfida, which is apparent in the lateral view. In both P. circumusta and P. perfida, the explanate borders of elytra are distinctly more coarsely punctate compared to the discal area, but this is more pronounced in the case of P. circumusta. Description: Male: TL: 3.63 mm; TW: 3.26 mm; TL/TW: 1.11; EL/EW: 0.89; PL/PW: 0.48. Form broadly rounded to almost circular in outline, dorsum strongly convex and hemispherical, glabrous except clypeal margin of head with yellowish hairs. Dorsal surface yellowish-orange to reddish in live specimens, yellowish in preserved specimens; scutellar shield black; pronotum with or without a small oval black spot on posterior margin above scutellar shield; elytral pattern with the following variations: (i) immaculate reddish to orange or yellow (Fig. 2a); (ii) with a narrow black lateral margin, anteriorly joining an elongate spot on humeral callus and posteriorly joining a larger apical black patch across elytra (Figs. 2e, f); (iii) each elytron with five black spots arranged in a 2-2-1/2 pattern (Figs. 2c, g), occasionally all spots coalescing to form larger fasciae (Fig. 2d) or reduced in size (Fig. 2b). Ventral side uniform yellowish brown except outer one-third of epipleura black. Head with anterior clypeal margin straight between lateral projections; eye canthus short but deep and broad; eyes prominent, interocular distance at its narrowest about as wide as an eye; head with even and fairly dense punctures, interspaces between punctures weakly reticulate to shiny. Pronotum with lateral sides strongly rounded, almost semicircular, slightly reflexed laterally, forming a gutter, anterolateral corners produced (Fig. 2h); punctures on disc of pronotum dense and slightly larger than those on elytra, shallowly impressed, separated by 1–3 diameters, closer and denser on lateral and posterior margins, interspaces between punctures smooth and shiny. Scutellar shield broadly triangular with few punctures. Elytral borders laterally explanate with a weak gutter; elytral disc densely punctate, punctures separated by 2–5 diameters, interspaces between punctures smooth and shiny; punctation on explanate lateral borders distinctly coarser and closer than that on elytral disc, particularly around base and humeral angles. Prothoracic hypomeron deeply foveolate near anterolateral corners. Prosternal intercoxal process (Fig. 3a) carinate, carinae reaching up to 2/3 length of prosternum; mesoventrite deeply foveate on anterior face, medially with a v-shaped notch and a short longitudinal carina in anterior half. Metaventrite with discrimen. Abdomen with six ventrites, abdominal postcoxal line incomplete (Fig. 3b), apically not recurved, parallel to posterior margin of ventrite 1; posterior margin of ventrite 5 medially very shallowly and that of ventrite 6 slightly more deeply emarginate (Fig. 3d). Legs lacking meso- and metatibial spurs; tarsal claws appendiculate. Epipleura very shallowly and deeply foveolate on level with mid- and hind femoro-tibial apices, respectively. Male genitalia (Figs. 3 h–k) with penis guide of tegmen in ventral view (Fig. 3h) broadest in basal two-fourths and abruptly narrowed around middle, apical half lanceolate, gradually narrowed towards apex, with a small rhomboidal expansion in apical 1/8 bearing finely dentate margins (Fig. 3i); parameres nearly as long as penis guide; penis (Fig. 3j) strongly curved, with a robust capsule, penis apex modified as illustrated (Fig. 3k). Female: TL: 3.93–4.10 mm; TW: 3.41–3.59 mm; TL/TW: 1.11; EL/EW: 0.93; PL/PW: 0.48. Externally similar to male except lateral margins of pronotum less strongly rounded with anterolateral corners produced to a lesser degree (Fig. 2i); posterior margin of ventrite 5 medially very shallowly produced, that of ventrite 6 arcuate (Fig. 3c). Female spermatheca (Figs. 3e, f) with a short sperm duct, infundibulum cup-shaped; coxites of blade and handle type. Material examined: Holotype: male, “ INDIA: Kerala: Kakkadampoyil, 06.xi.2018; H. Sankararaman ” (BMNH). Paratypes: One female with same data (BMNH); “ Kerala: Kayamkulam, 7.iii.2019, on sweeping, Anusree” (one female, NBAIR); “ Kerala: Vellayani, 24.ii.2018, on mulberry, Anusree ” (2 females, NBAIR); Other material: Kanara, S. India / Andrewes Bequest (3, BMNH); Kanara, S. India/ N. Kanara, T. R. Bell (1, BMNH). Distribution: India: Tamil Nadu (Nilgiri hills); Karnataka; Kerala. Etymology: The specific epithet is a Latin adjective in nominative case (perfida L.= ‘false’), alluding to its external similarity to C. circumusta. Notes: Gorham’s (1903) specimens of ‘ C. circumusta ’ from Kanara (in Karnataka, South India) were described by him as ‘with the basal half of the elytra black, running up the suture to the scutellum […] very near to a specimen in my collection labelled mandarinea [sic].’ This description applies to the variants of P. perfida sp. n. (Fig. 2e, f) illustrated here. His description of ‘ Coelophora circumusta var. rufula ’ from Nilgiri Hills (South India) corresponds to the immaculate form of P. perfida sp. n. (Fig. 2a). However, the spotted form of P. perfida is distinctive and P. circumusta does not have this kind of variation. One specimen from Sri Lanka examined at BMNH (“Nedunkeni, 4.04 / Ceylon 1924.37”) was similar to this species, but it could not be dissected for confirmation. Material examined at BMNH, London, indicates that P. circumusta also is distributed in South India and it is not always easy to separate it from P. perfida sp. n. by the punctation alone if the latter is lacking elytral spots. Hence, these two species are best separated by their genitalia., Published as part of Poorani, J., Sankararaman, H. & Anusree, S. S., 2021, A review of Indian species of Phrynocaria Timberlake (Coleoptera, Coccinellidae) with notes on synonymy and a new species from South India, hitherto misidentified as Coelophora circumusta (Mulsant), pp. 117-133 in Zootaxa 4926 (1) on pages 121-123, DOI: 10.11646/zootaxa.4926.1.8, http://zenodo.org/record/4500738, {"references":["Gorham, H. S. (1903) On Coleoptera collected in India by MM. H. E. and H. L. Andrewes, Families Malacodermata, Erotylidae, Endomychidae and Coccinellidae. Annales de la Societe Entomologique de Belgique, 47, 323 - 347.","Mulsant, E. (1850) Species des Coleopteres Trimeres Securipalpes. Annales des Sciences Physiques et Naturelles, d'Agriculture et d'Industrie, publiees par la Societe nationale d'Agriculture, etc., de Lyon, Deuxieme Serie, 2, 1 - 1104. https: // doi. org / 10.5962 / bhl. title. 8953"]}

Phrynocaria eberti (Bielawski) (Fig. 8c) Coelophora eberti Bielawski, 1972: 304. Phrynocaria eberti: Miyatake 1985: 21.— Poorani 2002: 338. Diagnosis: This species can be identified by its orange to yellow coloured body with five black spots on each elytron arranged in a 2-2-1 pattern as illustrated (Fig. 8c). Male genitalia illustrations by Bielawski (1972) confirm that it is indeed a Phrynocaria, but Iablokoff-Khnzorian (1982) included it under Lemnia (s. str.). Only a single female from northern India was examined and the genitalia could not be studied. Distribution: India. Nepal., Published as part of Poorani, J., Sankararaman, H. & Anusree, S. S., 2021, A review of Indian species of Phrynocaria Timberlake (Coleoptera, Coccinellidae) with notes on synonymy and a new species from South India, hitherto misidentified as Coelophora circumusta (Mulsant), pp. 117-133 in Zootaxa 4926 (1) on page 131, DOI: 10.11646/zootaxa.4926.1.8, http://zenodo.org/record/4500738, {"references":["Bielawski, R. (1972) Die Marienkafer (Coleoptera: Coccinellidae) aus Nepal. Fragmenta Faunistica, 18, 283 - 312. https: // doi. org / 10.3161 / 00159301 FF 1972.18.16.283","Miyatake, M. (1985) Coccinellidae collected by the Hokkaido University Expedition to Nepal Himalaya, 1968 (Coleoptera). Insecta Matsumurana, New Series, 30, 1 - 33.","Poorani, J. (2002) An annotated checklist of the Coccinellidae (Coleoptera) (excluding Epilachninae) of the Indian Subregion. Oriental Insects, 36, 307 - 383. https: // doi. org / 10.1080 / 00305316.2002.10417335","Iablokoff-Khnzorian, S. M. (1982) Les Coccinelles Coleopteres-Coccinellidae. Tribu Coccinellini des regions Palearctique et Orientale. Societe nouvelle des editions Boubee, Paris, 568 pp."]}