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Ladybirds (family Coccinellidae) are one of the most diverse groups of beetles and globally comprise over 6000 species. Despite their scientific and economic significance, the taxonomy of Coccinellidae remains unstable, and we still know little about their evolutionary history. By using a small number of genes, previous phylogenetic analyses have not reliably resolved the relationships among major ladybird lineages. In this study, we sequenced 94 nuclear protein-coding genes for 214 species of Coccinellidae and 14 outgroups, covering 90 genera and 35 tribes. We found that nucleotide compositional heterogeneity is present among ladybird tribes so that phylogenetic inference at the amino acid level is more reliable than at the DNA level. Based on the maximum likelihood analyses of the amino acid dataset, we recognize three subfamilies in Coccinellidae: Microweiseinae, Monocoryninae stat. nov., and Coccinellinae. The subfamily relationships are strongly supported as (Microweiseinae, (Monocoryninae stat. nov., Coccinellinae)). The tribes of ladybirds are mostly monophyletic, except Ortaliini, Sticholotidini, Scymnini, and Coccidulini. The phylogenetic relationships among tribes of Coccinellinae are still not well resolved, with many nodes weakly supported. Our divergence time analysis suggests that the crown group of extant lady beetles arose in the Early Cretaceous ~ 143 million years ago (Mya) and experienced a rapid diversification during the Late Cretaceous (120–70 Mya). We hypothesize that the boom of angiosperms in the Late Cretaceous promoted the diversification of herbivorous Sternorrhyncha insects, especially aphids, which in turn drove the rapid radiation of predatory lady beetles. In summary, our work provides a comprehensive time-calibrated phylogeny of Coccinellidae that provides a sound framework for revising their classification and understanding the origin of their biodiversity.

Argosadalia gen. nov. (Figs. 1���7, 11 a���e, 15���18) type species: Argosadalia priscilla sp. nov. Diagnosis. Distinguished from other members of the Neda group by a combination of: body oval, small-sized, with length 5.2 mm or less, elytral margins narrowly but distinctly explanate, with marginal bead (Figs. 1���4, 11a, d). Prosternal intercoxal process lacking carinae. Elytral epipleuron (Fig 11b, d) narrow, with maximum width about 1/7 width of body, subhorizontal in anterior half, planar to weakly concave. Hind margin of abdominal ventrites 1���4 (inclusive of flexure bands) sublinear when abdomen is held flat (Figs. 11b, c); postcoxal line of first abdominal ventrite joined to oblique line and obliterated beyond, forming modified V shape (Fig. 7, 11b). Basal lobe of male genitalia widest before apex, apically bifurcate, terminating in pair of parallel tapered projections separated by Ushaped emargination (Fig. 11e, 16). Spermatheca of female genitalia (Fig. 18) with nodulus joined to cornu to form distinct outer angle, shaped like the number ���2,��� but with apex of nodulus tapered distally, slightly down-curved at point of attachment to spermduct. Remarks. Within the Neda -group the new genus resembles some of the smaller members of Neoharmonia, but the latter have the prosternal intercoxal process with a pair of convergent carinae separated by a triangular depression, abdominal postcoxal line continued laterally beyond the juncture with the oblique line (Fig. 9), basal lobe of male genitalia with the apex undivided (Fig. 12e), and female spermatheca with a long straight tapered nodulus that flows directly from the cornu without a distinct outer angle (Fig. 19). The genus Neda has the same form of the female spermathecal capsule as the new genus and is probably closely related. However, it can be easily distinguished (Figs. 13 a���e) by the larger body size (5.2���10.0 mm in length), broadly explanate elytral margin, epipleuron strongly concave and steeply descending externally, from about �����1/3 body width at widest part, hind margin of abdominal ventrite 4 distinctly arcuately emarginate in both sexes, and the basal lobe of the male genitalia widened apically, with projections diverging distally. Mononeda (Figs. 14 a���e) also falls within the ��� Neda -group��� and differs from the new genus by the much larger body size (length 7.5���12.4 mm), complete lack of an elytral marginal bead, elytral margin broadly explanate, and elytral epipleuron very wide, concave and descending externally, from about 1/3���1/2 body width at widest part, with ventral epipleural carina complete to apex. The new genus superficially resembles the Holarctic genus Adalia in its relatively small size, general body shape, and similar form of the abdominal postcoxal line (Figs. 7, 8), but Adalia males have an undivided basal lobe, and a differently shaped penis with a strongly pigmented penis capsule. Adalia females have a smaller spermathecal accessory gland and the spermduct with a more robust and heavily sclerotized sheath. Etymology. Argosadalia (gender feminine) formed from a combination of Argos (= shining, bright, from Greek) + Adalia (= noble, from Hebrew). In Greek mythology, Argos was the name of the hundred-eyed servant of the goddess Hera, who had eyes all over his body. This name was selected because of the numerous white spots on the elytra which resemble eyes. The second part of the name was used because of the superficial similarity of this lady beetle to members of the lady beetle genus Adalia., Published as part of Vandenberg, Natalia J., 2019, A new monotypic genus and new species of lady beetle (Coleoptera: Coccinellidae Coccinellini) from western South America, pp. 413-422 in Zootaxa 4712 (3) on pages 417-418, DOI: 10.11646/zootaxa.4712.3.7, http://zenodo.org/record/3586422

A new monotypic genus Argosadalia gen. nov. and new species A. priscilla sp. nov. are described from the forests of Bolivia, Ecuador, and Peru. The new taxon is placed in the tribe Coccinellini in a group informally designated as the “Neda-group,” and comprising the genera Neda Mulsant; Mononeda Crotch; Neoharmonia Crotch and Argosadalia. Key morphological and anatomical characters of the new taxa are described and illustrated and the character states that define the Neda-group are reviewed.

Andrzej Ślipiński, 2007 Andrzej Ślipiński 2007:92. Type species: Andrzej antennatus Ślipiński, 2007, by original designation. Diagnosis. This monotypic genus is based on a single male specimen that has 11 antennomeres and the terminal antennomere extremely large and elongate (length more than 2�� width). Remarks. In a more recent work by Pang and Ślipiński (2009), another species, Diomus sedani (Blackburn,1889), is diagnosed with similar proportions to the terminal antennomere, although less enlarged overall than that of A. antennatus. Unfortunately, the latter species is known from females only, so characteristics of the male genitalia diagnosed for Andrzej cannot be compared with it. Ślipiński (2007) indicates that the antennal club of A. antennatus consists of a single antennomere; however, we observed that the antennomeres gradually increase in width beginning with antennomere 9, and therefore consider that the last 3 antennomeres together constitute the club., Published as part of Vandenberg, Natalia J. & Hanson, Paul E., 2019, Overview of the lady beetle tribe Diomini (Coleoptera: Coccinellidae) and description of a new phytophagous, silk-spinning genus from Costa Rica that induces food bodies on leaves of Piper (Piperaceae), pp. 255-285 in Zootaxa 4554 (1) on page 264, DOI: 10.11646/zootaxa.4554.1.9, http://zenodo.org/record/2623565, {"references":["Slipinski, S. A. (2007) Australian ladybird beetles (Coleoptera: Coccinellidae). Their biology and classification. ABRS, Canberra, 288 pp.","Pang, H. & Slipinski, A. (2009) Revision of the Australian Coccinellidae (Coleoptera). genus Diomus Mulsant. Part 1. Annales Zoologici, 59 (4), 641 - 698. https: // doi. org / 10.3161 / 000345409 x 485008"]}

Vandenberg, Natalia J., Hanson, Paul E. (2019): Overview of the lady beetle tribe Diomini (Coleoptera: Coccinellidae) and description of a new phytophagous, silk-spinning genus from Costa Rica that induces food bodies on leaves of Piper (Piperaceae). Zootaxa 4554 (1): 255-285, DOI: https://doi.org/10.11646/zootaxa.4554.1.9

Tribe Diomini Gordon, 1999 (Figs. 1 a���d) Diomini Gordon 1999:3. Type genus: Diomus Mulsant, 1850 Included taxa (new circumscription, excluding Magnodiomus and Erratodiomus which we transfer to Hyperaspidini; see ��� Remarks,��� below): Diomus, Decadiomus, Heterodiomus, Dichaina, Andrzej, and the new genus described herein Diagnosis. Size minute to small (1.1���3.5 mm), pubescent; antenna (Figs. 1a,b) composed of 10 or 11 antennomeres; pedicel bead like, articulated with and slightly narrower than scape, antennomere 3 elongate, 1.5���3�� as long as antennomere 4; distal 3, 4, or 5 antennomeres forming slightly flattened asymmetrical compact club with oblique to truncate apex bearing concentration of short sensory setae (=sensilla). Mentum (Fig. 1a) subtrapezoidal, tapered posteriorly, typically with anterior margin concave or bicuspidate. Terminal maxillary palpomere in repose free, not partially inserted beneath mentum, more or less expanded distally with sensory surface directed anteriorly or anteromedially. Tarsi trimerous (Fig. 1c). Tibial spurs lacking. Abdomen with 6 visible ventrites; ventrite 1 and 2 partially fused medially. Abdominal postcoxal line (Fig. 1d) curving posterolaterally, merging with posterior margin of ventrite. Male genitalia with basal lobe (=penis guide sens. Ślipiński) distinctly asymmetrical to roughly symmetrical at least in outline; penis capsule with outer arm distinct to obsolete (Figs. 9���12). Female genitalia with spermathecal capsule well developed, except vestigial or absent in some Australian species; capsule with simple angular or C-shaped form, vermiform throughout or with swollen basal chamber; cornu of moderate length, not convoluted; ramus and nodulus sessile to weakly projecting; ramus with or without short beak-like projection (apodeme) overhanging attachment of accessory gland. Coxites teardrop or rhombus-shaped, with widely arcuate posterior margin. Members of Diomini are most easily confused with other small pubescent lady beetles. The shape of the abdominal postcoxal line and wide posterior margin of the coxites will distinguish diomines from both Scymnus and Nephus (including its subgenera sens. Gordon 1985 which are now often accorded full generic status). In Scymnus and Nephus the postcoxal line does not reach the hind margin of the ventrite and the outer end is often recurved toward the base of the ventrite; the coxites are narrow and distally tapered rather than broad. Among New World species, the basal lobe of Diomini is distinctly asymmetrical, while that of Scymnus is bilaterally symmetrical. Furthermore, Scymnus can be distinguished from Diomini by the possession of cryptotetramerous tarsi, and Nephus is distinct in having the basal two antennomeres fused or tightly joined (non-articulated). Pubescent members of the newly recircumscribed tribe Hyperaspidini (see ���Remarks,��� below) differ from Diomini in possessing an antenna with a fusiform club (Figs. 2a,b) bearing a proliferation of setae in a membranous area on the inner (medially facing) surface of the terminal antennomere and smaller membranous area on the inner distal margin of the penultimate antennomere (antennae directed anteriorly); terminal maxillary palpomere in repose (Fig. 2a, left side of image) with inner edge contiguous with or partially inserted beneath anterolateral lobe of large cordate mentum, with oblique sensory surface facing dorsally and pressed against ventral surface of head, and tarsi cryptotetramerous (Fig. 2c). Also in Hyperaspidini the abdominal postcoxal line (Fig. 2d) does not reach the posterior margin of the ventrite and/or the outer end is recurved and directed toward or attains the anterior margin of the ventrite (see corresponding character states for Diomini (Fig. 1) listed in the paragraph above this one). Members of the subtribe Selvadiina (Hyperaspidini) can further be distinguished from Diomini based on the extremely elongate and convoluted cornu of the female spermathecal capsule (Figs. 3���4). Remarks. Ślipiński (2007) diagnoses the Diomini as lacking interfacetal setae, an ocular canthus, and stylus of the coxites (=female genital plates or hemisternites) (Table 2). Our own random sampling of Diomini exemplars in the USNM suggests that these structures are normally present, but at times difficult to see or lost through abrasion. Published digital images of Australian species also document the presence of these structures, at least in some species (e.g. Ślipiński 2007:figs. 298 & 309; Pang & Ślipiński 2010:25d, showing coxites bearing styli; Pang & Ślipiński 2009:fig. 17b, showing frontal view of specimen with eyes bearing canthi and interfacetal setae). Other character states involving the number of antennomeres in the club and the presence or absence of an infundibulum in the female genitalia are difficult to assess because no consistent method for delineating the club has been presented, and the term ���infundibulum��� has not been well defined nor treated consistently by various authors. When Ślipiński (2007) indicated that certain genera ���probably do not belong in Diomini ��� we assume from his later remarks that he intended to name the two taxa related to Selvadius ��� Erratodiomus and Magnodiomus ���but instead mentioned Erratodiomus and Heterodiomus. Gordon (1985) originally misinterpreted the unusual convoluted cornu of the spermathecal capsule in Selvadius (Fig. 3) as a continuation of the spermduct, and assumed that the spermatheca was lacking. He correctly interpreted the homologous structures in Magnodiomus and Erratodiomus (Fig. 4) (Gordon 1999) but failed to consider their placement in Selvadiini, possibly due to his original misinterpretation. Gordon placed his new tribe Selvadiini in the subfamily Scymninae, but discussed similarities between Selvadius and certain hyperaspidines, even referring to them as Hyperaspidinae in his tribal diagnosis. Vandenberg (2002) indicated that Selvadiini would be better placed in the Hyperaspidinae along with Hyperaspidini and Brachiacanthini. These similarities had been previously noted by Whitehead (1967) who suggested that Selvadius may bear a closer affiliation to Hyperaspidius than to Scymnus. In addition, Vandenberg and Perez-Gelabert (2007) noted that two South American genera, Mimoscymnus and Planorbata, originally placed in Coccidulini (Coccidulinae) (Gordon 1994), also belong in Hyperaspidinae. Ślipiński (2007) concurred about the improper placement of Planorbata but referred to it as ���a ���Scymninae��� genus,��� possibly using the latter subfamily in the broad sense of Sasaji, which would have included Hyperaspis and allied genera. Seago et al. (2011) reduced Hyperaspidinae to tribal level and placed Brachiacanthini as a synonym of Hyperaspidini. This action was followed by Gordon et al. (2014), although inconsistently within that paper, and mentioned but not employed in a subsequent work in that series (Canepari et al. 2016). We follow Seago et al. (2011) and various other modern authors in treating the Hyperaspidini at tribal level, but recognize four distinct subtribes: Hyperaspidina, Brachiacanthina, Selvadiina stat. nov. (for Selvadius, Magnodiomu s and Erratodiomus) and Mimoscymnina subtribe nov. (for Mimoscymnus and Planorbata; type genus= Mimoscymnus), as distinct lineages each defined by one or more autapomorphies of the male and female genitalia. The placement of the Selvadiina in Hyperaspidini is supported by the molecular phylogenies of Seago et al. (2011) and Robertson et al. (2015) who show Selvadius as clustering more closely with Brachiacantha and Hyperaspis than with Diomus. The placement of Mimoscymnina in Hyperaspidini is provisional, and based only on external morphological characters (Fig. 2) as there has not yet been a molecular study involving either Mimoscymnus or Planorbata. Members of Mimoscymnina can be readily distinguished from other members of Hyperaspidini by the elongate triangulate coxites of the female genitalia, and the male genitalia with the trabes much longer than the basal lobe and basal piece combined., Published as part of Vandenberg, Natalia J. & Hanson, Paul E., 2019, Overview of the lady beetle tribe Diomini (Coleoptera: Coccinellidae) and description of a new phytophagous, silk-spinning genus from Costa Rica that induces food bodies on leaves of Piper (Piperaceae), pp. 255-285 in Zootaxa 4554 (1) on pages 260-261, DOI: 10.11646/zootaxa.4554.1.9, http://zenodo.org/record/2623565, {"references":["Gordon, R. D. (1999) South American Coccinellidae (Coleoptera), Part VI: A systematic revision of the South American Diomini, new tribe (Scymninae). Annales Zoologici, 49 (1), 1 - 219.","Gordon, R. D. (1985) The Coccinellidae (Coleoptera) of America north of Mexico. Journal of the New York Entomological Society, 93, 1 - 912.","Slipinski, S. A. (2007) Australian ladybird beetles (Coleoptera: Coccinellidae). Their biology and classification. ABRS, Canberra, 288 pp.","Pang, H. & Slipinski, A. (2010) Revision of the Australian Coccinellidae (Coleoptera). Genus Diomus Mulsant. Part 2. Annales Zoologici, 60 (4), 645 - 702. https: // doi. org / 10.3161 / 000345410 X 550382","Pang, H. & Slipinski, A. (2009) Revision of the Australian Coccinellidae (Coleoptera). genus Diomus Mulsant. Part 1. Annales Zoologici, 59 (4), 641 - 698. https: // doi. org / 10.3161 / 000345409 x 485008","Vandenberg, N. J. (2002) Coccinellidae Latreille 1807. In: Arnett, R. H. Jr., Thomas, M. C., Skelley, P. E. & Frank, J. H. (Eds.), American Beetles. CRC Press, Boca Raton, pp. 371 - 389.","Whitehead, V. B. (1967) The validity of the higher taxonomic categories of the tribe Scymnini (Coleoptera: Coccinellidae). Dissertation, Doctor of Philosophy in Entomology, University of California, Berkeley, 312 pp. [unpublished thesis]","Vandenberg, N. J. & Perez-Gelabert, D. E. (2007) Redescription of the Hispaniolan ladybird genus Bura Mulsant (Coleoptera: Coccinellidae) and justification for its transfer from Coccidulinae to Sticholotidinae. Zootaxa, 1586 (1), 39 - 46. https: // doi. org / 10.11646 / zootaxa. 1586.1.4","Gordon, R. D. (1994) South American Coccinellidae (Coleoptera). Part IV: definition of Exoplectrinae Crotch, Azyinae Mulsant, and Coccidulinae Crotch; a taxonomic revision of Coccidulini. Revista Brasileira de Entomologia, 38, 681 - 775.","Seago, A. E., Giorgi, J. A., Li, J. & Slipinski, A. (2011) Phylogeny, classification and evolution of ladybird beetles (Coleoptera: Coccinellidae) based on simultaneous analysis of molecular and morphological data. Molecular Phylogenetics and Evolution, 60 (1), 137 - 151. https: // doi. org / 10.1016 / j. ympev. 2011.03.015","Gordon, R. D., Canepari, C. & Hanley, G. A. (2014) South American Coccinellidae (Coleoptera), Part XVI: systematic revision of Brachiacantha Dejean (Coccinellidae: Hyperaspidini). Insecta Mundi, 0 390, 1 - 76.","Canepari, C., Gordon, R. D. & Hanley, G. A. (2016) South American Coccinellidae (Coleoptera), Part XVII: Systematic revision of the genera Cyrea Gordon and Canepari and Tiphysa Mulsant (Hyperaspidinae: Brachiacanthini). Insecta Mundi, 0 486, 1 - 180.","Robertson, J. A., Slipinski, A., Moulton, M., Shockley, F. W, Giorgi, A., Lord, N. P., McKenna, D. D., Tomaszewska, W., Forrester, J., Miller, K. B., Whiting, M. F. & McHugh, J. V. (2015) Phylogeny and classification of Cucujoidea and the recognition of a new superfamily Coccinelloidea (Coleoptera: Cucujiformia). Systematic Entomology, 40, 745 - 778. https: // doi. org / 10.1111 / syen. 12138"]}

Heterodiomus Br��thes, 1925 Heterodiomus Br��thes 1925:155. Type species: Heterodiomus darwini Br��thes, 1925, by subsequent designation of Korschefsky 1931. Diagnosis. This genus was described but not diagnosed by Br��thes (1925). The species currently included (Gordon 1999; Gordon & Gonz��lez 2003) are distinguished from other members of the New World Diomini by the combination of: antennae composed of 11 antennomeres, prosternum shaped like a short stemmed T, long anterior to coxal cavities (subequal to length of cavity), intercoxal process with carinae short, not extending to anterior margin, and suture between abdominal ventrites 1 and 2 partially obliterated. Members of Heterodiomus are strongly united by the shape of the female genitalia which have a bulbous base to the spermatheca and distally tapered sclerotized sheath in the basal half of the sperm duct (= ���thorn-like infundibulum��� sens. Gordon 1999). Although this combination of genital characteristics does not occur in the other Diomini illustrated in Gordon (1999), the corresponding structures in most species have not yet been evaluated, and a very similar configuration (Fig. 6) is found in the new genus described herein., Published as part of Vandenberg, Natalia J. & Hanson, Paul E., 2019, Overview of the lady beetle tribe Diomini (Coleoptera: Coccinellidae) and description of a new phytophagous, silk-spinning genus from Costa Rica that induces food bodies on leaves of Piper (Piperaceae), pp. 255-285 in Zootaxa 4554 (1) on page 263, DOI: 10.11646/zootaxa.4554.1.9, http://zenodo.org/record/2623565, {"references":["Brethes J. (1925) Sur une collection de Coccinellides (et un Phalacridae) du British Museum. Anales del Museo de Historia Natural de Buenos Aires, 33, 145 - 175. [in French]","Korschefsky, R. (1931) Pars. 118, Coccinellidae. I. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus. W. Junk, Berlin, pp. 1 - 224.","Gordon, R. D. (1999) South American Coccinellidae (Coleoptera), Part VI: A systematic revision of the South American Diomini, new tribe (Scymninae). Annales Zoologici, 49 (1), 1 - 219.","Gordon, R. D. & Gonzalez, G. (2003) Descriptions of the male of Heterodiomus marchali Brethes and a new species of Chilean Heterodiomus Brethes (Coleoptera: Coccinellidae: Scymninae: Diomini). Insecta Mundi, 17 (3 - 4), 237 - 239."]}

Decadiomus Chapin, 1933 Decadiomus Chapin 1933:96. Type species: Diomus bahamicus Casey, 1899, by original designation. Diagnosis. In the New World fauna, Decadiomus species can be distinguished from most previously known Diomini by the combination of: antenna composed of 10 antennomeres, prosternum shaped like a short stemmed Y, short anterior to coxal cavities (about �� diameter of cavity), intercoxal process with carinae extending to anterior margin or nearly so. Decadiomus differs from the new genus described in the present paper (which also has only 10 antennomeres) by having free living, predatory larvae, and also by characteristics of the male genitalia: penis with apical flagellum (Fig. 7), and outer arm of capsule truncated or strongly reduced and not enclosing the ejaculatory duct prior to its entry into the main penis tube (Figs. 11���12). Genital characteristics of Decadiomus males and females are included in the generic key, below, and also discussed by Gordon (1976) (as Diomus, floridanus group and bigemmeus group), and by Segarra-Carmona and Otero (2014). These characteristics may serve to differentiate Decadiomus species from the occasional occurrence of Diomini with 10 antennomeres from areas outside of the New World (e.g. in some individuals of Diomus flavolaterus (Lea) from Australia). Remarks. The following species previously classified in Diomus are hereby transferred and represent new combinations within Decadiomus: D. balteatus (LeConte, 1878), D. floridanus (Mulsant, 1850), D. amabilis (LeConte, 1852), D. liebecki (Horn, 1895), D. myrmidon (Mulsant, 1850), D. humilis (Gordon, 1976), D. pseudotaedatus (Gordon, 1976), D. taedatus (Fall, 1901), D. bigemmeus (Horn, 1895), and D. austrinus (Gordon, 1976). Decadiomus seini Segarra, 2014 is placed as a junior synonym of D. austrinus based on the original type descriptions and illustrations, and a comparison between specimens from Puerto Rico and the type material of D. austrinus., Published as part of Vandenberg, Natalia J. & Hanson, Paul E., 2019, Overview of the lady beetle tribe Diomini (Coleoptera: Coccinellidae) and description of a new phytophagous, silk-spinning genus from Costa Rica that induces food bodies on leaves of Piper (Piperaceae), pp. 255-285 in Zootaxa 4554 (1) on pages 263-264, DOI: 10.11646/zootaxa.4554.1.9, http://zenodo.org/record/2623565, {"references":["Chapin, E. A. (1933) A new genus of West Indian Coccinellidae (Coleoptera). Proceedings of the Biological Society of Washington, 46, 95 - 99.","Casey, T. L. (1899) A revision of the American Coccinellidae. Journal of the New York Entomological Society, 7 (2), 71 - 169.","Gordon, R. D. (1976) The Scymnini of the United States and Canada: Key to genera and revision of Scymnus, Nephus and Diomus. Bulletin of the Buffalo Society of Natural Sciences, 28, 1 - 362.","Segarra-Carmona, A. E. & Otero, M. (2014) Four new ladybug species belonging to Decadiomus Chapin (Coleoptera: Coccinellidae) from Puerto Rico. Neotropical Entomology, 43 (6), 555 - 563. https: // doi. org / 10.1007 / s 13744 - 014 - 0243 - 8"]}

A new genus of lady beetle, Moiradiomus gen. nov. (Coleoptera: Coccinellidae Latreille, 1807: Diomini Gordon, 1999 ), and four new species are described from Costa Rica, representing the first known occurrences of obligate phytophagous lady beetle species outside of the tribe Epilachnini Mulsant, 1846 (sens. Ślipiński 2007). The new species are described, illustrated and keyed, and their life histories discussed. Each species of Moiradiomus occurs on a separate species of Piper L., 1753 (Piperaceae Giseke, 1792), where the larva constructs a small silken tent between leaf veins and inside this shelter induces the production of food bodies, which are its exclusive source of food. Background information is provided on lady beetle trophic relations and other insect/Piper symbioses. The taxonomic history of Diomus Mulsant, 1850 and related species in the tribe Diomini is reviewed and existing errors in observation, interpretation, identification, and classification are corrected in order to provide a more meaningful context for understanding the new genus. The tribe Diomini is rediagnosed and recircumscribed to include Diomus, Decadiomus Chapin, 1933, Heterodiomus Brèthes, 1925, Dichaina Weise, 1923, Andrzej Ślipiński, 2007, and Moiradiomus. Magnodiomus Gordon, 1999 and Erratodiomus Gordon, 1999 are removed from Diomini and transferred to Hyperaspidini Costa, 1849, subtribe Selvadiina Gordon, 1985 stat. nov. Mimoscymnus Gordon, 1994 and Planorbata Gordon, 1994, originally described in Coccidulini Mulsant, 1846 are also transfered to Hyperaspidini and placed in Mimoscymnina subtribe nov. (type genus Mimoscymnus). The main morphological characters distinguishing Diomini and Hyperaspidini are described and illustrated. A key to genera of Diomini sensu novo is provided. The identification of the Australian Diomus species illustrated in Gordon’s publication on North American lady beetles is corrected from D. pumilio Weise, 1885 to D. tenebricosus (Boheman, 1859), however specimens recently collected in California do not match these genitalic illustrations and are identified as true D. pumilio. The following species of Diomus are transferred to Decadiomus as new combinations: D. balteatus (LeConte, 1878), D. floridanus (Mulsant, 1850), D. amabilis (LeConte, 1852), D. liebecki (Horn, 1895), D. myrmidon (Mulsant, 1850), D. humilis (Gordon, 1976), D. pseudotaedatus (Gordon, 1976), D. taedatus (Fall, 1901), D. bigemmeus (Horn, 1895), and D. austrinus (Gordon, 1976). Decadiomus seini Segarra, 2014 is placed as a junior synonym of D. austrinus. The following new species of Moiradiomus are described: M. clotho sp. nov., M. lachesis sp. nov., M. atopos sp. nov., M. nanita sp. nov.

Dichaina Weise, 1923 Dichaina Weise 1923:145. Type species: Dichaina inornata Weise, 1923, by monotypy. Diagnosis. This monotypic genus is very similar to Diomus, but can be easily distinguished by the sharp submarginal carinae separating the anterolateral pronotal angles from the disc., Published as part of Vandenberg, Natalia J. & Hanson, Paul E., 2019, Overview of the lady beetle tribe Diomini (Coleoptera: Coccinellidae) and description of a new phytophagous, silk-spinning genus from Costa Rica that induces food bodies on leaves of Piper (Piperaceae), pp. 255-285 in Zootaxa 4554 (1) on page 264, DOI: 10.11646/zootaxa.4554.1.9, http://zenodo.org/record/2623565, {"references":["Weise, J. (1923) Results of Dr. E. Mjoberg's Swedish Scientific Expedition to Australia 1910 - 1913. 31. Chrysomeliden und Coccinelliden aus Queensland. Arkiv for Zoologi, 15 (12), 1 - 150."]}

Diomus lupusapudoves, sp. nov. (Figs. 1–7, 9, 11–15) Diagnosis. Due to variability in the dorsal color pattern (Figs. 4–7), the new species is best identified by the exact configuration of the adult male genital structures (Figs. 11–12) and by the myrmecophilous, onisciform (=platyform) larva (Figs. 14–15) possessing a finely granulate dorsum devoid of setae, and with the epipleurum of each abdominal segment extended into an oblique lateral plate, forming a protective skirt-like border to shield the underside of the body and appendages. This species closely resembles another myrmecophile, Diomus thoracicus (Fabricius, 1801), recorded from northern South America and the Antilles, to Mexico and southern Florida (Peck 2015), but the adult of our new species is smaller on average (1.7–2.1 mm vs. 2.0– 2.4 mm). Diomus thoracicus male genitalia (illustrated in Gordon 1999) are structurally quite similar as well, but possess at least 1.5× the number of setae along the outer margins of each paramere and have the trabes strongly inflated in apical 2/3. The larva of D. thoracicus is also onisciform (Vantaux et al. 2010; Roux et al. 2017), but has the dorsal surface a uniform light bluish gray except for the contrasting white epipleurum of each abdominal segment. The perimeter of the body is distinctly fimbriate in D. thoracicus, whereas the new species has only a few setae on the head and posterior end of the abdomen. In addition, the head of the D. thoracicus larva projects anteriorly, and is not enclosed laterally by the sides of the pronotum as it is in the new species. Although larvae of both species are myrmecophilous, D. thoracicus larvae are intranidal parasites that feed on ant brood, whereas larvae of the new species are coccidophagous and occur on open vegetation. Another species, D. urban Gordon, 1999 (Fig. 8), was found infrequently alongside the new species among colonies of C. viridis in the coffee agroecosystem that we studied in Chiapas, Mexico. Males and females of D. urban are superficially very similar to the unspotted male form of the new species (Figs. 5–6), but actually belong to an entirely different group within Diomus (Diomus Group G, sensu Gordon 1999) characterized by the male genitalia with extremely wide and densely setose, funnel-shaped parameres, and the penis with an apical flagellum. Females belonging to this group are distinguished by a heavily sclerotized, cylindrical or spindle-shaped nodulus and elongate tapered cornu of the spermathecal capsule. Externally, adults of D. urban can be separated from the new species by their slightly more robust form and the longer intercoxal process of the prosternum (Fig. 10) with the carinae more narrowly separated posteriorly and more weakly convergent anteriorly. Description of holotype (adult, male) (Fig. 4). Length 2.0 mm, width 1.5 mm. Form broadly oval, convex (height = 0.9 mm). Metathoracic wing present. Dorsal surfaces finely granulate, feebly shining, distinctly punctate, pubescent. Coloration of head including mouthparts pale yellow with mandibular apex reddish amber; pronotum pale yellow with large irregularly rounded basomedian reddish brown spot. Elytron predominantly black with faint bluish sheen, with diffusely lighter yellow brown elytral apex; disc bearing large teardrop shaped oblique reddish orange mark; elytral epipleuron yellow brown with margins narrowly dark brown. Scutellar shield dark brown. Venter predominantly brown, darkest on meso-, metaventrite; prosternum pale yellow with median half yellow brown; mesepimeron paler than surrounding sclerites with margins narrowly darkened; abdomen pale brown, lighter, yellower toward apex. Legs pale yellow with meso-, metacoxae slightly darker. Pubescence shiny off white. Dorsal punctation moderately coarse; punctures separated by approximately 1× diameter on elytron, equal in width to eye facet; punctures slightly coarser, more crowded on head, pronotum. Dorsal pubescence semierect, moderately dense, evenly distributed, lacking distinct setal pattern; individual setae equal to about 1/3 to 3/4× length of scutellar shield, arcuate; elytral setae directed more or less posteriorly except directed outwardly at sides of body, posteromedially near suture. Head large, 0.64× width of pronotum; eye large, finely facetted, with interfacetal setae, distinctly notched near antennal insertion by small triangular ocular canthus; inner orbits evenly arcuate, with minimum separation near midlength; interocular distance 1.6× width of eye in frontal view. Antenna composed of 11 antennomeres; antennomere 3 elongate, about 2.3× length of antennomere 4; last four antennomeres subequal in length, forming gradual oval club expanding apically from antennomere 8–10; antennomere 11 subrhomboidal, slightly narrower, than 10. Maxillary palp with terminal palpomere strongly expanded distally, triangulate, about as long as greatest width. Pronotum with basal width about 2× length, tapered toward apex, strongly convex. Elytron in dorsal view with lateral margin evenly arcuate; elytral apices dehiscent. Ventral surfaces pubescent, distinctly punctate; punctation less even, generally less dense than elytral punctation. Prosternum (Fig. 9) with intercoxal process short, broad, with convergent carinae extending to, merging near apex, framing ovotriangular depression. Suture between abdominal ventrites 1 and 2 distinct, linear; 5th ventrite with apex shallowly, roundly emarginate; 6th ventrite with apex bearing indistinct shallow notch at middle. Tarsal claw with broad scythe-like inner tooth extending from base to apical 1/3 or beyond. Male genitalia (Figs 11–12). Basal lobe (=penis guide sensu Ślipiński 2007) moderately flat, in ventral view expanding from base to apical 2/5, tapered beyond to pointed apex, asymmetrical, about 2/3 length of paramere. Paramere with apical 1/2 roundly expanded, with about 19–25 long setae along perimeter, with few shorter setae in more proximal position on each side; setae slightly flattened, broad at base, sharply pointed distally; inner (=ventral) surface of paramere with very short setae sparsely distributed. Penis (=aedeagus or sipho) with outer arm of capsule short, truncate, bearing semicircular crest on outer surface; inner arm short, roundly tapered; penis apex lacking flagellum, with slightly expanded membranous area. Female (Fig. 7). Similar to male except body form proportionally shorter, dorsal coloration uniformly medium brown. Prosternum and underside of head yellow brown. 5th abdominal ventrite with posterior margin roughly linear in median 1/3; 6th ventrite with posterior margin arcuate. Tarsal claw with short triangular tooth near base, not extending beyond apical 1/2. Spermathecal capsule of female genitalia (Fig. 13) moderately small, weakly bent near middle; nodulus (=collum) in the form of short annular projection; ramus sessile, with thorn-like apodeme (=beak sensu Gordon 1999) above accessory gland; cornu short, distinctly swollen distally. Sperm duct lacking sclerotized sheath or process (=infundibulum auctorum). Bursa with lightly sclerotized internal chamber (=bursal plate sensu Gordon 1999) visible as light brown ring-like structure. Variation (Figs 4–7). Length 1.7–2.1 mm. Male with basal mark on pronotum variable in size; reddish orange oblique mark on elytron may be absent or only faintly indicted as slightly lighter area on disc; pale band at elytral apex yellow to brown or indistinct. Female with dorsal coloration light brown to blackish, unicolorous or with anterior pronotal angles and extreme elytral apex defusely lighter. Larva (based on field collected 2 nd –4 th instars) (Fig. 2, 14-15). Form broadly oval, onisciform. Body light grayish tan with head, thoracic and terminal abdominal plates medium brown. Anterior margin of head, posterior margin of abdominal apex with few simple setae. Dorsum devoid of setae, with granulate texture consisting of tiny fingerprint-like ridges and pits (visible at 50× magnification or greater), with eight conspicuous pairs of intersegmental pores near anterior borders of abdominal segments 1–8; first pair situated more anteriorly, appearing to extend onto base of metanotum. Head with frontal arms of epicranial suture well developed; epicranial stem obsolete. Pronotum with pair of pigmented plates; external pronotal margins expanded, flattened, surrounding sides of head. Meso-, metanotum similarly expanded, each with two pairs of small pigmented plates. Abdominal segments transverse, with expanded, flattened epipleurum; tergum 9 with v-shaped pigmented plate. Venter concave, without strongly sclerotized plates; legs short, robust, not capable of extending beyond sides of body; pygopod well developed. Pupa (Fig. 3). Form broadly oval, attached at caudal end, not enclosed in shed larval skin; surface bearing glandular hairs. Coloration yellow to yellowish brown. Trophic relations. Larvae and adults have been observed feeding on Coccus viridis (Green) on coffee plants, especially those tended by Azteca sericeasur Longino and Pheidole synanthropica Longino ants. Etymology. The species name is a concatenation of the Latin phrase “lupus apud oves” (=wolf among the sheep), in reference to its presence among coccids tended by ants. Type material. Holotype (male) with labels: “MEX: Chiapas: Finca Irlanda, 15.173583°−92.336081°, 28.July.2010, coll. H. Liere / On Coffea arabica plants among C. viridis, newly eclosed adult / HOLOTYPE, Diomus lupusapudoves Vandenberg, Iverson & Liere, 2018 ” (USNM); 19 paratypes (9 females, 10 males): 2 with same labels as type except third label “PARATYPE, Diomus lupusapudoves Vandenberg, Iverson & Liere, 2018 ” (USNM); 3 with labels “MEX: Chiapas: Finca Irlanda, 15.173583° −92.336081°, 24.May.2010, coll. A. Iverson / On Coffea arabica plants / PARATYPE, Diomus lupusapudoves Vandenberg, Iverson & Liere, 2018 ” (USNM); 14 with labels “MEX: Chiapas: Finca Irlanda, 15.173583° −92.336081°, 10.June.2010, coll. A. Iverson / Reared from late instar onisciform larva collected on Coffea arabica plants / PARATYPE, Diomus lupusapudoves Vandenberg, Iverson & Liere, 2018 ” (13 USNM, 6 ECO-TAP-E) Other material examined. A small amount of additional material of the new species bearing the same collection data as the 24.May.2010 and 10.June.2010 paratypes has been deposited in the USNM. These represent less well preserved exemplars (either disarticulated, fragmented, or rubbed), but still deemed useful for studying certain morphological or anatomical details. Specimens from each collection event have been placed together on a single pin using multiple points and/or within a gelatin capsule. A vial of mixed second through fourth instar larvae of the new species from the 10.June.2010 collection event has been deposited in the USNM alcohol collection. A male and a female specimen of D. urban from the 24.May.2010 collection event have been point-mounted and deposited in the USNM dry collection. The later collection also contains a large quantity of material of other South American Diomini documented in the Gordon (1999) monograph of this group and utilized for comparative purposes during our study. Remarks. Gordon (1999) informally divided the genus Diomus into eight species groups (Diomus groups A through H) based almost entirely on male genitalic characteristics. He indicated that some of these groups may be paraphyletic, but still have a value for identification purposes. Using his system, our new species belongs in group B and appears to be closely allied to D. thoracicus, the type species of the genus. Our new species exhibits strong sexual dimorphism with respect to the dorsal color pattern (Figs. 4–7). This situation is not uncommon in the tribe Diomini, and probably explains why the females of many other species have not been associated with their male counterparts. We have managed to include females in our type series because both sexes were reared together from the same distinctive onisciform larvae, and have the same configuration of the antenna, prosternal process, punctation, and pubescence (dorsal setal pattern).

Vandenberg, Natalia J., Iverson, Aaron, Liere, Heidi (2018): A new species of myrmecophilous lady beetle in the genus Diomus (Coleoptera: Coccinellidae: Diomini) from Chiapas, Mexico that feeds on green coffee scale, Coccus viridis (Green) (Hemiptera: Coccidae). Zootaxa 4420 (1): 113-122, DOI: https://doi.org/10.11646/zootaxa.4420.1.6

Diomus Mulsant Scymnus (Diomus) Mulsant 1850: 951. Type species: Coccinella thoracica Fabricius, 1801, by subsequent designation of Korschefsky 1931. Diomus: Weise 1895:144. Nephus (Diomus): Iablokoff-Khnzorian 1976: 377. Amidellus Weise 1923: 141. Type species: Scymnus ementitor Blackburn, 1895 by original designation. Synonymized by Ślipiński 2007: 87 (see Gordon (1976) and Pang & Gordon (1986) for a more complete bibliography). Diomus is the most speciose genus in the tribe Diomini Gordon, 1999, and possibly the largest genus in the entire family Coccinellidae Latreille, 1807 (Pang & Ślipiński 2009, 2010). Diomus was originally placed as a subgenus of Scymnus Kugellan, 1794, and classified, until recent times, in the tribe Scymnini Mulsant, 1846 along with a miscellaneous assortment of other small pubescent lady beetles. Gordon (1999) recognized Diomus and allies as deserving of their own tribe Diomini, distinct from Scymnini, but unfortunately his circumscription of the former included some genera belonging to another tribe���Selvadiini Gordon, 1985 (Vandenberg 2002; Vandenberg & Hanson in review). A re-circumscription and review of Diomini is detailed in Vandenberg and Hanson (in review) and will not be repeated here., Published as part of Vandenberg, Natalia J., Iverson, Aaron & Liere, Heidi, 2018, A new species of myrmecophilous lady beetle in the genus Diomus (Coleoptera: Coccinellidae: Diomini) from Chiapas, Mexico that feeds on green coffee scale, Coccus viridis (Green) (Hemiptera: Coccidae), pp. 113-122 in Zootaxa 4420 (1) on page 114, DOI: 10.11646/zootaxa.4420.1.6, http://zenodo.org/record/1247206, {"references":["Slipinski, A. (2007) Australian Ladybird Beetles (Coleoptera: Coccinellidae): Their Biology and Classification. Australian Biological Resources Study, Canberra, ACT, 288 pp.","Gordon, R. D. (1976) The Scymnini of the United States and Canada: Key to genera and revision of Scymnus, Nephus and Diomus. Bulletin of the Buffalo Society of Natural Sciences, 28, 1 - 362.","Pang, X. F. & Gordon, R. D. (1986) The Scymnini (Coleoptera: Coccinellidae) of China. The Coleopterists Bulletin, 40, 157 - 199.","Gordon, R. (1999) South American Coccinellidae (Coleoptera), Part VI: A systematic revision of the South American Diomini, new tribe (Scymninae). Annales Zoologici, Warsaw, 49, 1 - 219.","Pang, H. & Slipinski, A. (2009) Revision of the Australian Coccinellidae (Coleoptera). Genus Diomus Mulsant. Part 1. Annales Zoologici, 59, 641 - 698. https: // doi. org / 10.3161 / 000345409 x 485008","Pang, H. & Slipinski, A. (2010) Revision of the Australian Coccinellidae (Coleoptera). Genus Diomus Mulsant. Part 2. Annales Zoologici, 60, 493 - 545. https: // doi. org / 10.3161 / 000345410 X 550382","Vandenberg, N. J. (2002) Coccinellidae Latreille, 1807. In: Arnett, R. & Thomas, M. (Eds.), American Beetles. Vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton, FL, pp. 371 - 389."]}

A new species of myrmecophilous lady beetle, Diomus lupusapudoves, sp. nov. (Coleoptera: Coccinellidae: Diomini), is described from a coffee agroecosystem in Chiapas, Mexico. The new species was found preying on the green coffee scale pest, Coccus viridis (Green), tended primarily by Azteca sericeasur Longino and Pheidole synanthropica Longino ants. The larval, pupal, and adult stages of the new species are described and habitus illustrations or photos provided along with anatomical details of the adult male and female genitalia. The species is most similar to Diomus thoracicus Fabricius (=type species of Diomus), another myrmecophile, which inhabits ant nests and feeds on ant brood. The new species has a peculiar onisciform larva that lacks dorsal setae, features that it shares with D. thoracicus. The new species is only the second species in the genus reported as a myrmecophile, although the life histories of most species have been poorly documented.

Anisandrus maiche Stark, an ambrosia beetle native to Asia, is reported for the first time in North America based on specimens from Pennsylvania, Ohio, and West Virginia. This is the twentieth species of exotic Xyleborina documented in North America. This species, along with three others occurring in North America, were formerly placed in Xyleborus Eichhoff, but currently are assigned to Anisandrus Ferrari. Descriptions of generic characters used to separate Anisandrus from Xyleborus, a re-description of the female A. maiche, and an illustrated key to the four North American species of Anisandrus are presented.

The Hispaniolan genus Bura Mulsant is removed from Coccidulinae and placed in Sticholotidinae. The characteristics which justify this transfer are discussed and an historical review of the classification of the Sticholotidinae is presented. Bura is diagnosed and redescribed, and its affinities to other Sticholotidini are discussed. Illustrations of key generic characters are provided.

1. Phaenochilus flaviceps Miyatake 1970 Phaenochilus flaviceps Miyatake 1970b: 111 (original description; morphological details and male genitalia figured). Type material: Holotype, male (BPBM); paratype, female (BPBM). Type locality: “ Coimbatore, Madras State, South India.” Remarks: This is one of three known Phaenochilus species with black to piceous elytra, and the only one of that coloration known to occur in India. It differs from P. ruficollis in having the median part of the pronotum black instead of red, and from P. mikado in having the basal lobe of the male genitalia (Fig. 9) with a sharply pointed median apical projection, and the parameres spoonshaped. It resembles Chilocorus nigritus (Fabricius), but differs in possessing the slender terminal article of the maxillary and labial palpi, narrow frons, and elongate basal tooth of the tarsal claw., Published as part of Giorgi, José Adriano & Vandenberg, Natalia J., 2012, Review of the lady beetle genus Phaenochilus Weise (Coleoptera: Coccinellidae: Chilocorini) with description of a new species from Thailand that preys on cycad aulacaspis scale, Aulacaspis yasumatsui Takagi (Hemiptera: Sternorrhyncha:, pp. 239-255 in Zootaxa 3478 on page 248, {"references":["Miyatake, M. (1970 b) Two new species of the genus Phaenochilus Weise from South India (Coleoptera: Coccinellidae). Transactions of the Shikoku Entomological Society, 10, 111 - 115."]}

3. Phaenochilus kashaya Giorgi & Vandenberg, new species Phaenochilus kashaya Giorgi & Vandenberg, current article (original description; adult habitus, morphological details, and male genitalia figured). Phaenochilus punctifrons: Chunram & Sasaji 1980: 481 (provisional ID in a checklist of the Coccinellidae of Thailand) (misidentification); Chunram 2002: 112 (habitus photo) (misidentification), not Weise 1895. Type material: Holotype, male (USNM); paratypes (14 total), 8 males, 6 females (2, USNM; 2, AMNH; 2, CASC; 2, TDOA; 2, ECMP; 2, EUCJ; 2, IOZB). Type locality: THAILAND: Nakhon, Ratchasima, Sub Tao, N 14°29.45’ E 101°58.60’ Remarks: The broad oval parameres of the male genitalia (Fig. 13) will distinguish this species from congeneric species with a similar yellow to reddish orange or ferrugineus coloration. Phaenochilus kashaya further differs from P. metasternalis in possessing narrow, weakly explanate elytral margins (Figs 5, 27, 32–34), and from P. punctifrons in possessing a more expansive pronotum with the anterior pronotal margins on each side of the emargination subtruncate (Figs 28, 31). Chunram and Sasaji (1980) tentatively assigned a specimen from Chiang Mai Province to P. punctifrons, and subsequently published a photographic image of a specimen from the same locality in a book on the lady beetles of Thailand (Chunram 2002), but we believe these records to be a misidentification of P. kashaya., Published as part of Giorgi, José Adriano & Vandenberg, Natalia J., 2012, Review of the lady beetle genus Phaenochilus Weise (Coleoptera: Coccinellidae: Chilocorini) with description of a new species from Thailand that preys on cycad aulacaspis scale, Aulacaspis yasumatsui Takagi (Hemiptera: Sternorrhyncha:, pp. 239-255 in Zootaxa 3478 on page 248, {"references":["Chunram, S. & Sasaji, H. (1980) A contribution to the Coccinellidae (Coleoptera) of Thailand. Oriental Insects, 14 (4), 473 - 491.","Chunram, S. (2002) Lady beetles in Thailand. Division of Entomology and Zoology. Department of Agriculture Science, 209 pp. (in Thai).","Weise, J. (1895) Neue Coccinelliden sowie Bemerkungen zu bekannten Arten. Annales de la Societe Entomologique de Belgique, 39, 120 - 146 (in German)."]}

Genus Phaenochilus Weise 1895 Phaenochilus Weise 1895: 135 (original description); Korschefsky 1932: 237 (catalogue); Chapin 1965a: 234–235, 267–269 (key to genera of Chilocorini; rediagnosis and historical review of Phaenochilus); Miyatake 1970a: 20, 49–51 (key to east Asian Chilocorini; rediagnosis and historical review of Phaenochilus); Poorani 2002: 7 (checklist, Coccinellidae of the Indian subcontinent). Type species: Phaenochilus punctifrons Weise, by subsequent designation of Korschefsky 1932. Diagnosis: Members of the genus share a similar body form (Figs 1–7) which is compact and roughly hemispherical with the elytral margin narrowly to broadly explanate. Phaenochilus can be easily distinguished from other Chilocorini genera by the combination of: antennae with 8 antennomeres (Figs 18–19), eyes large and elongate (Fig. 25–27, 33), frons narrow and distally tapered, interocular distance less than to 1.5X width of an eye, labrum ovotriangulate, mandible weakly curved with outer edge nearly linear, terminal maxillary palpomere (Figs 14–17) elongate, slender, with length equal to or greater than twice basal width, subcylindrical to tapered with oblique apex, or somewhat swollen with subtruncate apex, terminal labial palpomere (Fig. 20) elongate, slender, rodlike, elytral epipleuron deeply foveate for reception of femoral apices, tibial spurs lacking, tarsal claws (Fig. 21) with elongate basal tooth half to more than two thirds length of claw, male genitalia (known for 6 of 9 species) (Figs 8–13) with basal lobe weakly to distinctly asymmetrical, parameres unequal, spermatheca of female genitalia (known for 3 of 9 species) stout with an elongate appendix or apodeme on distal end of cornu., Published as part of Giorgi, José Adriano & Vandenberg, Natalia J., 2012, Review of the lady beetle genus Phaenochilus Weise (Coleoptera: Coccinellidae: Chilocorini) with description of a new species from Thailand that preys on cycad aulacaspis scale, Aulacaspis yasumatsui Takagi (Hemiptera: Sternorrhyncha:, pp. 239-255 in Zootaxa 3478 on page 242, {"references":["Weise, J. (1895) Neue Coccinelliden sowie Bemerkungen zu bekannten Arten. Annales de la Societe Entomologique de Belgique, 39, 120 - 146 (in German).","Korschefsky, R. (1932) Pars 120: Coccinellidae. II. In: Junk, W. & Schenkling, S. (Eds), Coleopterorum Catalogus. W. Junk, Berlin, pp. 225 - 659.","Chapin, E. A. (1965 a) New species of Chilocorini (Coleoptera: Coccinellidae). Psyche, 72 (2), 148 - 151.","Miyatake, M. (1970 a) The East - Asian coccinellid beetles preserved in the California Academy of Sciences. Tribe Chilocorini. Memoirs of the College of Agriculture, Ehime University, 14 (3), 19 - 56.","Poorani, J. (2002) An annotated checklist of the Coccinellidae (Coleoptera) (excluding Epilachninae) of the Indian subregion. Oriental Insects, 36, 307 - 383."]}

4. Phaenochilus metasternalis Miyatake 1970 Phaenochilus metasternalis Miyatake 1970a: 50–52 (original description; male and female genitalia figured); Hoang 1981: 12, 15, 16 (first report from Vietnam); Ren Shunxiang et al. 2009: 138–139 (photos of dorsal habitus, male genitalia, abdominal postcoxal line). Phaenochilus punctifrons: Chapin 1965b: 267 & Fig. 18A–G (female genitalia and morphological details figured) (misidentification). Type material: Holotype, male (CASC); paratypes (2 total), 1 male, 1 female (CASC). Type locality: “ Ta Hau, Hainan Is., S. China.” Remarks: This species can be distinguished from similarly colored congenerics by the broadly explanate elytral margins (Figs 3, 26), the central part of the metasternum densely covered with long hairlike setae (may be lost through abrasion), and the male genitalia which has the basal lobe distinctly undulate on one side and constricted near base (Fig. 12a). A small black leather notebook associated with Chapin’s slide collection (USNM) documents that he performed a whole body dissection of a female of “ P. punctifrons ” from Java (Java, Feb. 1911, R.L.Woglum). Although we could not find the resulting slide mounted material which he designated “X-15,” the remaining specimens from Java, Buitenzorg (=Bogor), and Singapore in the series that Chapin studied all belong to P. metasternalis. Ren Shunxiang et al. (2009) report the presence of P. metasternalis in the Chinese provinces of Yunnan, Anhui, Guangdong, Guangxi, and Hainan. Hoang (1981) reports this species from Trung Quoc, Viet Nam. (See also “Remarks” section for P. ruficollis, below.), Published as part of Giorgi, José Adriano & Vandenberg, Natalia J., 2012, Review of the lady beetle genus Phaenochilus Weise (Coleoptera: Coccinellidae: Chilocorini) with description of a new species from Thailand that preys on cycad aulacaspis scale, Aulacaspis yasumatsui Takagi (Hemiptera: Sternorrhyncha:, pp. 239-255 in Zootaxa 3478 on page 250, {"references":["Miyatake, M. (1970 a) The East - Asian coccinellid beetles preserved in the California Academy of Sciences. Tribe Chilocorini. Memoirs of the College of Agriculture, Ehime University, 14 (3), 19 - 56.","Hoang, D. N. (1981) New data on the subfamily Chilocorinae (Coccinellidae, Coleoptera) in Vietnam. Tap chi Sinh Vat Hoc, 3 (1), 11 - 17 (in Vietnamese).","Chapin, E. A. (1965 b) The genera of the Chilocorini (Coleoptera, Coccinellidae). Bulletin of the Museum of Comparative Zoology, 133 (4), 227 - 271."]}

7. Phaenochilus punctifrons Weise 1895 Phaenochilus punctifrons Weise 1895: 136 (original description); Chapin 1965b: 267, in part (illustration of male genitalia). Type material: Syntype, female (MNHB) (digital images of dorsal habitus and SEMs of morphological details taken by P. Laczynski (MIZW) examined). Type locality: “Insula Banguey.” Remarks: Both P. metasternalis and P. kashaya have been confused with this species in the past (see the corresponding remarks sections, above). Phaenochilus punctifrons can be distinguished from both of these other species by the less expansive pronotum with the lateral part on each side of the head only weakly prolonged and tightly, evenly rounded apically (Figs 25, 28, 29), by the roughly semicircular shape of the elytral dorsum in a frontal view (Fig. 25), and by the nearly symmetrical basal lobe of the male genitalia (Fig. 11a). The male genitalia of P. punctifrons were illustrated by Chapin (1965b) based on an individual from Mindanao Island which he recorded as “Mindanao, Kolambugan, C.F.Baker #16609” in his little black notebook, and designated as “X-14” (see also remarks under P. metasternalis, above). This slide could not be located in Chapin’s slide cabinet (USNM), but a second specimen from Mindanao Island labeled “Surigao, Mindanao, Baker” was dissected and found to have matching male genitalia except for the shape of the sipho. When we transferred our new dissection between media of different viscosities, the sipho temporarily assumed an S-shape matching Chapin’s drawing, apparently due to a difference in the hydrostatic pressure on the outside and inside of the siphonal tube. When allowed to equilibrate, the structure resumes the simple curved shape seen in our figure (Fig. 11c). (See also “Remarks” section for P. monostigma, above.), Published as part of Giorgi, José Adriano & Vandenberg, Natalia J., 2012, Review of the lady beetle genus Phaenochilus Weise (Coleoptera: Coccinellidae: Chilocorini) with description of a new species from Thailand that preys on cycad aulacaspis scale, Aulacaspis yasumatsui Takagi (Hemiptera: Sternorrhyncha:, pp. 239-255 in Zootaxa 3478 on pages 251-252, {"references":["Weise, J. (1895) Neue Coccinelliden sowie Bemerkungen zu bekannten Arten. Annales de la Societe Entomologique de Belgique, 39, 120 - 146 (in German).","Chapin, E. A. (1965 b) The genera of the Chilocorini (Coleoptera, Coccinellidae). Bulletin of the Museum of Comparative Zoology, 133 (4), 227 - 271."]}

6. Phaenochilus monostigma Weise 1895 Phaenochilus monostigma Weise 1895: 241 (original description); Schultze 1916:38 (catalogue of Philippine Coleoptera). Type material: Holotype, sex unknown (? ECMP). Type locality: “ Agusan River, Mindanao.” Remarks: This is the only species of Phaenochilus with a single common dark sutural spot on the elytra (Fig. 7). The male genitalia are unknown. Weise indicated that the type of this species was from the collection of W. Schultze and was deposited in the collection of the Bureau of Science, Manila. The USNM has a specimen that was acquired with the Korschefsky collection bearing the labels “Mindanao, P. I. / Phaenochilus monostigma Ws. Det W. Schultze ” that may belong to the same series as the type. The specimen is disarticulated and mounted on two points. Recuenco-Adorada (2008) contributed a paper on Philippine Chilocorini, but apparently misidentified or confused some of the species. The included redescription and habitus illustration of P. monostigma portray a specimen with an immaculate elytral disc and narrow black lateral border on the basal two thirds of each elytron. The male genitalia (lateral views of the sipho and phallobase) and anatomical details incorporated on the same plate suggest instead the species P. punctifrons, although we have not seen a comparable example with the dark elytral border. In the same publication, another plate labeled Eguis sp. shows a specimen which matches the USNM example of P. monostigma in such details as the dorsal color pattern and the shape of the maxillary palpomeres, antenna, pronotum, prosternum, abdominal postcoxal line, and tarsal claw., Published as part of Giorgi, José Adriano & Vandenberg, Natalia J., 2012, Review of the lady beetle genus Phaenochilus Weise (Coleoptera: Coccinellidae: Chilocorini) with description of a new species from Thailand that preys on cycad aulacaspis scale, Aulacaspis yasumatsui Takagi (Hemiptera: Sternorrhyncha:, pp. 239-255 in Zootaxa 3478 on page 251, {"references":["Weise, J. (1895) Neue Coccinelliden sowie Bemerkungen zu bekannten Arten. Annales de la Societe Entomologique de Belgique, 39, 120 - 146 (in German).","Schultze, W. C. M. (1916) A catalogue of Philippine Coleoptera. Philippine Journal of Science. Section D, General Biology, Ethnology, and Anthropology, 11 (1 - 2), 194.","Recuenco-Adorada, J. D. (2008) Systematics of Philippine Chilocorinae. I. Tribe Chilocorini (Coleoptera: Coccinellidae). Philippine entomologist, 22 (2), 147 - 162."]}

2. Phaenochilus indicus Miyatake 1970 Phaenochilus indicus Miyatake 1970b: 113 (original description; morphological details and male genitalia figured). Type material: Holotype, male (EUCJ). Type locality: “ Chinchona, Anamalai Hills, 1,050 m (3,500 ft.), Madras State, South India.” Remarks: Three other congeners possess immaculate yellow to reddish orange or ferrugineus elytra, but this is the only species of that appearance known to occur in India. It also differs from the similarly colored species in possessing a more compact antennal club with the last antennomere not much longer than wide (Fig. 19). The male genitalia (Fig. 10) and morphological details illustrated by Miyatake (1970b) suggest a close relationship to the melanic P. flaviceps (Fig. 9), also described from Madras State., Published as part of Giorgi, José Adriano & Vandenberg, Natalia J., 2012, Review of the lady beetle genus Phaenochilus Weise (Coleoptera: Coccinellidae: Chilocorini) with description of a new species from Thailand that preys on cycad aulacaspis scale, Aulacaspis yasumatsui Takagi (Hemiptera: Sternorrhyncha:, pp. 239-255 in Zootaxa 3478 on page 248, {"references":["Miyatake, M. (1970 b) Two new species of the genus Phaenochilus Weise from South India (Coleoptera: Coccinellidae). Transactions of the Shikoku Entomological Society, 10, 111 - 115."]}

Sticholotis gibbula Weise, stat. rev. (Figs. 8���10) Sticholotis (Apterolotis) gibbula Weise, 1908: 226 (Lectotype female, BMNH). Stictobura gibbula: Sicard, 1911: 385.��� Korschefsky, 1931: 211. The syntype female of this species (BMNH, examined) indeed belongs to Sticholotis (stat. rev.) and is very small, unlike other members of Stictobura. It is not clear why Sicard (1911) and subsequently Korschefsky (1931) chose to place it under Stictobura. The syntype from BMNH is designated here as lectotype (lectotype designation) and briefly redescribed below. Redescription. Length: 1.60 mm; width: 1 mm. TL/TW: 1.21; EL/EW: 0.96; PL/PW: 0.46. Form short oval, dorsum strongly convex (Fig. 8) and dumpy, with a prominent, hump-backed outline (Fig. 10). Dorsum and ventral side yellow except scutellum, edges of elytra and epipleura darker reddish brown, elytra with an ill-defined stria marked by darkly pigmented dots on either side of sutural line, antenna with flagellomeres darker brownish in posterior half. Head (Fig. 9) with clypeal margin shallowly emarginate, pubescence silvery white. Eyes small, coarsely faceted, posteriorly strongly divergent, with an orbital carina, interocular distance ca. 3.2 x as wide as eye; punctures separated by 2���5 diameters, interspaces with strong reticulate microsculpture. Antennal insertions exposed, frons emarginate around antennal insertions. Antenna 11 -segmented, with a distinct club. Pronotum transverse; basal margin strongly arcuate, with submarginal line; lateral margins nearly linear, with short scattered hairs; anterior margin deeply trapezoidally emarginate around head; lateral and anterior margins with fine bead; punctures on pronotum smaller than those on head, shallow, widely separated, interspaces with reticulate microsculpture. Elytra with punctures larger, closer and denser than those on head and pronotum, separated by 2���4 diameters, interspaces smooth, shiny. Prosternal process obscured by forelegs. Abdomen with five visible ventrites; abdominal postcoxal line incomplete, curved posterolaterally, joining posterior margin of ventrite 1; ventrites 2���4 subequal; ventrite 5 elongate, tapering, posteriorly arcuate. Legs with cryptotetramerous tarsi; tarsal claws narrow, sickle-shaped, basally slightly swollen, lacking a distinct basal tooth. Genitalia not dissected. Specimen examined: Lectotype of gibbula, female (designated here) : ���Type (circular Red bordered Label)/ Madura, S. India / Sticholotis gibbula m (in Weise���s handwriting)/Type (rectangular orange label)/ Madura��� (BMNH). Distribution. India: Tamil Nadu. Note. Weise (1908) did not discuss variation within the species, nor indicate the number of specimens examined., Published as part of Vandenberg, Natalia J., 2011, A revision of the genus Stictobura Crotch and description of a new species of Sticholotis Crotch (Coleoptera: Coccinellidae: Sticholotidinae), pp. 1-13 in Zootaxa 3031 on pages 8-9, DOI: 10.5281/zenodo.202599, {"references":["Weise, J. (1908) Coleopteren aus Ostindien. Stettiner Entomologische Zeitung, 69, 213 - 230.","Sicard, A. (1911) Especes et varietes nouvelles de Coccinellides de l'Inde [Coleopt.] appartenant a la collection de M. Andrewes (de Londres). Annales de la Societe Entomologique de France, 79 [1910], 377 - 389.","Korschefsky, R. (1931) Coleopterorum Catalogus. Pars 118. Coccinellidae I. Berlin, 224 p."]}

1. melanaria (Weise) (Figs. 7, 28��� 32) Sticholotis melanaria Weise, 1903: 232 (Type depository:?ZMB; Type locality: Pondicherry). Stictobura melanaria: Korschefsky, 1931: 211. Diagnosis. This species can be identified by the fully black, strongly convex dorsum with dense, closely placed elytral punctures. The male genitalia (Figs. 28���31) are diagnostic. Redescription. Length: 3.1���3.6 mm; width: 3.0��� 3.5 mm; TL/EW: 1.03���1.06; EL/EW: 0.83���0.95; PL/PW: 0.40���0.44. Form (Fig. 7) more or less circular, dorsum strongly hemispherical and dome-like. Dorsal side fully black, ventral side dark pitchy brown to black; head with silvery white pubescence; pronotum and elytra with sparse but distinct, uniform, suberect to erect silvery white hairs, more noticeable on anterior, lateral and posterior margins of elytra, than on disk of elytra. Head with clypeal margin shallowly emarginate; eyes widely separated by more than 3 x eye width; punctures very shallowly impressed, widely separated by 3���6 diameters, interspaces between punctures strongly reticulate. Pronotum with lateral sides linear, antero- and posterolateral corners broadly rounded, posterior margin strongly sinuate with submarginal line, lateral sides narrowly beaded; punctures shallowly impressed, separated by 2���5 diameters. Scutellum small, triangular. Elytra slightly wider than long, with denser, more closely placed punctures, only slightly more deeply impressed than those on head and pronotum, separated by 1���4 diameters, more or less of one size though with some coarser punctures, interspaces more or less smooth. Prosternal process broad, quadrate with a pair of carinae, subparallel posteriorly, gradually divergent towards anterior. Tarsal claws almost simple, with a weak basal tooth. Elytral epipleura shallowly depressed on level with middle and hind legs. Abdominal postcoxal lines incomplete; ventrite 5 in female with a depressed area on either side below anterior margin, posterior margin broadly truncate; ventrite 5 in male apically truncate. Female genitalia with a long bursa and a prominent sclerotized structure (Fig. 32), probably a poorly differentiated spermatheca. Male genitalia (Figs. 28���31) with basal lobe of tegmen (Fig. 29) elongate cylindrical, apically rounded and produced into a short, blunt process; sipho (Fig. 30, 31) elongate, stout, with a large capsule. Specimens examined. INDIA: Tamil Nadu: Trichinopoly, Ind. Or., 6 females, 2 ex (MNHN); Kerala: without locality data, 1 female (NBAII); Shembaganur, Mad. 1904���1905, P. du Breuil / BMNH, 1 female, 3 ex (BMNH); S. India, Shambaganur, Madura, 1921 -146, 1 female (BMNH, dissected); Trichinopoli, Ind.-Or./ Nunenmacher Collection, 1 male, 1 female (MNHN); Shembaganur, Sud-India/ St. melanaria, 1 female (MNHN). Distribution. India: Pondicherry; Tamil Nadu. Note. Weise (1903) described Sticholotis melanaria from Staudinger material and it is expected that Weise might have kept some examples in Berlin (ZMB) as well as returning material to Staudinger. Efforts to find Staudinger Coccinellidae material have proved unsuccessful in the past., Published as part of Vandenberg, Natalia J., 2011, A revision of the genus Stictobura Crotch and description of a new species of Sticholotis Crotch (Coleoptera: Coccinellidae: Sticholotidinae), pp. 1-13 in Zootaxa 3031 on pages 3-4, DOI: 10.5281/zenodo.202599, {"references":["Weise, J. (1903) Neue Coccinelliden. Deutsche Entomologische Zeitschrift, 1903, 229 - 232.","Korschefsky, R. (1931) Coleopterorum Catalogus. Pars 118. Coccinellidae I. Berlin, 224 p."]}

Vandenberg, Natalia J. (2011): A revision of the genus Stictobura Crotch and description of a new species of Sticholotis Crotch (Coleoptera: Coccinellidae: Sticholotidinae). Zootaxa 3031: 1-13, DOI: 10.5281/zenodo.202599

2. pallideguttata (Mulsant) (Figs. 1, 19��� 23) Calvia? pallideguttata Mulsant, 1853: 289 (holotype, UCCC). Stictobura pallidiguttata: Crotch, 1874: 201 (name misspelt).��� Weise, 1923: 137.��� Korschefsky, 1931: 211.��� Gordon, 1987: 25 (unnecessary lectotype designation). Sticholotis (Apterolotis) andrewesi Weise, 1908: 225; 1923: 127 (Lectotype, BMNH).���Synonymised by Sicard, 1911: 385.- Korschefsky, 1931: 211. Diagnosis. It can be identified by the dark brown to dull black elytra with eight yellow spots (Fig. 1) and dual punctation. The male genitalia (Figs. 19���23) are diagnostic. Redescription. Length: 3.78 mm; width: 3.50 mm; TL/EW: 1.03���1.06; EL/EW: 0.93���0.94; PL/PW: 0.42��� 0.44. Form (Fig. 1) subcircular, broadest around middle; dorsum strongly convex. Head and pronotum yellowish brown to testaceous, elytra dull to dark brown, with four oblique, elongate oval yellowish spots on each elytron, one pair in anterior half and one in posterior; antenna yellowish to brown, occasionally pedicel and scape lighter, yellowish; legs sometimes darker brown. Ventral side yellowish brown to testaceous except elytral epipleura yellowish brown with darker brown borders or fully dark brown. Head with shallowly impressed punctures, separated by 1���3 diameters, interspaces with reticulate microsculpture; pubescence on head silvery white. Antenna 11 -segmented with a long club, terminal antennomere elongate oval. Pronotum with shallowly impressed punctures, more widely spaced than those on head, separated by 2���5 diameters. Elytra slightly wider than long, punctation distinctly dual, with a mixture of fine and somewhat irregular, larger and coarser punctures, widely separated, interspaces weakly alutaceous. Prosternal process broadly quadrate, carinae subparallel, anteriorly slightly divergent. Lateral margins of elytra with a marginal bead. Anterior margin of mesoventrite shallowly, broadly emarginate in middle. Elytral epipleura shallowly impressed on level with hind legs. Tarsal claws basally somewhat swollen, lacking a distinct basal tooth. Posterior margin of ventrite 5 broadly truncate in female, double emarginate in male. Male genitalia (Figs. 19���23) as illustrated, basal lobe slightly longer than parameres in lateral view (Fig. 19), slightly narrower at middle, apically narrowed and produced into a short tubular process; basal lobe in ventral view (Fig. 20) elongate cylindrical and subparallel, apically narrowed, triangular; sipho (Fig. 21) with a large capsule, apex (Figs. 22, 23) somewhat lanceolate. Specimens examined. Lectotype of Sticholotis andrewesi (designated here): ���Type (circular red bordered label)/ Nilgiri Hills, H.L. Andrewes/ Andrewes Bequest B.M. 1922 - 221 /Nilgiri Hills/ Sticholotis andrewesi m. (in Weise���s handwriting)/Type (rectangular orange label)/Syntype (blue label)���, abdomen missing (BMNH); Others: India: Tamil Nadu, Kotagiri, 23.x. 1975, C.A. Viraktamath (NBAII); Coonoor, 22.vi. 59, CIBC-BS, Ex. Red scale on mulberry, 1 female, 1 male (PDBC); Keti, June 1929, M.S.K. Coll., 1 female, 1 male (NBAII); Keti, 17.IV. 29, M.S.K.coll., 1 male; Nilgiris waterfalls, 1���6 May 15, Ponniah coll., 1 male (NBAII). Distribution. India: Kerala; Tamil Nadu. Notes. Mulsant (1853) observed that his type specimen was in poor condition. He felt it belonged in ���Halyziaires���, but assigned it to the genus Calvia with reservation. He referred clearly to ���L���individu���, which is therefore a holotype. The species of Stictobura are much larger than the average size of most sticholotidines, which might have led to some confusion regarding its placement. Crotch (1874) proposed the genus Stictobura, with Calvia pallideguttata Mulsant as the type species���he also misspelled the name of the type species as S. pallidiguttata. Sicard (1911) compared a specimen of Stictobura pallideguttata (labeled by Crotch) with the type specimen of Sticholotis (Apterolotis) andrewesi and pronounced them synonyms. Weise (1923) accepted S. andrewesi and S. pallideguttata Crotch (not Mulsant) as synonyms. He noted differences between Mulsant���s original description and Crotch���s redescription which led him to believe the Crotch material was misidentified. Korschefsky (1931) retained Mulsant���s species under the genus Calvia and Crotch���s species under Stictobura. Gordon (1985) designated one of the three specimens in Crotch���s collection, a ���severely damaged��� one, as the lectotype, unnecessarily so. The syntype of Sticholotis andrewesi (BMNH, examined), designated as lectotype (above), is similar to the nominate form of the species and matches well to Weise���s original species description (1908). Weise did not discuss variation within the species, nor indicate the number of specimens examined. Biology / associated habitat. Red scale, Aonidiella aurantii (Maskell) on mulberry; Coccus viridis (Green) on citrus; collected on tea (label data). Rao et al. (1970) reported it as feeding on tea mites., Published as part of Vandenberg, Natalia J., 2011, A revision of the genus Stictobura Crotch and description of a new species of Sticholotis Crotch (Coleoptera: Coccinellidae: Sticholotidinae), pp. 1-13 in Zootaxa 3031 on pages 4-5, DOI: 10.5281/zenodo.202599, {"references":["Mulsant, E. (1853) Supplement a la monographie des coleopteres trimeres securipalpes. Annales de la Societe Linneenne de Lyon (N. S.), 1, 129 - 333. [reprinted as Opuscules Entomologiques, 3, 1 - 205.]","Crotch, G. R. (1874) A Revision of the Coleopterous Family Coccinellidae, E. W. Janson, London. 311 p.","Weise, J. (1923) Results of Dr E. Mjoberg's Swedish scientific expeditions to Australia, 1910 - 1913. 31. Chrysomeliden und Coccinelliden aus Queensland. Arkiv fur Zoologi, 15 (12), 1 - 150.","Korschefsky, R. (1931) Coleopterorum Catalogus. Pars 118. Coccinellidae I. Berlin, 224 p.","Gordon, R. D. (1987) A catalogue of the Crotch collection of Coccinellidae (Coleoptera). Occasional Papers on Systematic Entomology, 3, 1 - 46.","Weise, J. (1908) Coleopteren aus Ostindien. Stettiner Entomologische Zeitung, 69, 213 - 230.","Sicard, A. (1911) Especes et varietes nouvelles de Coccinellides de l'Inde [Coleopt.] appartenant a la collection de M. Andrewes (de Londres). Annales de la Societe Entomologique de France, 79 [1910], 377 - 389.","Rao, V. P., Datta, B. & Ramaseshaiah, G. (1970) Natural enemy complex of flushworm and phytophagous mites on tea in India. Scientific Publication Series No. 5, Tea Board, India. 53 p."]}

Delphastus hirtulus (Kirsch), new combination Figs. 2���5, 9 Alexia hirtula Kirsch, 1876: 132. (in part) Rhymbus hirtula (Kirsch): Csiki, 1901: 42. (in part) Rhymbus hirtulus (Kirsch): Csiki, 1910: 53. (in part) Bystus hirtulus (Kirsch): Strohecker, 1953: 22. (in part) Type material. Lectotype (here designated to ensure nomenclatural stability): ���Poznzn [sic, apparent misspelling of Pozuzu, a river in Peru] Coll Kirsch [green label] / Typus [red label] / Staatl. Museum f��r Tierkunde, Dresden / Lectotype Alexia hirtula Kirsch, 1876, des. F.W. Shockley & N.J. Vandenberg, 2010 [red label]��� (male). Paralectotypes: 3, all with identical labels, ���Pozuzu Coll. Kirsch [green labels] / Staatl. Museum f��r Tierkunde, Dresden / Paralectotype Alexia hirtula Kirsch, 1876 [yellow labels].��� Additional labels have been added to differentiate among the 3 paralectotypes: 1, a disarticulated, cleared specimen, returned in genitalia vial on original pin, ���# 1 / Delphastus hirtulus (Kirsch), 1876, det. F.W. Shockley & N.J. Vandenberg, 2010 ��� (female); 1, an intact point-mounted specimen, ���# 2 / Microscymnus n.sp., det. F.W. Shockley & N.J. Vandenberg, 2010 ��� (male); and 1, a partial point-mounted specimen lacking head, thorax and one elytron, ���# 3 / Scotocryptini (Leiodidae), probably Aglyptinus sp., det. F.W. Shockley & N.J. Vandenberg, 2010 ��� (sex not determined) (SMTD). Diagnosis. Delphastus hirtulus can be distinguished from its congeners by the combination of a short clypeus (Fig. 2) and distinctly punctate elytra bearing long erect hairlike setae sparsely distributed over the dorsal surfaces (Fig. 9), and the presence of a dense patch of short, decumbent setae on the lateral margin of the elytra beginning just above the epipleural fovea for reception of the metafemoral apex (Fig. 4). Delphastus anthracinus Gordon resembles this species in dorsal color pattern, vestiture, and general body form, but can be easily distinguished by the elongate clypeus which lends a nearly triangular shape to the head (Fig. 1). Delphastus hirtulus can be distinguished from other ���micrococcinellidae��� of the New World by the traits mentioned above together with the generic characteristics of an expanded prosternum concealing the mouthparts, angulate tibiae, trimerous tarsi, and antennal club composed of a single elongate segment. Redescription. Lectotype (male). Length 1.5 mm, width 1.0 mm. Form ovoid, slightly elongate, broadest in basal half, tapered posteriorly (Figs. 3���5). Color on dorsal surfaces deep reddish brown, nearly black, paler reddish brown near anterior and lateral margins of pronotum; head orange-brown; venter dark reddish brown except lighter brown on abdominal ventrites II���V, especially near external margins; appendages yellow brown. Dorsal surfaces polished, shiny, distinctly punctate, with sparse pubescence consisting of long erect to suberect yellowish hairlike setae; dense patch of shorter decumbent hairlike setae near lateral margin of elytron in apical 2 / 3. Head (Fig. 2) transverse, oval, tapered toward clypeus, evenly convex in lateral view (Fig. 4), weakly opisthognathous in repose, with intermixed fine and coarse punctures mostly concentrated in band between eyes (Fig. 2), with few decumbent hairs present (specimen apparently abraded); clypeus short, shallowly arcuate, projecting beyond ventral cusp of emargination for antennal insertions by about 1 / 4 distance separating ventral cusps. Pronotum with unevenly scattered, intermixed fine and coarse punctures; punctures separated by less than one to several puncture diameters; with scattered decumbent to erect hairlike setae. Elytron with disc evenly punctate; punctures fine, separated by 3��� 5 times their diameter; some punctures bearing long, erect, hairlike seta �� or more length of lateral pronotal margin, estimated number of setae 65, of which only about a dozen remain due to abrasion (see setal map Fig 9 and description based on paralectotype, below), remaining punctures each bearing microseta scarcely projecting beyond rim of puncture; dense elongate pubescent patch of more than 20 short decumbent setae near lateral margin beginning just above epipleural fovea for reception of metafemoral apex, continued posteriorly to area above hind margin of 4 th abdominal ventrite (Fig. 4); lateral margin of elytron slightly undulate, with weakly raised lateral bead; epipleuron with depression for reception of mid-, hind femora. Prosternum convex with anterior margin arcuate, with few very fine scattered punctures. Meso-, metasternum obscured by glue and paper point. Abdomen with intercoxal process of ventrite I with few fine scattered punctures; ventrites II���IV finely rugostriate; ventrite V with narrow rugostriate band near base, remainder polished with moderately coarse scattered setiferous punctures separated by 3���5 times their diameter; each seta about �� to 1 / 3 length of segment along midline. Meso-, metatibiae with median cusplike angulation on outer margin of ventral face. Paralectotype # 1 (female). Similar to male except head, pronotum entirely dark reddish brown, nearly black. Dissection and clearing provided the following information not observable in the point-mounted, undissected lectotype. Elytron with approximately 65 punctures each bearing long erect hairlike seta �� or more length of lateral pronotal margin (count based on scattered remaining setae and setal bases of broken setae, Fig. 9), forming about 7 striae: 4 uniseriate discal striae, 2���3 somewhat confused marginal/submarginal striae; lateral patch of short decumbent setae not seen (apparently abraded). Mesosternum with shallow intermediate-sized punctures. Metasternum with coarse to intermediate-sized punctures separated by less than to 5 times their diameter; punctures more deeply impressed in median half of sclerite, nearly obsolete laterally. Details of female genitalia not clearly observed in dissection. Original Latin description (Kirsch 1876): ���Subhemisphaerica, nigra, capite prothoracisque lateribus saepe rufescentibus, hujus angulis posticis subrectis, scutello triangulari; elytris levissime parce punctatis, pilis longis erectis sparse obsitis pedibus flavis. Long. 1 ��, lat. 1 Mill.��� (English translation: Subhemispherical, black; head and prothorax with sides often reddish, its [the prothorax���] hind angles nearly straight [=right angled]; scutellum triangular, elytra very sparsely punctate, everywhere sparsely covered with long erect hairs; legs yellow. Length 1 ��, width 1 mm.), Published as part of Shockley, Floyd W. & Vandenberg, Natalia J., 2011, Notes on the taxonomic identity of Bystus hirtulus (Kirsch) and transfer from Endomychidae to Coccinellidae (Coleoptera: Cucujoidea), with designation of a lectotype for Alexia hirtula Kirsch, pp. 62-68 in Zootaxa 2868 on pages 64-65, DOI: 10.5281/zenodo.201518, {"references":["Kirsch, T. (1876) Beitrage zur Kenntnifs der Peruanischen Kaferfauna auf Dr. Abendroth's Sammlungen basirt. Deutsche Entomologische Zeitschrift, 20, 6 - 144.","Csiki, E. (1901) Catalogus Endomychidarum. Termeszetrajzi Fuzetek, 24, 2 - 53.","Csiki, E. (1910) Pars 12: Fam. Endomychidae. In: S. Schenkling (Ed), Coleopterorum Catalogus. W. Junk, Berlin, pp. 1 - 68.","Strohecker, H. F. (1953) Coleoptera, Endomychidae. In: Wytsman, P. (Ed), Genera Insectorum. Louis Desmet-Verteneuil, Bruxelles, pp. 1 - 145."]}

Sticholotis magnostriata sp. n. (Figs. 33���39) As mentioned earlier, some species of Sticholotis seem to be rather larger than average and are similar to Stictobura. A new species of Sticholotis from northeastern India (from the state of Assam), which was sent as part of a loan of Stictobura specimens received from BMNH, is described below. Diagnosis. The elytral pattern and presence of characteristic dual punctures on elytra with larger punctures arranged in striae readily separate this species from the other species of Sticholotis and Stictobura of this region. The male genitalia are diagnostic. Description. Male: Length: 4.50���4.75 mm; width: 4.00��� 4.50 mm; TL/EW: 1.04���1.07; EL/EW: 0.88���0.91; PL/PW: 0.43���0.44. Form (Fig. 33) more or less circular, moderately convex; dorsum apparently glabrous. Head and pronotum reddish brown; ground colour of elytra dark brown, with seven spots arranged in a 2 - 2 - 1-2 pattern as follows: three pairs of yellowish spots���one on basal margin on either side of scutellum, one on lateral side just before middle, and one in posterior half; and one reddish brown, much larger, transverse oval spot around middle of elytra; lateral margins of elytra paler, reddish brown (Figs. 33, 34). Antenna and mouthparts reddish brown. Ventral side more or less uniform reddish brown, metaventrite slightly darker brown in the paratype. Head (Fig. 35) with anterior clypeal margin distinctly deeply emarginate medially; eyes broadly separated, interocular distance ca. 2.7 x as wide as an eye; densely punctate with punctures separated by their own diameter or less, slightly more widely spaced towards clypeal margin; interspaces with strong reticulate microsculpture. Antenna 11 -segmented. Pronotum with much finer punctures than those on head and more widely separated by 3���5 diameters, interspaces apparently smooth, punctures on anterior margin much finer than those on disc, punctures on anterolateral corners slightly more closely spaced, with weakly reticulate microsculpture between interspaces. Elytra slightly broader than long, with lateral margins weakly explanate; punctures distinctly dual, large punctures mainly arranged in slightly irregular rows, at least six rows clearly visible from sutural line, particularly around middle, somewhat tapering off towards apices, one complete row of larger punctures apparent on lateral margin; punctures between rows much finer, widely separated,; interspaces between punctures smooth, shiny. Prosternal intercoxal process trapezoidal and strongly narrowed towards posterior, anterior margin triangularly emarginate with a short median tubercle, anterior margin of mesoventrite shallowly emarginate. Epipleura foveolate on level with middle and hind legs. Tarsi cryptotetramerous. Abdominal postcoxal lines incomplete, ventrite 5 apically shallowly emarginate. Male genitalia (Figs. 36���39) with basal lobe elongate cylindrical anteriorly, gradually widened towards apex, apically rounded and produced into a short, tubular process in ventral view (Fig. 37), parameres longer than basal lobe, apically densely hairy; sipho (Figs. 38, 39) as illustrated with a distinct capsule. Female: Not known. Specimens examined. Holotype male: Doherty/ Assam, Patkai Mts / Fry Coll. 1905.100 (dissected, male genitalia in vial on same pin) (BMNH); Paratype male: With the same data as holotype (BMNH). Etymology. The species epithet is in reference to its large size and arrangement of larger elytral punctures apparently in the form of striae., Published as part of Vandenberg, Natalia J., 2011, A revision of the genus Stictobura Crotch and description of a new species of Sticholotis Crotch (Coleoptera: Coccinellidae: Sticholotidinae), pp. 1-13 in Zootaxa 3031 on page 12, DOI: 10.5281/zenodo.202599

Sticholotis buruensis Korschefsky, n. comb. (Fig. 11) Stictobura buruensis Korschefsky, 1944: 48 (Holotype:?ZMAN).���Miyatake, 1997: 265. Redescription. Length: 2.2 mm; width: 2.0 mm. Form slightly elongate, globose; dorsum strongly convex. Dorsal and ventral surfaces including appendages yellow ferrugineous, edges of sclerites and elytra narrowly darker reddish amber; six more or less defined rows of elytral striae marked by darkly pigmented dots. Head (Fig. 11) and pronotum with shallowly impressed punctures separated by about a diameter, punctures on pronotum slightly larger than on head, interspaces with reticulate microsculpture on head, much smoother with only trace of reticulation on pronotum; elytra smooth, shiny with extremely shallow punctures separated by 2���4 times a diameter; appearing almost impunctate except at high magnification. Head with fine sparse even decumbent pubescence, each hair separated by about its length; clypeus with apex nearly linear. Eyes small, separated by more than 3 times eye width, coarsely faceted, posteriorly divergent towards temples, with weak orbital carina; ocular canthus triangulate, extending onto eye for distance of about 3 facets. Antennal insertions exposed, frons around antennal insertions emarginate. Antenna with 10 segments, with three-segmented oval club. Terminal segment of maxillary palpus narrowly conical, apex attenuate; membrane of sensory surface appearing as a narrow seam on mesal surface in distal one-fourth, slightly expanded at distal end. Pronotum transverse; basal margin strongly arcuate, with submarginal line; lateral margins nearly linear, with short scattered hairs; anterior margin deeply trapezoidally emarginate around head; lateral and anterior margins with fine bead. Scutellum minute. Elytra lacking distinct humeral calli, appearing glabrous, but with very short, erect, scattered setae visible at high magnification; lateral borders narrowly reflexed. Prosternum lacking carinae, slightly convex, anteriorly arcuate and flanged, tapered to a blunt point posteriorly. Mesoventrite trapezoidal, widest anteriorly, weakly emarginate medially with small cavity on anterior face to receive tip of prosternum. Abdomen with five visible ventrites; abdominal postcoxal line incomplete, curved posterolaterally, joining posterior margin of ventrite 1; ventrites 2���4 short, subequal; ventrite 5 elongate, tapered, posteriorly arcuate, extreme apex obscured by paper mount. Condition of metathoracic wing not assessed. Elytral epipleura slightly narrower than width of mesocoxa, tapered in posterior half of length, complete, with inner carina reaching apex, shallowly foveolate or depressed to receive middle and hind femora. Legs with cryptotetramerous tarsi; tarsal claws narrow, sickle-shaped, basally slightly swollen, lacking a distinct basal tooth. Genitalia not dissected. Specimen examined. Paratype: ���L.J. TOXOPEUS/Buru.Station 17 / 22-23 Oct. ��� 21 (rectangular white label)/ TYPUS (retangular reddish brown label)/ Korschefsky collection/ 1952 (rectangular white label)��� (USNM). Distribution. Known only from the type locality, Buru Island, Indonesia. Notes. Korchefsky (1944) described this species from two specimens acquired from J. B. Corporaal, Zoologischen Museum, Amsterdam (ZMAN). The holotype was supposedly returned to the Amsterdam museum, but it is not listed in the online database on Coleoptera types from the Orient hosted by ZMAN (2010). The second specimen in the type series was retained in Korschefsky���s private collection which was later acquired by the USNM. The USNM specimen of S. buruensis is presumed to be the paratype. It agrees with the label data in the original description except that Korschefsky���s writeup transposes the L. and the J. (i.e. He wrote ���J. L. Toxopeus���) and there appears to be a transcription error in the date which was written as ��� 22���230 et., 1921.��� Examination of the USNM specimen confirms our suspicion that the species is not correctly classified in Stictobura, but belongs instead in the genus Sticholotis (Slipinski, 2004; Wang et al., 2010). Korschefsky stated that this species is closely related to S. gibbula (see preceding entry). Of the Neotropical members of the tribe Sticholotidini, S. buruensis is most similar to members of the genus Nexophallus Gordon (1969), but differs in having an incomplete postcoxal line and tarsal claw lacking the small basal tooth., Published as part of Vandenberg, Natalia J., 2011, A revision of the genus Stictobura Crotch and description of a new species of Sticholotis Crotch (Coleoptera: Coccinellidae: Sticholotidinae), pp. 1-13 in Zootaxa 3031 on page 9, DOI: 10.5281/zenodo.202599, {"references":["Korschefsky, R. (1944) Neue altweltliche Coccinelliden (Coleoptera: Coccinellidae). Arbeiten uber morphologische und taxonomische Entomologie aus Berlin-Dahlem, Band 11, Nr. 1, 47 - 56.","Slipinski, A. (2004) Revision of the Australian Coccinellidae (Coleoptera). Part 2. Tribe Sticholotidini. Annales Zoologici, 54 (2), 389 - 402.","Wang, X. - M., Ren, S. - X. & Chen, X. - S. (2010) The genus Nesolotis Miyatake (Coleoptera: Coccinellidae) from China with descriptions of eight new species. Annales Zoologici, 60, 1 - 13.","Gordon, R. D. (1969) A new genus and two new species of Sticholotini (Coleoptera: Coccinellidae) from South America. Coleopterists Bulletin, 23, 93 - 99."]}

Stictobura Crotch, 1874 Stictobura Crotch, 1874: 201; Sicard, 1911: 385. Type species: Calvia? pallideguttata Mulsant, 1853, by monotypy. Apterolotis Weise, 1908: 225; 1923: 127.��� Sicard, 1911: 385 (synonymy).��� Korschefsky, 1931: 211. Type species: Sticholotis (Apterolotis) andrewesi Weise, 1908 (= Stictobura pallideguttata (Mulsant, 1853), by subsequent designation of Korschefsky, 1931. Diagnosis. Body medium to large, with a round or subcircular outline (Figs. 1���7); dorsum very strongly convex and hemispherical, without apparent vestiture but characterized by sparse, thin, suberect to erect hairs on pronotum and elytra, easily noticeable particularly on anterior, lateral and posterior margins of elytra, those on disc noticeable only at high magnifications. Head (Fig. 12) with clypeal margin always distinctly semi-circularly emarginate. Eyes small, widely separated, coarsely faceted, inner margins with distinct orbital carinae, posteriorly divergent towards temples; ocular canthus extending shortly into eyes. Antennal insertions exposed, frons around antennal insertions distinctly, deeply emarginate. Antenna (Fig. 13) 11 -segmented, with a three-segmented, elongate, fusiform club, terminal antennomere apically elongate, conical-oval. Terminal segment of maxillary palpi (Fig. 15) elongate conical, apical margin strongly and obliquely truncate, shorter than outer margin. Pronotum transverse, anterior margin deeply trapezoidally emarginate around head; posterior margin strongly arcuate, with submarginal line; lateral margins linear to broadly rounded, with short scattered hairs, finely carinate, narrowly reflexed. Interspaces between punctures on head and pronotum always with strong reticulate microsculpture. Elytra lacking humeral calli, lateral borders with a distinct marginal bead; punctation on elytra often conspicuously dual, with fine and coarser punctures intermixed. Prosternum T-shaped, anterior margin not lobed in front of coxa, prosternal process (Fig. 17) very broad, quadrate with a pair of carinae. Anterior margin of mesoventrite broadly shallowly emarginate medially. Abdomen with five visible ventrites in both sexes, ventrites 1 and 5 subequal, longer than rest; abdominal postcoxal lines (Fig. 16) incomplete, parallel to or approaching posterior margin of ventrite 1. Functional wings absent. Elytral epipleura broad, complete with inner carina reaching apex of elytra, shallowly foveolate or depressed on level with middle and hind legs. Legs with cryptotetramerous tarsi, tarsal claws swollen basally, lacking a distinct basal tooth. Male genitalia (Figs. 19���31) with phallobase having an additional median strut besides trabes; basal lobe [=���penis guide��� sensu Ślipiński (2007), or ���median lobe��� auctorum] of tegmen elongate, symmetrical, parameres long with several marginal and apical setae; sipho [=���penis��� sensu Ślipiński (2007)] stout, with a large, prominent capsule. Female genitalia with coxites elongate triangular, bearing short styli, with a very long bursa and a rather large and moderately sclerotized structure (Fig. 32), representing a poorly differentiated spermatheca, or possibly just a lobe of the bursa. Distribution. This genus is apparently endemic to the Western Ghats range in southern India. Its members are rarely collected and have been found almost exclusively in plantations at high altitudes. Related genera. Stictobura is very closely related to Sticholotis. Its species differ from those of the latter only by their distinctly larger size, strongly convex / hemispherical dorsum, strongly reticulate microsculpture on interspaces between punctures on head and pronotum, elytra with a distinct marginal bead, absence of functional wings, and male genitalia with a very stout sipho having a large capsule. Stictobura species have sparse, somewhat inconspicuous, but more or less uniform, fine, suberect to erect pubescence on the pronotum and elytra, though most of the old specimens examined in this study appeared to be glabrous due to abrasion. Many species of Sticholotis appear to be glabrous, but have sparse, very short erect hairs on the elytra. Functional wings are absent in many Australian species of Sticholotis and all the species of Nesolotis Miyatake (1966). Ślipiński (2004) regarded Nesolotis as a synonym of Sticholotis, but Wang et al. (2010) resurrected the genus based on its distinctly foveate elytral epipleura, tibiae of front legs strongly expanded externally, and other features. Some unusually large species of Sticholotis that resemble Stictobura occur in Vietnam, northeastern India, and Indonesia. Among other Asian Sticholotidini, Jauravia Motschulsky (1858) shares many features with Stictobura, but has conspicuous dense pubescence all over the body. Filipinolotis Miyatake (1994) appears to be close to Stictobura by virtue of its strongly convex dorsum and atrophied hind wings. Among the Neotropical genera, the Hispaniolan genus Bura Mulsant (1850) has a similar size and body form to Stictobura, but differs in the highly polished appearance of the head and pronotal interspaces, and in possessing fully developed metathoracic wings, 10 -segmented antennae, clypeus anteriorly truncate and not emarginate around antennal insertions, and tarsal claws with a large triangular basal tooth (Vandenberg and Perez-Gelabert, 2007). The composition of Sticholotidini and the taxonomic status of many genera therein have not been fully resolved yet and the ongoing comprehensive studies including molecular studies on the world taxa of Sticholotidinae by other coccinellid workers may throw more light on the interrelationships among these genera. Hence, in spite of its very close relationship with Sticholotis, Stictobura is treated as a distinct genus in this paper. Biology. Detailed biology is not known for all the species, but prey records from label data indicate a preference for Coccoidea (Diaspididae; Coccidae). Rao et al. (1970) reported a Stictobura sp. as feeding on tea red spider mites. Almost all available specimens in collections have been collected on plantation crops such as tea and coffee., Published as part of Vandenberg, Natalia J., 2011, A revision of the genus Stictobura Crotch and description of a new species of Sticholotis Crotch (Coleoptera: Coccinellidae: Sticholotidinae), pp. 1-13 in Zootaxa 3031 on pages 2-3, DOI: 10.5281/zenodo.202599, {"references":["Crotch, G. R. (1874) A Revision of the Coleopterous Family Coccinellidae, E. W. Janson, London. 311 p.","Sicard, A. (1911) Especes et varietes nouvelles de Coccinellides de l'Inde [Coleopt.] appartenant a la collection de M. Andrewes (de Londres). Annales de la Societe Entomologique de France, 79 [1910], 377 - 389.","Mulsant, E. (1853) Supplement a la monographie des coleopteres trimeres securipalpes. Annales de la Societe Linneenne de Lyon (N. S.), 1, 129 - 333. [reprinted as Opuscules Entomologiques, 3, 1 - 205.]","Weise, J. (1908) Coleopteren aus Ostindien. Stettiner Entomologische Zeitung, 69, 213 - 230.","Korschefsky, R. (1931) Coleopterorum Catalogus. Pars 118. Coccinellidae I. Berlin, 224 p.","Slipinski, A. (2007) Australian ladybird beetles (Coleoptera: Coccinellidae): their biology and classification. Australian Biological Resources Study, Canberra, 306 p.","Miyatake, M. (1966) A new genus belonging to the tribe Pharini from the northern part of the Ryukyu Islands (Coleoptera: Coccinellidae). Transactions of the Shikoku Entomological Society, 9, 47 - 50.","Slipinski, A. (2004) Revision of the Australian Coccinellidae (Coleoptera). Part 2. Tribe Sticholotidini. Annales Zoologici, 54 (2), 389 - 402.","Wang, X. - M., Ren, S. - X. & Chen, X. - S. (2010) The genus Nesolotis Miyatake (Coleoptera: Coccinellidae) from China with descriptions of eight new species. Annales Zoologici, 60, 1 - 13.","Motschulsky, V. (1858) Insectes des Indes Orientales. Etudes Entomologiques, 7, 117 - 122.","Miyatake, M. (1994) Revisional studies of Asian genera of the subfamily Sticholotidinae (Coleoptera: Coccinellidae). Memoirs of the College of Agriculture-Ehime University, 38 (2), 223 - 293.","Mulsant, E. (1850) Species des Coleopteres trimeres securipalpes. Annales des Sciences Physiques et Naturelles, d'Agriculture et d'Industrie, Lyon, (2) 2, 1 - 1104.","Vandenberg, N. J. & Perez-Gelabert, D. E. (2007) Redescription of the Hispaniolan ladybird genus Bura Mulsant (Coleoptera: Coccinellidae) and justification for its transfer from Coccidulinae to Sticholotidinae. Zootaxa, 1586, 39 - 46.","Rao, V. P., Datta, B. & Ramaseshaiah, G. (1970) Natural enemy complex of flushworm and phytophagous mites on tea in India. Scientific Publication Series No. 5, Tea Board, India. 53 p."]}

3. semipolita Sicard (Figs. 2 ���6, 18a���e, 24���27) Stictobura semipolita Sicard, 1911: 384 (Lectotype; BMNH. Type locality: Nilgiri Hills).��� Korschefsky, 1931: 211 (cat.) Stictobura semipolita ab. fuscipes Sicard, 1911: 385.��� Korschefsky, 1931: 211. Stictobura semipolita ab. testaceicollis Sicard, 1911: 385.��� Korschefsky, 1931: 211. Stictobura rubroguttata Sicard, 1925: 449 (Lectotype, BMNH).���Synonymised by Korschefsky, 1933: 7. Diagnosis. Stictobura semipolita has variable elytral colour and pattern (Figs. 2 ���6, 18a���e). It is somewhat similar in terms of general colour scheme and elytral pattern to Sticholotis obscurella Weise (1908), but it can be easily differentiated from the latter by the much larger size, more strongly convex body and the male genitalia. Redescription. Length 3.8 ���4.0 mm; width 3.5���3.7 mm; TL/EW: 1.12���1.14; EL/EW: 1.03���1.12; PL/PW: 0.42���0.45. Form circular (Fig. 5) to distinctly obovate (Figs 2 ���4, 6), with elytra posteriorly narrowed towards apex, dorsal side strongly convex. Head and pronotum yellowish-testaceous, elytral colour and pattern variable as follows: (i) elytra reddish with a broad black border (ab. fuscipes Sicard) (Figs. 5, 18e), (ii) sutural border also black with a median circular spot, with two reddish brown, elongate discal spots (nominate form) (these spots medially interrupted partially or fully in some examples) (Figs. 3, 4, 18 b���d), (iii) elytra dark brown to black with four distinct reddish brown spots (S. rubroguttata) (Figs. 2, 18a), and (iv) elytra more or less fully reddish brown (S. testaceicollis) (Fig. 6). Scutellum reddish brown. Ventral side more or less uniform yellowish brown, except antennomeres 5���11 and outer margin of elytral epipleura darker brown. Head with silvery white pubescence, punctures shallowly impressed, separated by 2���4 diameters, interspaces strongly reticulate. Antenna 11 -segmented, clothed with prominent yellowish white pubescence, club long, three-segmented, terminal antennomere elongate oval. Pronotum with shallow punctures, slightly more deeply impressed than those on head, separated by 2���5 diameters on disc, slightly denser and closer on lateral sides, interspaces strongly reticulate; posterior margin strongly arcuate, with submarginal line. Elytral punctures distinctly dual, large punctures separated by 2���4 diameters, with finer punctures in interstices, interspaces smooth, shiny. Prosternal process broad, subtrapezoidal, carinae distinctly divergent towards anterior. Anterior margin of mesosternum broadly, semicircularly emarginate in middle. Ventrite 5 conical with posterior margin arcuate in female, broader with posterior margin faintly emarginate in male. Male genitalia (Figs. 24���27) with basal lobe longer than parameres, apically very strongly arched and narrowed towards parameres in lateral view (Fig. 24); in ventral view (Fig. 25) elongate cylindrical, apically narrowed; sipho (Fig.26, 27) with a large, prominent, spatulate capsule. Specimens examined. Lectotype of Stictobura semipolita (designated here) : Male, labelled Type [printed, red-bordered circular Museum label]/ Nilgiri Hills. H.L. Andrewes. [printed]/ H.L. Andrewes Nilgiri Hills [printed]/ Type [printed in red ink]/ Andrewes Bequest B.M. 1922 - 221. [printed]/ Stictobura semipolita n.sp. Sic [Sicard's handwriting] (BMNH). Other specimens: 6 specimens on 1 card, labelled Type [printed, red-bordered circular Museum label]/ Nilgiri Hills. H.L. Andrewes. [printed]/ H.L. Andrewes Nilgiri Hills [printed]/ Type [printed in red ink]/ Andrewes Bequest B.M. 1922 - 221. [printed]/ Stictobura semipolita n.sp. Sic [Sicard's handwriting]. 10 specimens on 3 cards, labelled Co-type [printed, green-bordered circular Museum label]/ Nilgiri Hills. H.L. Andrewes. [printed]/ H.L. Andrewes Nilgiri Hills [printed]/ Andrewes Bequest B.M. 1922 - 221. [printed]. 20 specimens on 7 cards, labelled Co-type [printed, green-bordered circular Museum label]/ Nilgiri Hills. H.L. Andrewes. [printed]/ Nilgiri Hills [printed]/ Andrewes Bequest B.M. 1922 - 221. [printed] (BMNH). S. semipolita ab. fuscipes (2 specimens): 1 directly pinned specimen, labelled Type [printed, red-bordered circular Museum label]/ Nilgiri Hills. H.L. Andrewes. [printed]/ H.L. Andrewes Nilgiri Hills [printed]/ Type [printed on orange background, Andrewes collection label]/ Type [printed in red ink]/ Andrewes Bequest B.M. 1922 - 221. [printed]/ Stictobura semipolita S. v. fuscipes [Sicard's handwriting]. 1 directly pinned specimen, labelled Co-type [printed, green-bordered circular Museum label]/ Nilgiri Hills. H.L. Andrewes. [printed]/ H.L. Andrewes Nilgiri Hills [printed]/ Andrewes Bequest B.M. 1922 - 221. [printed] (BMNH). S. semipolita ab. testaceicollis : 1 carded specimen, labelled Type [printed, red-bordered circular Museum label]/ Nadgani Malabar [hand written]/ Type [printed on orange background, Andrewes collection label]/ Type [printed in red ink]/ Andrewes Bequest B.M. 1922 - 221. [printed]/ Stictobura semipolita Sic. v. testaceicollis S [Sicard's handwriting] (BMNH). Lectotype of rubroguttata (designated here): ���Type (circular red bordered Label)/ Nilgiri Hills, H.L. Andrewes/ Andrewes Bequest B.M. 1922 - 221 / Nilgiri Hills / Stictobura rubroguttata Sic. n. s/r 9 s??? (in Sicard���s handwriting)/Type (rectangular orange label)/H.L Andrewes, Nilgiri Hills ��� (BMNH). Others: On coffee green bug, Gulikere estate, June 56, G.P.C. Basavanna 1368 / Com. Inst. Ent. Coll. No. 14949 / Stictobura nr. semipolita Sic. R.D. Pope det. 1956 / Pres. Comm. Inst. Ent. B.M. 1981 - 315 (BMNH); South India, Kerala State, Trivandrum, Ponmudi range, 10.v. 1973, Leg. T.R.S. Nathan, 2 ex.; Anamalai hills, 2400, Madras, JCM Gardner, 6.V. 1930 / 420 / Korschefsky collection, 1952 / Stictobura rubroguttata det. A.S. Ślipiński, 1 male; Kerala, Calicut dist., Chembra Peak, V. 1970, leg. T.R. Susainathan/ Collection of Verne Reaves, 1 female (MNHN). Distribution. This appears to be most common species of the genus in southern India, but is confined to the Western Ghats range (Tamil Nadu; Karnataka; Kerala). Notes. One of the long series of Sicard���s syntypes of S. semipolita (BMNH) has been designated as lectotype. The median lobe in the lectotype male (designated above) is rather less strongly curved than shown in the figures, but the sipho agrees well (dissected and studied by RGB). Although Sicard labelled his varieties "v.", he published them as "ab.", so the names are automatically infrasubspecific and therefore not available under the ICZN Code. The syntype of S. rubroguttata (BMNH, examined) is designated as lectotype (above) in order to fix the identity of the species and confirm its synonymy, because the original description gave no indication of the number of original specimens examined. It was earlier synonymised with S. semipolita by Korschefsky (1933), who mentioned that this is the darkest form of S. semipolita. The lectotype has yellowish-testaceous head and pronotum as the nominate form of S. semipolita, but the elytra are dark pitchy brown to black with four reddish spots, one pair in each half. The body outline is variable in S. semipolita ���elytra somewhat obovate and posteriorly distinctly narrowed towards apex (Figs. 2 ���4, 6) or distinctly rounded (Fig. 5). Korschefsky (1933) and Chatterjee and Bose (1933) provided brief notes on the species and illustrated the variations in elytral pattern. Biology. Predatory on coffee green scale, Coccus viridis (Green) (Hemiptera: Coccidae) (label data). Associated with scales infesting coffee and sandal (Chatterjee and Bose, 1933)., Published as part of Vandenberg, Natalia J., 2011, A revision of the genus Stictobura Crotch and description of a new species of Sticholotis Crotch (Coleoptera: Coccinellidae: Sticholotidinae), pp. 1-13 in Zootaxa 3031 on pages 5-8, DOI: 10.5281/zenodo.202599, {"references":["Sicard, A. (1911) Especes et varietes nouvelles de Coccinellides de l'Inde [Coleopt.] appartenant a la collection de M. Andrewes (de Londres). Annales de la Societe Entomologique de France, 79 [1910], 377 - 389.","Korschefsky, R. (1931) Coleopterorum Catalogus. Pars 118. Coccinellidae I. Berlin, 224 p.","Sicard, A. (1925) Descriptions de Coccinellides appartenant a la Collection de M. Andrewes, de Londres. Annals and Magazine of Natural History, (9) 15, 447 - 449.","Korschefsky, R. (1933) Entomological investigations on the spike disease of sandal (16). Coccinellidae (Col.). Indian Forest Records, 19 (6), 1 - 9.","Weise, J. (1908) Coleopteren aus Ostindien. Stettiner Entomologische Zeitung, 69, 213 - 230.","Chatterjee, N. C. & Bose, M. (1933) Entomological investigations on the spike disease of sandal (17). Coccinellidae (Col.). Supplementary data. Indian Forest Records, 19 (7), 1 - 10."]}

DNA barcodes have proven invaluable in identifying and distinguishing insect pests, most notably for determining the provenance of exotic invasives, but relatively few insect natural enemies have been barcoded. We used Folmer et al.'s (1994) universal invertebrate primers and Hebert et al.'s (2004) for Lepidoptera, to amplify 658 bp at the 5' end of the mitochondrial cytochrome oxidase c subunit I (COI) gene in five species of lady beetles from crop fields in six states in the US Mid-Atlantic, Plains and Midwest: three native species, Hippodamia convergens Guerin-Meneville, H. parenthesis (Say) and Coleomegilla maculata (De Geer); and two exotic species, Harmonia axyridis (Pallas) and Coccinella septempunctata Linnaeus. Sequence divergences within species were low, never exceeding 0.9% (Kimura 2-parameter distances). Sequence divergences between the two Hippodamia species ranged from 14.7 to 16.4%, mirroring the relationships found for other arthropod taxa. Among the exotic species, C. septempunctata sequences were as variable as those of the three native species, while H. axyridis populations comprised a single haplotype. Limited data on two Coleomegilla subspecies, C. m. lengi Timberlake and C. m. fuscilabris (Mulsant), are consistent with their belonging to the same species, although morphological and reproductive data indicate that they represent separate species. Our results support the general utility of COI barcodes for distinguishing and diagnosing coccinellid species, but point to possible limitations in the use of barcodes to resolve species assignments in recently divergent sibling species.

Anovia Casey 1908 (Figs. 1���41) Anovia Casey, 1908: 408. Leng, 1920: 214. Korschefsky, 1931: 96. Gordon, 1972: 26. (Type species: Scymnus virginalis Wickham, by monotypy). The type species for Anovia was originally described as Scymnus virginalis Wickham, but subsequent authors questioned the placement in Scymnus (Casey 1908). Casey (1908), noting several morphological similarities to both Rodolia Mulsant and Novius Mulsant, erected Anovia to accommodate this species and included all three genera in Exoplectrini. Leng (1920) included Anovia, Novius, and Rodolia in Noviini for the first time. Diagnosis. Adults of Anovia are diagnosed by the following combination of characters: body convex, subhemispherical dorsum that is widest just posterior to humeral angles (Figs. 17���22); all surfaces including eye facets covered with pale, posteriorly-directed vestiture; eye margin entire, not interrupted by an ocular canthus (Fig. 23); clypeal apex horizontal (Fig. 23); antenna with 8 articles, weakly clubbed (Fig. 25); and tarsi trimerous (Figs. 32���34)., Published as part of Forrester, Juanita A., Vandenberg, Natalia J. & Mchugh, Joseph V., 2009, Redescription of Anovia circumclusa (Gorham) (Coleoptera: Coccinellidae: Noviini), with first description of the egg, larva, and pupa, and notes on adult intraspecific elytral pattern variation, pp. 25-40 in Zootaxa 2112 on page 27, DOI: 10.5281/zenodo.187887, {"references":["Casey, T. L. (1908) Notes on the Coccinellidae. The Canadian Entomologist, 40, 393 - 421.","Leng, C. W. (1920) Catalogue of the Coleoptera of America, north of Mexico. Mount Vernon, New York, 470 pp.","Korschefsky, R. (1931) Pars 118, Coccinellidae. Coleopterorum Catalogus, W. Junk, Berlin, 224 pp.","Gordon, R. D. (1972) The tribe Noviini in the New World (Coleoptera: Coccinellidae). Journal of the Washington Academy of Sciences, 62 (1), 23 - 31."]}

Anisandrus maiche Stark (Figs. 1, 2, 4���6) Anisandrus maiche Stark 1936: 142. Anisandrus maiche Eggers 1942: 36 (Synonymy: Pfeffer 1944) Diagnosis. Specimens of A. maiche can be distinguished from other members of Anisandrus occurring in North America by the smaller body size (A. dispar, also a non-native species in North America, but is easily distinguished by its smaller body size ( 3.2 mm for A. dispar), and the interstriae on the elytral disc which have only one row of seriate punctures (Fig. 2) compared with 2���3 rows of punctures in A. dispar (Fig. 3). Distinguishing Anisandrus from other North American Xyleborina. Hulcr et al. (2007) restored the genus Anisandrus, and included species formerly in Xyleborus, Ambrosiodmus Hopkins and Cyclorhipidion Hagedorn. They state that Anisandrus belongs to a group of genera that are defined by a short, stout body, characteristically flattened antennal club, and, most importantly, the presence of a pronotal mycangial tuft of hairs (Fig. 1). Anisandrus is very similar to Xylosandrus except that the procoxae of Anisandrus are contiguous (Fig. 8), whereas those of Xylosandrus are separated (Fig. 7). In the key to species of Xyleborus north of Mexico by Rabaglia et al. (2006), species that are now recognized as species of Anisandrus are characterized by having the antennal club distinctly and obliquely truncate with segment 1 corneous (couplet 1), the anterior margin of the pronotum distinctly armed by several coarse serrations, and the body A. maiche with alterations in bold type. An illustrated, online key (http://xyleborini.tamu.edu/keys.php) to North American Xyleborina will be updated to include Anisandrus in the generic key and the following key to the four species in North America., Published as part of Rabaglia, Robert J., Vandenberg, Natalia J. & Acciavatti, Robert E., 2009, First records of Anisandrus maiche Stark (Coleoptera: Curculionidae: Scolytinae) from North America, pp. 23-28 in Zootaxa 2137 on page 24, DOI: 10.5281/zenodo.188523, {"references":["Stark, V. N. (1936) Novye vidy koroedov iz Aziatsko chasti SSSR [New species of bark beetles from the Asian part of the USSR]. Bulletin of the Far East Branch, Academy of Science USSR, Vladivostok (Akademiia Nauk SSSR, Dal' nevostochnyi filial, Vestnik) 18: 141 - 154 [In Russian].","Eggers, V. H. (1942) Zur palaarktischen Borkenkaferfauna (Coleoptera: Ipidae), VIII. Borkenkafer aus dem asiatischen Russland. Arbeiten uber morphologische und taxonomische Entomologie [In German].","Pfeffer, A. (1944) Bemerkungen zur Arbeit von Hans Eggers: Zur Palearktischen Borkenkaferfauna. VIII. Borkenkafer aus dem asiatischen Russland (Col.: Ipidae). Arbeiten uber Morphologische und Taxonomische Entomologie, 11,130 - 131 [In German].","Hulcr, J., Dole, S. A., Beaver, R. A., & Cognato, A. I. (2007) Cladistic review of taxonomic characters in Xyleborina (Coleoptera: Curculionidae: Scolytinae). Systematic Entomology, 32 (3), 568 - 584.","Rabaglia, R. J., Dole, S. A. & Cognato, A. I. (2006) Review of American Xyleborina (Coleoptera: Curculionidae: Scolytinae) occurring north of Mexico, with an illustrated key. Annals of the Entomological Society of America, 99 (6), 1034 - 1056."]}

Anovia circumclusa (Gorham) (Figs. 1���41) Zenoria circumclusa Gorham, 1899: 262. Korschefsky, 1931: 108. Blackwelder, 1945: 443. Anovia circumclusa Gordon, 1971: p. 1; Gordon, 1972: 27 ���29. (Type depository: BMNH). Diagnosis: The larva of this species resembles all other known noviine larvae, but is distinguishable by the presence of many chalazae on the lateral strumae of the abdominal segments (R. cardinalis has 2, and R. koebelei has 4). Anovia circumclusa adults are best recognized by the structure of the male genitalia. In A. circumclusa, the basal lobe is slender and does not extend laterally beyond the internal paramere margin, while in all other Anovia species the basal lobe is quite broad distally, and overlaps the medial paramere margin. Also, the basal piece is widest basally in A. circumclusa, not distally as in A. virginalis. Egg. Length 0.5 mm, width 0.25 mm. Elongate-oval, color bright magenta. Surface granular, often covered with waxy exudate (Fig. 1). Eggs typically oriented horizontally, not placed on end; laid singly or in small clusters on exposed leaf surfaces; often laid on or under prey (Majerus 1994, JAF pers. obs.) Mature Larva. Length 5���7 mm, (Figs. 2���4). Body ovoid, convex, widest at midpoint, laterally arcuate. Color bright magenta with waxy, white exudate. Dorsal surface moderately setose, finely granulate, covered with waxy exudate (Figs. 2���4). Setae pale, erect, simple, length variable. Head (Figs. 5���7) prognathous, darkly pigmented, subquadrate, at least twice as long as wide; dorsal and lateral surfaces with several chalazae; seta-like asperities lateral to frontal arms (Fig. 5). Frontal arms vshaped; epicranial stem short, about as wide as long; median endocarina absent. Stemmata arranged in triangular pattern, three on each side. Antenna inserted anteromesad to stemmata, 2 -segmented (Fig. 9). Antennomere I robust, length ~ 1 / 3 width; II small, length subequal to width, sensorium longer than antennomere I. Labrum distinct, subrectangular, weakly bilobed apically (Figs. 5, 7). Mandible triangular, enlarged basally, falcate apically (Figs. 7, 8). Maxillolabial complex retracted (Figs. 6, 7). Maxilla with cardo and stipes fused to form solid, sclerotized structure with slender, arm-like extensions passing anteriorly and laterally around labial palpi; maxillary palpomere 2 -segmented; I much broader than long; II about as broad as long (Figs. 6, 7). Mala membranous, transverse. Hypopharyngeal bracon present, well-developed. Thoracic segments each with a pair of sclerotized plates; meso-and metathorax each with a pair of lateral strumae; struma bearing many chalazae (Figs. 2���4). Legs long, robust, strongly sclerotized dorsally, semimembranous and unpigmented ventrally (Figs. 10���12). Coxa transverse (Fig. 3). Femur robust, almost as broad as long (Figs. 3, 10���12). Tibia elongate, ventral surface distally setose; distal setae flat, clavate (Figs. 10���13). Tarsungulus strongly curved, basal tooth well-developed. Abdomen 10 -segmented; segments I���IX with 2 pairs of sclerotized tubercles, 1 pair of chalazate strumae, and 1 pair of annular spiracles; X bearing pygopod (Fig. 3). Pupa. Length 4.5���5.5 mm, width 2.5 ��� 3.5 mm, exarate (Figs. 14���16). Dorsal habitus elliptical, convex, partially covered in last larval exuvium, attached by cauda to substrate. Color (excluding exuvium) magenta with pale setae (Fig. 15, 16). Dark, stout, bristle-like setae present on dorsal surface of head, pronotum, and humeral angles (Fig. 15). Head length subequal to width. Antenna short, not extending beyond outer margin of eye, club indistinguishable from flagellum. Apical maxillary palpomere strongly securiform (Fig. 16). Abdomen with 9 ventrites, I and II reduced and hidden beneath metacoxae; dorsal surface of abdomen with paired transverse tubercles on segments I���VIII; anterolateral angles with annular spiracles; IX with bipartite urogomphi. Adult. Length 4 ���4.5 mm. Dorsal habitus hemispherical, laterally arcuate, convex; head strongly deflexed, not visible from above; color variable (Figs. 17���22). Vestiture pale, short, moderately dense, posteriorlydirected. Head width about twice head length; dorsal surface with evenly spaced, small, shallow punctures; ventral surface narrower; postoccipital margin sinuate (Figs. 23, 24). Eyes large, covered entirely by pale, suberect setae. Antennal insertion exposed, anteromesad to inner eye margin. Antenna with 8 articles; antennomere I asymmetrical, laterally expanded; II subglobose; III���V subequal in length and width; VI���VIII forming loose club, VI���VII asymmetrical, expanded medially; VI about as long as IV + V, VII shorter, VIII broadly tapered apically (Fig. 25). Clypeus small, fused to frons (Figs. 23, 24). Frontoclypeal suture absent. Labrum (Fig. 26) emarginate medially, expanded beyond clypeus laterally. Mandible apically bidentate, teeth sickle-shaped, not in same plane, ventral tooth longer than dorsal one; prosthecal fringe well-developed (Fig. 27). Lacinia slender, elongate, apically setose (Fig. 28). Galea broad, elongate, truncate and apically setose. Maxillary palp 3 -segmented, palpifer well-developed; palpomere I elongate, about three times as long as basal width, broadest apically, membranous surface exposed; II apically divergent, mesal edge short, membranous surface exposed; III with distal edge almost twice the length of proximal edge, lateral edge twice the length of mesal one (Fig. 28). Labium narrow, labial palp 2 -segmented; palpomeres I and II subequal in size, palpomere II gradually narrowed distally to apical sensory area (Fig. 29). Pronotum with dorsal surface punctate, moderately setose; anterior angles extending forward just beyond lower margin of eye (Figs. 18, 20, 22); anterior edge horizontal just behind head capsule; posterior edge markedly sinuate, slightly notched at scutellum (Fig. 30). Prosternum narrow; prosternal process abruptly raised, rectangular with margins entire; procoxal cavities slightly transverse, closed internally (Fig. 31). Scutellum triangular. Meso- and metathorax ventrally flattened, pubescent (Fig. 35). Mesoventrite short, narrowest posteriorly. Metaventrite wider than long, finely punctate. Legs (Figs. 32���34) flattened, broad and stout. Femur deeply grooved ventrally for reception of tibia; groove bicarinate, sharply defined, extending almost entire length of femur. Profemur with anterior groove expanded prior to apex. Tibia slightly widened at mid-length, ventral surface broader than dorsal, deeply grooved for reception of tarsus; groove bicarinate. Tarsal formula 3 - 3 - 3; tarsomeres I and II elongate, lobed ventrally with spongy pubescence; III elongate, cylindrical; male tarsal claw bifid; female tarsal claw with long triangular tooth (Fig. 38). Elytron subhemispherical to hemispherical, laterally arcuate, finely punctate, non-striate; epipleuron complete to posterior margin, ventral surface moderately rugose. Wing with reduced cantharoid venation, absent in distal half, with strong medial and cubital veins, one anal vein, jugal lobe present. Abdomen with broad, slightly cleft intercoxal process; postcoxal line incomplete to lateral margin; 6 ventrites; I���V rectangular, progressively narrower in width posteriorly; VI narrower, tapering slightly to rounded apex; male with emarginate apex (Fig. 37), female with apex entire. Pygidium subrectangular, setose, broadly rounded apically (Fig. 36). Male genitalia with phallobase widest anteriorly; basal lobe slender, not extended laterally beyond internal margin of parameres (Figs. 39���41). Sipho as in Fig. 41. Material examined: see Table 2., Published as part of Forrester, Juanita A., Vandenberg, Natalia J. & Mchugh, Joseph V., 2009, Redescription of Anovia circumclusa (Gorham) (Coleoptera: Coccinellidae: Noviini), with first description of the egg, larva, and pupa, and notes on adult intraspecific elytral pattern variation, pp. 25-40 in Zootaxa 2112 on pages 27-32, DOI: 10.5281/zenodo.187887, {"references":["Gorham, H. S. (1899) Insecta. Coleoptera. Supplement to Endomychidae and Coccinellidae. Biologia Centrali- Americana. R. H. Porter, London, 276 pp, 13 pl.","Korschefsky, R. (1931) Pars 118, Coccinellidae. Coleopterorum Catalogus, W. Junk, Berlin, 224 pp.","Blackwelder, R. E. (1945) Checklist of the Coleopterous Insects of Mexico, Central America, the West Indies, and South America. Part 3. United States National Museum Bulletin 185. 188 pp.","Gordon, R. D. (1971) A revision of the genus Zenoria Mulsant (Coleoptera: Coccinellidae). Smithsonian Contributions to Zoology, 86, 1 - 22.","Gordon, R. D. (1972) The tribe Noviini in the New World (Coleoptera: Coccinellidae). Journal of the Washington Academy of Sciences, 62 (1), 23 - 31.","Majerus, M. E. N. (1994) Ladybirds. Harper Collins, London, 367 pp."]}

Shockley, Floyd W., Vandenberg, Natalia J. (2009): Catalogue of the primary types of Cerylonidae, Endomychidae and Latridiidae (Coleoptera: Cucujoidea) deposited in the National Museum of Natural History, with additional notes and clarification of the status of several types 2229. Zootaxa 2229 (1): 1-64, DOI: 10.11646/zootaxa.2229.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.2229.1.1

Cycloneda pretiosa Vandenberg & Gonz��lez, new species (Figs.1, 2 c, 3���8) Diagnosis: Distinguished from other Cycloneda species by tarsal claw lacking a basal tooth (Fig. 2 c), and distinctive tricolored elytron featuring a large sinuous black vitta and 3 or 4 yellowish cream-colored dots against a saturated red to orange background (Fig. 1). At present this is the only known species of Cycloneda where the tarsal claw is simply widened basally with a superficial medioventral notch (Fig. 2 c), however two other members of the Cycloneda germainii species complex (C. lacrimosa Gonz��lez & Vandenberg and C. disconsolata Vandenberg & Gonz��lez) have a basal tooth that is greatly reduced in length (Fig. 2 b). Although the one-two-one arrangement of yellowish cream-colored elytral maculae is consistent with other members of the C. germainii species complex, the appearance of the new species is perhaps more evocative of C. pulchella (Klug) from Brazil and Argentina and C. bioculata (Korschefsky) from Brazil, both tricolored species with a yellow to red background and round cream-colored spots embedded in larger black maculae. Upon closer examination, however, it is clear that the arrangement of the tricolored elements differs among these taxa, and both C. pulchella and C. bioculata are more orbicular in body form, with a much broader elytral epipleuron (subequal to width of frons) and different form of the male genitalia. Cycloneda pulchella has the basal lobe strongly, almost angularly curved down and then up, with a truncate apex and a dorsal patch of setae on each side near base, suggesting a closer affinity to C. emarginata (Mulsant) or C. ecuadorica (Timberlake), than to C. pretiosa. Cycloneda bioculata has a basal lobe that is much shorter and broadly rounded apically, with parameres that project well beyond the basal lobe. Description (Holotype male): Length 3.5 mm, width 2.3 mm. Form elongated oblong, convex, apically pointed; elytral and pronotal margins narrowly reflexed. Punctation on dorsal surface fine, regular, with each puncture separated by 2��� 3 X its diameter; surface between punctures shiny, with only faint trace of reticulation on head, not reticulate on pronotum or elytron. Dorsal color pattern as follows: Head black with two irregular cream-colored spots, one at medial margin of each eye, extending from eye canthus to just below level of upper 1 / 3 of eye; eye black; labrum translucent brown. Pronotum (Fig. 4) black with triangulate yellow cream-colored spot beginning at anterior margin over medial margin of eye, tapered obliquely along lateral margin to basal 1 / 2, enclosing entire anterior angle. Scutellum black. Elytral ground color reddish orange with black sinuous band, and four yellow cream-colored spots. Black band about 2 / 5 elytral width, filling most of disc, beginning near suture just beyond scutellum at basal 1 / 8, curved obliquely toward lateral margin just before basal ��, then closely following elytral curvature, joined to lateral margin at apical ��, continued along margin to apex. Four yellow cream-colored spots irregularly rounded to oblong, disposed as follows: first spot at base adjacent to scutellum, touching base of black band; second and third spots in a row slightly before middle of elytron, outer spot of pair adjacent to reflexed margin, inner spot midway between outer spot and suture; fourth spot oblong, situated in apical declivity at apical 1 / 5 adjacent to reflexed outer margin; first, second and forth spots forming oblique line, equally spaced. All margins of elytron with same orange of background color, except apex black. Lateral orange margins narrowly transparent. Ground color of ventral surfaces black; elytral epipleuron orange, except apex black. Pronotal hypomeron straw-colored in apical 2 / 3; antenna, mouthparts brown; legs black, except tarsus obscure brown. Ventral surfaces including appendages clothed in decumbent silvery pubescence. Eyes finely facetted separated by 2 ��X eye diameter; inner orbits nearly parallel in lower half, diverging at upper and extreme basal level. Antenna (Fig. 5) of 11 antennomeres, total length slightly greater than distance between eyes; third antennomere elongate, 1 ��X length of second, subequal to four plus five combined. Pronotum evenly convex except for very narrowly reflexed lateral margin; in outline with basal margin subsinuate, lateral margin strongly evenly arcuate, anterior margin subtrapezoidally emarginate; anterior angles subtriangulate, not conspicuous in dorsal view. Elytron elongate, in dorsal view with humeral angle rounded, weakly arcuate from beyond humeral angle to apical 1 / 3, slightly sinuate at 2 / 5 length, this more evident in ascendant side of elytron than in the reflexed margin; in lateral view (Fig. 3) unevenly arcuate, dorsally somewhat flattened in anterior 2 / 3, more abruptly declivitous in posterior 1 / 3, of approximately equal height at anterior, posterior ��; epipleuron flat, horizontal in anterior 1 / 3, progressively inwardly sloping in posterior half, with greatest width equal to about �� width of frons. Prosternum T-shaped, in ventral view with lateral arms gently folded back from stem, forming shallow arc in cross section; intercoxal process convex, inflated, apex truncate. Mesosternum trapezoidal; anterior border approximately linear with raised margin. Metasternum broad, with post mesocoxal line reaching lateral margin. Abdomen shortened semi-oval, broadest at second ventrite; postmetacoxal line of first abdominal ventrite curved posterolaterad, closely paralleling posterior margin for much of length, not attaining lateral margin; posterior margin of ventrites 1���5 linear; exposed portion of ventrite 6 rounded. Tarsal claw (Fig. 2 c) without tooth, but with a superficial medioventral notch, wider in proximal ��, evenly curved distally. Male genitalia (Fig. 6 a���c): Basal lobe elongate, roughly parallel-sided in basal ��, distal �� tapered, apex attenuate; parameres slender, reaching 4 / 5 distance to apex of basal lobe; sipho (Fig. 6 c) curved, widened in last 1 / 3; abruptly constricted, inflected in apical 1 / 6. Female: Similar to male, color slightly less reddish, black band narrower. The color pattern on the head as in the male, not gender specific. Female genitalia (Fig. 7) with spermathecal capsule weakly curved; infundibulum elongate, tubular. Variation: Length 3.4 to 3.7 mm. Fourth yellow cream-colored spot can disappear leaving only a fine yellow point. Black band can be expanded in lateral 2 / 3, or can be partially dissolved, leaving a mackerel pattern. Type material (map, Fig. 8): Holotype (male), ��� CHILE, Lo [Lago] Pirihueico [Comuna Panguipulli, Provincia Valdivia] fc [fecha] 15.01.0 2, coll: A.L��er��� (MNHN). Paratypes, Total 5 (females), 1 same data as holotype, 1 with date ��� 18.01.02 ��� and 3 with date ��� 19.01.02 ��� (2, MNHN; 1, USNM; 2, ALPC). FIGURE 8. Distribution of Cycloneda pretiosa, Vandenberg & Gonz��lez new species. Etymology: From the Latin ��� pretiosa ��� meaning very beautiful and of great value. Remarks: Cycloneda pretiosa can be incorporated into the existing key to species in the Cycloneda germainii species complex (Gonz��lez and Vandenberg, 2006) by adding an unnumbered couplet before existing couplet 1: Tarsal claw simply widened at base, lacking tooth, but with superficial medioventral notch (Fig. 2 c); ground color of elytron saturated red or orange with contrasting markings in black and yellowish cream (Fig. 1); black mark beginning near elytral base and continued to apex, sinuous, not interrupted at midlength .................................................................................. C. pretiosa Vandenberg & Gonz��lez, n. sp. Tarsal claw with basal tooth (Figs. 2 a���b); elytral color pattern not as above: ground color of elytron blackish, ferrugineus, or straw-colored; if orange then with a pair of irregular black marks generally well separated near midlength, or rarely with narrow connection at inner margins, and with cream colored markings absent or strongly suffused with orange ground color................................................................................. 1, Published as part of Vandenberg, Natalia J. & Gonz��lez, Guillermo, 2008, A new Chilean species of Cycloneda Crotch (Coleoptera: Coccinellidae: Coccinellinae: Coccinellini), pp. 63-68 in Zootaxa 1772 on pages 64-68, DOI: 10.5281/zenodo.182164, {"references":["Gonzalez, G. & Vandenberg, N. J. (2006) Review of lady beetles in the Cycloneda germainii species complex (Coleoptera: Coccinellidae: Coccinellinae: Coccinellini) with descriptions of new and unusual species from Chile and surrounding countries. Zootaxa, 1311, 13 - 50."]}

Bura Mulsant Bura Mulsant, 1850: 374, 419. Type species: Bura cuprea Mulsant, by monotypy. Diagnosis: Distinguished from most other Sticholotidini by the combination of circular, hemispherical to superhemispherical body form (Figs. 2���3), by the shape and development of the eye canthus (Fig. 1), clypeus not emarginate around antennal insertion, distal maxillary palpomere nearly parallel-sided (Fig. 7), and tarsal claw with a large subtriangular basal tooth (Fig. 8). Redescription: Form (Figs. 2���3) compact, hemispherical to superhemispherical, shiny with metallic sheen; dorsum apparently glabrous except for sparse marginal setae. Punctation on dorsal surfaces distinct, with intermixed large/small punctures on elytron, single-sized punctures on head and pronotum; minute seta associated with each puncture, scarcely visible using standard light microscopy. Head (Figs. 1���3) vertical, widest at middle of eye, tapered toward clypeus with scattered setae at inner margin of eye and anterior margin of clypeus; labrum distinctly narrower than clypeal margin, moderately setose, joined to clypeus by conspicuous trapezoidal membrane. Head capsule with slightly raised bead along inner margin of eye. Eyes (Fig. 1) well-developed, dorsally divergent; facets somewhat flattened, eye canthus long and narrow, with distal end slightly up-curved, partially dividing eye, subcarinate proximally, with several rows of facets visible below its ventral margin. Anterior, lateral margins of clypeus reflexed, not emarginate at antennal insertion. Antenna (Figs. 1, 5���6) inserted laterally beneath clypeal margin, moderately short and compact, composed of ten antennomeres; club gradual, well-developed, spindle shaped; mesal surface of penultimate antennomere projecting anterolaterally, with brush of short setae in small semi-membranous patch near apex; last antennomere with scattered short setae in distal half and dense concentration of short setae in large semi-membranous patch on mesal surface; both setal patches with indistinct boundaries. Mandible with bifid apex of which inner tooth shorter and thicker than outer tooth. Mentum moderately broadly joined to submentum. Distal maxillary palpomere (Fig. 7) elongate, nearly parallel-sided, apically pointed, with long oblique sensory surface. Distal labial palpomere elongate, tapered to very small round sensory surface. Pronotum with marginal bead continuous along base and lateral margin to inner anterior angle directly behind inner margin of eye, very narrowly, faintly indicated beyond; anterior angle and lateral margin narrowly reflexed, with short widely spaced setae along reflexed edge. Elytron with lateral margin narrowly, sharply reflexed with distinct lateral bead bearing sparse short setae. Scutellum small, triangular. Pronotal hypomeron with depression to receive retracted antennal club. Prosternum forming a modified T or Y-shape (Fig. 4); short stem strongly raised above level of lateral arms with carinae distinct at least in anterior half and joined to form an inverted U. Coxae broadly separated; meso- and metasternites compactly joined with dividing sutures partially obliterated. Femur robust; tibia simple, slender, not externally dentate, apical spurs lacking; tarsi cryptotetramerous; claw with well developed triangular tooth at base (Fig. 8). Elytral epipleuron broad in anterior half, may be subfoveolate to receive femoral apices in repose, complete, inner margin narrowed by arcuate expansion of abdomen, tapered toward apex. Abdomen with five ventrites, extreme tip of sixth may be visible in males; postcoxal line of first ventrite curved posterolaterally, not quite attaining junction of posterior and lateral margins. Male genitalia with slender curved sipho; basal lobe and parameres elongate, bilaterally symmetrical or nearly so; basal piece symmetrical, lacking eccentric dorsal strut. Remarks: In general facies, Bura looks very similar to many of the photographs of Australian Sticholotis species in a recent publication by Ślipiński (2004), but the latter have the clypeus emarginate around antennal insertions, distal maxillary palpomere spindle-shaped (apically convergent), median part of prosternum anteriorly produced, tarsal claw simple or merely broadened at base (lacking triangular tooth), and basal piece of male genitalia asymmetrical, with an eccentric dorsal strut. Bura appears to be closely allied to many of the Neotropical sticholotidine genera described by Gordon (1969, 1977, 1991, 1994 c), but these are generally small to minute beetles (1.8 to1.25 mm in length) while Bura cuprea exceeds 2.5 mm and can reach up to 3.2 mm in length. In addition to the greater size, Bura can be distinguished primarily by the well-developed triangular tooth of the tarsal claw (Fig. 8). It is perhaps most similar to the genus Lenasa Gordon (1994 c), but the latter is much smaller, more oval and less convex in body form, and with only a small acute tooth at the base of the tarsal claw. Bura also shares some features with the ���cocciduline��� taxa of the West Indies (Chapin 1930, Gordon 1994 b), but in this case the overall similarity is not sufficient to cause any confusion in identification. In his recent study of Sticholotis, Ślipiński (2004) chose to recognize a single broadly defined genus rather than continuing to segregate specialized species based on loss of flight wings or reduction in number of antennomeres. Presently, the Neotropical Sticholotidini consists of a number of small genera which share many features with Bura and with each other. If the trend begun with Sticholotis is continued in the world revision of Sticholotidinae (Ślipiński in prep.), we might expect a number of Neotropical sticholotidine genera to fall into synonymy with Bura. In particular the genus Lenasa does not differ from Bura except in the overall size and proportions of various body parts, not normally considered significant differences at the generic level., Published as part of Vandenberg, Natalia J. & Perez-Gelabert, Daniel E., 2007, Redescription of the Hispaniolan ladybird genus Bura Mulsant (Coleoptera: Coccinellidae) and justification for its transfer from Coccidulinae to Sticholotidinae, pp. 39-46 in Zootaxa 1586 on pages 44-45, DOI: 10.5281/zenodo.178466, {"references":["Gordon, R. D. (1969) A new genus and two new species of Sticholotini (Coleoptera: Coccinellidae) from South America. Coleopterists Bulletin, 23, 93 - 99.","Gordon, R. D. (1977) Classification and phylogeny of the New World Sticholotidinae (Coccinellidae). Coleopterists Bulletin, 31,185 - 228.","Gordon R. D. (1994 c) West Indian Coccinellidae VI (Coleoptera): New genera and species of Sticholotidini and a cladistic analysis of included genera. Journal of the New York Entomological Society, 102 (2), 232 - 241.","Chapin, E. A. (1930) New Coccinellidae from the West Indies. Journal of the Washington Academy of Sciences, 20 (20), 488 - 495.","Gordon, R. D. (1994 b) West Indian Coccinellidae V (Coleoptera): A review of Coccidulini and additions to Psorolyma Sicard. Journal of the New York Entomological Society, 102 (2), 222 - 231."]}

Cycloneda boliviana (Mulsant) new combination, status revised (Figs. 2 C; 3 G; I; 5 I–K; 7 B; 8 G; 9 H; 11) Coccinella boliviana Mulsant 1866: 75; Crotch 1874: 106 (as synonym of C. areata Mulsant); Weise 1910: 21 (as synonym of C. areata Mulsant); Gordon 1987: 12 (as synonym of C. areata Mulsant). Coccinella areata ab. boliviana Korschefsky 1932: 510; Blackwelder 1945: 454. Diagnosis: Distinguished from similar appearing members of the genus by the combination of antenna composed of 11 antennomeres (Fig. 9 H), pronotum with an even cream­colored or yellow anterior and lateral border (Fig. 3 I), and elytron with a pale subapical mark centered relatively distant from the apex (Fig. 5 I–K). This species is the largest in the C. germainii species complex, and has a distinctly matt or pruinose appearance to the elytra. It is most closely related to C. sicardi (see diagnosis above), but has even cream­colored pronotal borders. The male genitalia of this species are distinctive (Fig. 7 B). Description (Holotype female) (Fig. 2 C): Length: 4.6 mm, width 3.3 mm. Form ovoid, weakly convex, narrowly explanate, lateral margin broadly weakly arcuate, apically tapered, somewhat pointed, extreme apex rounded. Punctation on dorsal surfaces shallow, regular, each puncture separated by 1.5–2.5 X its diameter; surface between punctures matt, strongly reticulate; punctures on head, pronotum easily discernable; punctures on elytra more shallow, nearly obsolete, more noticeable along lateral margins. Dorsal color pattern as follows (Fig. 2 C): Head black with two cream­colored spots, one at inner margin of each eye extending from eye canthus to mid level of eye; eye dark with hint of silvery reflections; labrum very dark brown. Pronotum predominantly black; anterior, lateral margins with narrow even cream­colored border (Fig. 3 I), slightly wider in lateral than anterior margin. Elytron predominantly black, with narrow light tan to ferrugineous band along anterior, lateral borders; band abruptly angularly expanded near elytral apex (Fig. 3 G); sutural border much narrower, very dark reddish brown. 5 creamcolored irregularly rounded maculae disposed as follows: one resting on elytral base narrowly separated from scutellar notch; two in row near midline, outer macula just touching ferrugineous lateral band, inner macula closer to suture than to outer macula; one in apical third between sutural, lateral margins. Additional small comma­shaped creamcolored mark, resting on ferrugineous band, positioned between humeral bulge, lateral margin. Outer margins of all cream­colored maculae narrowly ferrugineous. Punctures in pale areas of elytron sometimes distinguished by pinpoint of brown pigmentation particularly noticeable along lateral edge of outer maculae, inner edge of lateral ferrugineous band. Blackish areas on dorsal surfaces appear pruinose, reflecting bluish tint when brightly illuminated. Ground color of ventral surfaces black or very dark brown; elytral epipleuron ocher; lateral ½ of pronotal hypomeron, mesepimeron cream­colored; mouthparts, antenna, tarsi dark reddish brown. Ventral surfaces including appendages clothed in fine decumbent silvery pubescence. Eyes finely facetted, separated by 2 X eye diameter; inner orbits nearly parallel in lower half, diverging at upper level. Antenna of 11 antennomeres (Fig. 9 H), combined length distinctly greater than distance between eyes; third antennomere slightly longer than fourth. Pronotum weakly convex, lateral margin explanate beginining just before transparent border. Pronotal outline with basal margin subsinuate; lateral margin strongly arcuate in basal half, weakly arcuate beyond; anterior margin subtrapezoidally emarginate; medially linear; anterior angles subtriangulate, projecting anteroventrally. Elytron elongate (Figs. 5 I–K), in dorsal view with humeral angle rounded, evenly weakly arcuate for rest of length, broadest at apical 2 / 5, apex rounded, outer margin weakly explanate; epipleuron weakly concave, descending externally. Prosternum T­shaped, with lateral arms of transverse basal piece flat, weakly folded away from midline on each side, weakly convex at middle; intercoxal process nearly flat with median third weakly impressed. Mesosternum trapezoidal; anterior border nearly linear, indistinctly emarginate at middle, with faint raised margin. Metasternum broad, with postmesocoxal line reaching lateral margin; transversely rugulopunctate; discrimen present, distinct except for extreme ends. Abdomen shortened semi­oval, broadest at second ventrite; posterior margin of ventrites 1–4 linear, of 5 approximately linear but slightly wavy, weakly protuberant at middle, 6 rounded; postmetacoxal line of first abdominal ventrite curved posterolaterad, closely paralleling posterior margin for much of length, not attaining lateral margin. Tarsal claw with rectangular basal tooth. Female genitalia (specimen from Molinos, Salta) as in figure 8 G. Male: Similar to female except head black with cream­colored band filling most of space between clypeal margin and level of upper 1 / 3 of eye including canthus; border separating black and cream­colored areas wavy; area near clypeus slightly infuscate. Abdomen with posterior margin of ventrite 5 linear, exposed portion of ventrite 6 rounded with shallow emargination in median third. Male genitalia as in figure 7 B: basal lobe elongate, with greatest width at basal one fourth; tapered beyond with sides slightly sinuate; parameres slightly thick, reaching 4 / 5 distance to apex of basal lobe. Variation: Length 4.2–4.6 mm. Elytral color pattern varies primarily in the width of the tan to ferrugineous band and size of the cream­colored maculae as shown (Figs. 5 I–K). Elytral ground color varies from deep maroon brown to black. Type material: Lectotype of C. boliviana, designated Gordon 1987, “TYPE [blue paper]/TYPE, boliviana, Deyr. ” (UCCC, specimen examined). Remarks: Mulsant (1866) regarded this as a good species, but subsequent authors, until the present, treated it as a synonym or mere aberration of C. areata (= sicardi). Summary of data from specimens examined (Map, Fig. 11): ARGENTINA: SALTA: Molinos, 21.I. 1950 (Monros­Willink), 2 specimens; BOLIVIA: LA PAZ: Sorata, 21.II. 1953 (F. Monros), 1 specimen; Sorata, II 1941 (W. Wittmer), 2 specimens (BMNH, IML, UCCC), Published as part of González, Guillermo & Vandenberg, Natalia J., 2006, Review of lady beetles in the Cycloneda germainii species complex (Coleoptera; Coccinellidae: Coccinellinae: Coccinellini) with descriptions of new and unusual species from Chile and surrounding countries, pp. 13-50 in Zootaxa 1311 on pages 46-48, DOI: 10.5281/zenodo.173868, {"references":["Crotch, G. R. (1874) A revision of the coleopterous family Coccinellidae. University Press, London, 311 pp.","Weise, J. (1910) Chrysomeliden und Coccinelliden, Sonderabdruck aus dem XLVIII. Bande der Verhandlungen des naturforschenden Vereins zu Brunn, 20 - 29.","Gordon, R. D. (1987) A catalogue of the Crotch collection of Coccinellidae (Coleoptera). Occasional Papers on Systematic Entomology, 3, 1 - 46.","Korschefsky, R. (1932) Coleopterorum Catalogus, Pars 120, Coccinellidae II. W. Junk, Berlin, 659 pp.","Blackwelder, R. E. (1945) Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 3. Bulletin of the United States National Museum, 185, 343 - 550."]}

Cycloneda lacrimosa González & Vandenberg, new species (Figs. 1 A; 3 A, H; 4 M–N; 6 A; 8 A; 9 A–C; 10) Diagnosis: Distinguished from other Cycloneda species by the bicolored or very weakly tricolored elytra with a large ocher­colored teardrop­shaped sutural macula, and by the robust convex oblong body form (Figs. 1 A, 3 A). This species appears to be closely related to C. disconsolata, new species, but the latter has the elytron distinctly tricolored and the dark discal figure extended obliquely toward the humeral angle. If females of the two species are compared, C. lacrimosa is distinctly broader. Cycloneda lacrimosa is similar in color and convexity to C. eryngii (Mulsant), but the latter has an ovoid body form, with the lateral elytral margin more evenly arcuate (Fig 2 A), and the elytron bearing a pair of transverse discal maculae that are separated at the midline, or joined only at the inner edges (Figs. 4 A–H). The male genitalia of C. lacrimosa (Fig. 6 A) have the basal lobe more strongly dilated in apical 1 / 4 compared with the other species in this complex (except males not known in C. disconsolata). Description (Holotype male): Length 3.0 mm, width 2.3 mm. Form shortened oblong, nearly parallel­sided, convex, apically rounded; elytral, pronotal margins very narrowly reflexed. Punctation on dorsal surfaces fine, regular, with each puncture separated by 2.0–3.0X its diameter; surface between punctures shiny, reticulate on head, pronotum, with only faint trace of reticulation visible on elytron. Dorsal color pattern as follows: Head black with two irregular cream­colored spots, one at inner margin of each eye extending from eye canthus to just below level of upper 1 / 3 of eye; eye silvery; labrum brown. Pronotum predominantly dark reddish brown, nearly black; lateral margins yellow cream­colored, beginning at posterior angle as narrow band, widened apically to enclose entire anterior angle, continued as broken trace on anterior margin (Fig. 3 H). Scutellum black. Elytron with ground color pale ocherous or straw with dark brown irregular figure filling most of disc (Fig. 4 N); discal figure with incised borders as follows: outer border with anteromedial triangular emargination pointing toward sutural apex; inner border with large teardrop­shaped emargination followed by smaller semicircular emargination, the latter deeply penetrating the dark zone leaving only a slender dark hook; union of dark, light areas irregular, suffused, somewhat reddish. Each puncture of dorsal surfaces with pinpoint of brown at center, visible only in areas with light to medium background coloration; staggered double row of punctures nearest suture, single to double staggered row along lateral margin beginning just outside of humeral bulge with more pronounced pigmentation. Anterior, lateral margins of pronotum, all margins of elytron narrowly transparent to light amber, sutural margin somewhat darker. Ground color of ventral surfaces dark brown; elytral epipleuron, pronotal hypomeron except basally at inner margin straw­colored; mesepimeron cream­colored; antenna, mouthparts yellow brown with antennal club slightly darker; legs with coxa brown, femur dark brown to blackish; tibia yellow brown with darker brown narrowly along outer margin; tarsus translucent yellow brown with last tarsomere, base of claw darker. Ventral surfaces including appendages clothed in decumbent silvery pubescence. Eyes finely facetted, separated by 2 ½X eye diameter; inner orbits nearly parallel in lower half, diverging at upper level. Antenna of 10 antennomeres (Fig. 9 B), combined length slightly greater than distance between eyes; third antennomere elongate, 1 ½X length of second, subequal to four plus five combined. Pronotum (Fig. 3 H) evenly convex except for very narrowly reflexed transparent lateral margin; in outline with basal margin subsinuate, lateral margin strongly evenly arcuate, anterior margin subtrapezoidally emarginate, medially slightly arcuately produced; anterior angles subtriangulate, conspicuous in dorsal view, projecting anteroventrally. Elytron broad (Figs. 4 M–N), in dorsal view with humeral angle rounded, weakly arcuate from beyond humeral angle to apical 1 / 3, broadest near apical 1 / 3, roundly tapered distally; epipleuron flat, horizontal in anterior half, progressively inwardly sloping in posterior half. Prosternum T­shaped, with lateral arms gently folded back from stem, in cross section forming shallow arc; intercoxal process weakly convex, bearing fine superficial median sulcus on distal 1 / 4, apex truncate. Mesosternum trapezoidal; anterior border approximately linear with raised margin. Metasternum broad, with postmesocoxal line reaching lateral margin; surface polished, obsoletely rugulose; discrimen present, indistinct. Abdomen shortened semi­oval, broadest between first, second ventrites; posterior margin of ventrites 1–4 linear, of 5 broadly shallowly emarginate, of 6 apically rounded; postmetacoxal line of first abdominal ventrite curved posterolaterad, closely paralleling posterior margin for much of length, not attaining lateral margin. Tarsal claw with shallow subquadrate basal tooth. Male genitalia as shown (Fig. 6 A): basal lobe elongate, roughly parallel­sided in basal half, subapically distinctly swollen with greatest width at apical 1 / 4; apex tapered, slightly attenuate; parameres slender, reaching three quarters distance to apex of basal lobe. Female: Similar to male except slightly larger on average and proportionally broader, apex slightly pointed. Abdomen with posterior margin of ventrite 5 nearly linear, apex of ventrite 6 pointed. Female genitalia as in figure 8 A. Variation: Length 3.0 to 3.6 mm. Pale markings on head variable in size but always well separated by dark frons; not gender­specific. Pale anterior margin of pronotum obsolete to narrow but entire. Elytron may have additional dark spot at apex (Fig. 1 A), or faint dark spot at humeral bulge; sometimes with hook shaped mark at apex of dark discal figure disconnected (Fig. 4 M) or entirely absent; some specimens with suffused creamcolored maculae nested within 3 incised areas of dark discal figure; more commonly with ground color only slightly lighter on disc or apparently unicolorous. Dark areas on dorsal surfaces nearly black in many specimens. Pronotal hypomeron unicolorous or basally darkened as in holotype. Tibia totally yellowish in some. Antenna may have 9, 10 or 11 antennomeres (Figs. 9 A–C). Sulcus on prosternal intercoxal process may be less distinct or wavering. Type material: Holotype (male) “ Agua Verde, Antofagasta, Chile, 25 ­dic­ 1986. G. González F.”(MNHN); Allotype (female), “ CHILE, Antofagasta, Agua Verde. 21 ­ diciembre­ 1991, leg. G. Gonzalez. ” (MNHN); Paratypes (total= 24), 1 with same data as holotype (MNHN), 18 with same data as allotype (4, AMNH; 3, CAS; 7, MNHN; 4, USNM), 4 “ Oruro, Bolivia, 14.1. 40 3700 m, W. Wittmer /Brit.Mus., 1945 ­ 33.[one paratype mounted along with an incomplete specimen, the latter not designated as a paratype] (BMNH), 1 “ ARGENTINA, SALTA, Nevado de Cachi, 5200 m _ 6m, 6.I­ 1973, Col: Stephan Halloy /COLECCION, INST.– FUND M. LILLO (4000) – S.M.TUCUMAN, TUCUMAN – ARGENTINA”(IML) Etymology: From the Latin lacrimosus (adj.) meaning “prone to tears or crying,” the name is a reference to the large teardrop­shaped mark at the elytral suture. Remarks: The specimens from Agua Verde were collected at an isolated gas station in the Atacama Desert, on plants in flower­pots, about 800 kilometers north of the northern most locality for C. germainii. From this single collection locality specimens were obtained with either nine, ten, or eleven antennomeres. These variants are assumed to be conspecific as they show no other apparent differences. Figure 9 A–C suggests one possible interpretation for the observed variation based on fusion in the region between the basal two and distal six antennomeres. This hypothesis is supported by a corresponding increase in the length of antennomere four in specimens with only ten antennomeres, or antennomere three in specimens with only nine antennomeres. Antennal polymorphism was previously reported in the coccinellid species Catana clauseni Chapin (Serangiini). Chapin (1940) distinguished Catana Chapin from other members of the tribe by the possession of only eight antennomeres, but he noted that the fifth antennomere sometimes shows the beginning of a division and is therefore morphologically equivalent to the fourth and fifth combined. Miyatake (1961) subsequently discovered a specimen of C. clauseni with a complete division resulting in nine distinct antennomeres as in the related genus Serangium. It is probable that Weise was familiar with the new species described here, but regarded it as a variation of C. eryngii. The Smithsonian Entomology Library has a copy of Crotch (1874) that was signed by Weise and apparently extensively annotated by him. In the left hand margin of page 107, below the caption for Coccinella eryngii, is a handpenned figure (left half of pronotum, left elytron) of a specimen with a pattern identical to the example illustrated in figure 4 N. The sketch is labeled with the female symbol and the words “var. Moreno, Argentin. Mus. [illegible word]”. Data from specimens examined (Map, Fig. 10): see “ Type material,” above., Published as part of González, Guillermo & Vandenberg, Natalia J., 2006, Review of lady beetles in the Cycloneda germainii species complex (Coleoptera; Coccinellidae: Coccinellinae: Coccinellini) with descriptions of new and unusual species from Chile and surrounding countries, pp. 13-50 in Zootaxa 1311 on pages 20-23, DOI: 10.5281/zenodo.173868, {"references":["Chapin, E. A. (1940) New genera and species of lady-beetles related to Serangium Blackburn (Coleoptera: Coccinellidae). Journal of the Washington Academy of Sciences, 30, 263 - 272.","Miyatake, M. (1961) The East-Asian coccinellid-beetles preserved in the California Academy of Sciences, tribe Serangiini. Memoirs of the Ehime University, Sect. VI (Agriculture), 6 (2), 135 - 146.","Crotch, G. R. (1874) A revision of the coleopterous family Coccinellidae. University Press, London, 311 pp."]}

Cycloneda disconsolata Vandenberg & González, new species (Figs. 1 B; 3 B; 4 O–P; 8 B; 10) Diagnosis: Distinguished from other Cycloneda species by tricolored elytra with a pair of cream­colored teardrop­shaped marks, one on each side of the suture (Fig. 1 B), convex oblong body form (Fig. 3 B), and antenna composed of 10 antennomeres. This species appears to be closely related to C. lacrimosa, but, unlike the preceding species, the suture is ferrugineous and distinctly darker than the pale elytral maculae (Figs. 4 O–P). Cycloneda disconsolata can be distinguished from all other species except C. lacrimosa by the teardrop­shaped mark near the elytral suture. Description (Holotype female): Length 3.3 mm, width 2.3 mm. Form shortened oblong, nearly parallel­sided, convex, apically pointed; elytral, pronotal margins very narrowly reflexed. Punctation on dorsal surfaces fine, regular, with each puncture separated by 2.0–3.0X its diameter; surface between punctures shiny, reticulate on head, pronotum, with only faint trace of reticulation visible on elytron. Dorsal color pattern as follows: Head black with two irregular cream­colored spots, one at inner margin of each eye extending from eye canthus to just below level of upper 1 / 3 of eye; eye dark with hint of silvery reflections; labrum dark brown. Pronotum predominantly black; lateral margins narrowly cream­colored, widened apically to enclose entire anterior angle, terminating opposite inner orbit of eye, faintly suggested beyond by cream­colored stippling. Scutellum black. Elytron with deeply incised irregular black figure against cream­colored background as shown (Fig. 1 B, 4 P); figure offset from lateral margin by cream­colored band about equal to scutellar width, offset from sutural margin by equally broad ferrugineous band; black figure with incised borders as follows: base with semicircular emargination; outer border with anteromedial trapezoidal emargination, apical 1 / 4 with deep spatulate incision beginning at outer border, nearly attaining ferrugineous sutural band; inner border with large teardrop­shaped emargination near midline; union of dark, light areas irregular, narrowly ferrugineous, strongly suffused at elytral apex, humeral angle. Each puncture of dorsal surfaces with pinpoint of brown at center, visible only in areas with light to medium background coloration; staggered double row of punctures nearest suture with more pronounced pigmentation; scattering of similar punctures visible on margins of cream­colored maculae near suture. Anterior, lateral margins of pronotum, all margins of elytron narrowly transparent, pale amber on outer elytral margins, dark reddish amber on sutural margin. Ground color of ventral surfaces dark reddish brown, nearly black; elytral epipleuron, pronotal hypomeron except basal dark spot at inner margin cream­colored; mesepimeron cream­colored; antenna, mouthparts amber brown with first, last two antennomeres dark brown; legs nearly black with all tarsi, protibia, apex of meso­, metatibia slightly lighter. Ventral surfaces including appendages clothed in decumbent silvery white pubescence. Eyes finely facetted separated by 2 ½X eye diameter; inner orbits nearly parallel in lower half, diverging at upper level. Antenna of 10 antennomeres, combined length slightly greater than distance between eyes; third antennomere elongate, 1 ½X length of second, subequal to four plus five combined. Pronotum evenly convex except for very narrowly reflexed transparent lateral margin; in outline with basal margin subsinuate, lateral margin strongly evenly arcuate, anterior margin subtrapezoidally emarginate, medially slightly arcuately produced; anterior angles subtriangulate, in dorsal view obscured by curvature of pronotum with only extreme apex visible, projecting ventrally, slightly anteriorly. Elytron broad (Figs. 4 O–P), in dorsal view with humeral angle rounded, weakly arcuate from beyond humeral angle to apical two­fifths, broadest just beyond middle, almost linearly tapered distally, rounded at extreme apex; epipleuron weakly concave, horizontal in anterior half, progressively inwardly sloping in posterior half. Prosternum T­shaped, with lateral arms gently folded back from stem, in cross section forming shallow arc; intercoxal process weakly convex, shallowly triangularly impressed along midline in distal 1 / 4 th. Mesosternum trapezoidal; anterior border approximately linear, with raised margin. Metasternum broad, with postmesocoxal line reaching lateral margin; surface polished with faint transverse rugulae, more apparent near midline; discrimen present, indistinct. Abdomen shortened semi­oval, broadest between first, second ventrites; posterior margin of ventrites 1–4 linear, of 5 weakly arcuate; exposed portion of 6 subtriangular; postmetacoxal line of first abdominal ventrite curved posterolaterad, closely paralleling posterior margin for much of length, not attaining lateral margin. Tarsal claw with shallow subquadrate basal tooth. Genitalia as in figure 8 B. Male: Unknown. Variation: Length 2.9 to 3.3 mm. Pale anterior margin of pronotum obsolete to narrow but entire; one specimen with pair of linear cream­colored marks enclosed within dark pronotal disc, situated one on each side at anterior 1 / 3, equidistant from lateral margin and midline. Elytron with pale marks as in holotype or with cream­colored markings more extensive, with anterior mark joined to teardrop­shaped mark as shown (Fig. 4 O). Dark coloration on pronotum, elytron varies from blackish to dark reddish brown. Venter blackish to medium brown. Tarsi dark brown to light yellow brown or amber; pro­, mesotibia, apex of metatibia yellow brown in one specimen. Type material: Holotype (female), “ 37 [round label]/Lupica, Parinacota, [Tarapaca,] Chile, 3 2000 m.s.n.m., Mayo 20, 1982 /En paja brava, D. Bobadilla coll./ Coccinellina eryngii? (Mulsant) det. R.Gordon 87 " (INIA); Paratypes (total= 2 females), 1 “ Valcheta, [Rio Negro,] Arg./No. 392, So Amer, Montevideo, Paras Lab, Date 1­20 ­ 43 Host ” (USNM), 1 “RIO SECO, Cord: Arica, 18 ­Nov­ 1952, Coll: L.E.Pena ” (USNM). Etymology: From the Latin disconsolatus (L. dis ­ + consolatus, p. p. of consolari to console), meaning “incapable of being consoled, filled with grief, hopelessly sad,” the name is a reference to the pair of teardrop­shaped marks, one on each side of the elytral suture. Remarks: The antenna of this species appears to have a similar shape and proportions to examples of C. lacrimosa with 10 antennomeres (Fig. 9 B). The antenna of C. disconsolata was not dissected and slide mounted due to the paucity of material available, and the desire to keep the type specimens intact. Data from specimens examined (Map, Fig. 10): see “ Type material,” above., Published as part of González, Guillermo & Vandenberg, Natalia J., 2006, Review of lady beetles in the Cycloneda germainii species complex (Coleoptera; Coccinellidae: Coccinellinae: Coccinellini) with descriptions of new and unusual species from Chile and surrounding countries, pp. 13-50 in Zootaxa 1311 on pages 23-25, DOI: 10.5281/zenodo.173868

Cycloneda eryngii (Mulsant) new combination (Figs. 2 A; 3 E; 4 A–H; 6 D; 8 E; 9 F; 11) Coccinella eryngii Mulsant 1850: 100, 1866: 83; Crotch 1874: 107; Philippi 1887: 173; Brèthes 1921: 454; Brèthes 1925: 152; Korschefsky 1932: 510; Blackwelder 1945: 454. Coccinellina eryngii: Timberlake 1943: 15; Aguilera 1995: 99; Vandenberg 2002: 226 (transfer to Cycloneda). Coccinellina eringii: Aguilera 1995: 99 (typo). Coccinella interrupta Germain 1854: 334; Philippi 1887: 173; Brèthes 1921: 454 (As synonym of C. eryngii); Blackwelder 1945: 454. Diagnosis: Distinguished from other members of the genus by the combination of antenna composed of 11 antennomeres (Fig. 9 F) and elytron with a pair of dark transverse fasciae which lack a median longitudinal connection (Figs. 4 A–F), but may be connected near their inner edges (Fig. 4 G). The elytral color patterns found in this species distinguish it from all others except some uncommon examples of C. germainii (4 I, L) that lack a median longitudinal connection between the dark elytral maculae. In such examples, the antenna with 10 antennomeres and more parallel­sided elytral base will identify these deceptively similar color morphs. In C. eryngii, the basal lobe of the male genitalia is more slender and apically simpler than in the other species (Fig. 6 D). Description (male from El Melocotón): Length 3.3 mm. Form ovoid, convex, feebly explanate, lateral margin evenly arcuate, apically tapered, somewhat pointed, extreme apex rounded; elytral, pronotal margins narrowly weakly reflexed. Punctation on dorsal surfaces shallow, regular, with each puncture separated by 3.0–4.0X its diameter; surface between punctures weakly shiny, reticulate. Dorsal color pattern as follows: Head black with cream­colored figure filling most of space between clypeal margin and middle level of eye including canthus, resembling an elongated “M” with sloping sides; two small dark triangles at clypeal margin form cut away lower border of figure. Pronotum predominantly black; anterior, lateral margins with even cream­colored band of little more than ½ diameter of eye. Scutellum black. Elytron with ground color orange yellow with pair of irregular black transverse fasciae slightly offset from sutural, lateral margins (Fig. 4 D); anterior fascia occupying basal half, posterior fascia occupying apical 1 / 3; fasciae broader near suture, constricted in lateral third. Each puncture of dorsal surfaces with pinpoint of brown at center; double staggered row of punctures nearest suture, single to double staggered row along lateral margin beginning just outside of humeral bulge with more pronounced pigmentation. Anterior, lateral margins of pronotum narrowly transparent to light amber; all margins of elytron darker orangy amber, darkest on outer margin toward apex. Ground color of ventral surfaces black; elytral epipleuron pale yellow orange; outer half of pronotal hypomeron, mesepimeron cream­colored; mouthparts, antenna, brownish, darkened toward apices; posterior, lateral margins of last abdominal segments dark reddish brown; leg black, with protibia, apex of meso­, metatibiae brown; all tarsi amber brown. Ventral surfaces including appendages clothed in decumbent to semi­erect pubescence of variable length, greyish white to golden in color. Eyes finely facetted, separated by 2 X eye diameter; inner orbits nearly parallel in lower half, diverging at upper level. Antenna of 11 antennomeres (Fig. 9 F), combined length equal to distance between eyes; third antennomere elongate, about 1 ¼ length of second, subequal to fourth plus fifth combined. Pronotum convex, lateral margin weakly reflexed beginning just before transparent border; base of pronotum weakly explanate beginning at middle of cream­colored border; pronotal outline with basal margin subsinuate, lateral margin strongly evenly arcuate, anterior margin subtrapezoidally emarginate, medially slightly arcuately produced; anterior angles subtriangulate, projecting anteroventrally. Elytron broad (Figs. 2 A, 4 A–H), in dorsal view with humeral angle rounded, evenly arcuate for rest of length, broadest near middle or just beyond, apex rounded; with outer margin weakly explanate; in lateral view (Fig. 3 E) evenly arcuate dorsally, somewhat wedge shaped, broader at anterior 1 / 4 than at posterior 1 / 4; epipleuron approximately horizontal, weakly concave in medial half. Prosternum T­shaped, with lateral arms of transverse basal piece flat, folded away from midline on each side, rounded off at middle, in cross section forming a broad angle with apex blunt; intercoxal process weakly convex with narrow median sulcus along most of length. Mesosternum trapezoidal; anterior border approximately linear, with raised margin. Metasternum broad, with postmesocoxal line reaching lateral margin; shallowly transversely rugulose; discrimen present, distinct except for extreme ends. Abdomen shortened semi­oval, broadest at apex of first ventrite; posterior margin of ventrites 1–4 linear, of 5 arcuately emarginate, 6 rounded at sides, arcuately emarginate in middle third; postmetacoxal line of first abdominal ventrite curved posterolaterad, closely paralleling posterior margin for much of length, not attaining lateral margin. Tarsal claw with rectangular basal tooth. Male genitalia as shown (Fig. 6 D): basal lobe elongate, slender, roughly parallel­sided in basal half, very slightly swollen at apical one third, tapered beyond; width at base of basal lobe slightly exceeding width at apical one third; parameres slender, reaching three quarters distance to apex of basal lobe. Female: Similar to male except larger on average; head black with irregular creamcolored patch on lower half adjacent to each eye, including eye canthus. Abdomen with posterior margin of ventrite 5 linear or slightly wavy, exposed portion of ventrite 6 short, apically rounded. Female genitalia as in figure 8 E. Variation: Length 3.2–4.6 mm. Elytral color pattern varies as shown (Figs. 4 A–H). Elytral ground color apparently unicolorous or gradually lightened from outer edge to disc; sometimes faintly suggesting one or more of the cream­colored markings found in C. germanii (e.g. oblique oval subapical mark) but less well defined; transition more abrupt in specimens from Baños de Cauquenes (Fig. 4 H) which have a paler yellow cream elytron with an orange band at sutural and lateral borders. The latter morph also occurs in specimens marked “ Chile ” without additional locality data. Some lightly marked northern specimens possess a pair of linear cream­colored marks enclosed within the dark pronotal disc, situated one on each side at anterior 1 / 3, equidistant from lateral margin and midline. Type material: Type material of C. eryngii, lost?, not in MHNL (location suggested in Gordon 1987); holotype of C. interrupta, “Prov. Valparaiso / Holotipo No. 2159 ” (MNHN) (specimen examined). Prey species: Aphididae (Hemiptera): Metopolophium dirhodum (Walker), Schizaphis graminum (Rondani), Sitobion avenae (F.), Acyrthosiphon pisum (Harris), A. kondoi Shinji, Aphis gossypii Glover, A. craccivora Koch, Macrosiphum euphorbiae (Thomas), and Uroleucon ambrosiae (Thomas) (data taken from Aguilera 1995). Remarks: Bréthes placed Coccinella interrupta as a synonym of C. eryngii based on similarities in the type descriptions, and that placement is followed here. Coccinella chilena (Weise) (MNHUB, type examined) and C. limbicollis Fairmaire (MNHP, type examined) are recognized here as distinct from C. eryngii. They were formerly treated as varieties of C. eryngii by Korschefsky (1932), and as varieties of Coccinella interrupta Germain by Blackwelder (1945). The unusual tricolored morph of C. eryngii (see figure 4 H) is tentatively included here based on the antenna with 11 antennomeres, the elytron somewhat wedgeshaped in profile, with lateral margin distinctly arcuate in dorsal view, and the male genitalia with a slender basal lobe that is broadest at base (single example dissected). Summary of data from specimens examined (Map, Fig. 11): CHILE: ANTOFAGASTA: Chiu­Chiu, 28.IX. 1986 (G. González F.), 1 specimen; Calama, Ojo Opache, 3.X. 1982 (G. Arriagada), 4 specimens; Prov. [Provincia] El Loa, Calama, 31.I. 1993 (G. González F), 1 specimen; Calama, Asen. Pedro A. Cerda, en alfalfa, 13.I. 1972 (R. Mendoza), 1 specimen; Rio Loa, Calama, 8.II. 1987 (G. González F.), 3 specimens; San Pedro de Atacama, 2500 m [m.s.n.m], 16.XI. 1946 (G. Kuschel), 1 specimen; San Pedro de Atacama, en alfalfa, 14.I. 1972 (R. Mendoza), 1 specimen; San Pedro de Atacama, en alfalfa, 7.XII. 1986 (G. González F.), 2 specimens; San Pedro de Atacama, en alfalfa, 29.IX. 1986 (G. González F.), 2 specimens; Toconao, 18.IV. 1946 (Kuschel), 1 specimen; Toconao, 17.XI.1986, 1 specimen. ATACAMA: El Salvador, V. 1982 (G. González F.), 2 specimens; Copiapó, 19.VIII. 1940 (P. A. Berry), 3 specimens; Copiapó, 25.XI. 1944 (M. Marió), 2 specimens; Cachiyuyo, 19.X. 1966 (R. Wagenknecht), 1 specimen; Cachiyuyo, 29.XI.1966, 1 specimen; Chañaral, 4.I. 1987 (G. González F.), 3 specimens; Bahía Inglesa, 8.IV.1991, 1 specimen. COQUIMBO: Chapilca Huanta, km 12, 1960 m.s.n.m., en Chilla, 11.XII. 1975 (A. Aguilera P.), 1 specimen; 20 miles E of La Serena, 3.XII. 1950 (Ross and Michelbacher), 4 specimens; La Serena, on Baccharis, 9.XII. 1950 (Ross and Michelbacher), 1 specimen; Rivadavia, Elqui, 950 m. s.n.m., 25.VIII. 1973 (H. Vasquez C.), 6 specimens; 5 miles N of Laguna Dam, 8000 feet, 6.XII.1950, 1 specimen; Paihuano, II.1983, 1 specimen; 12 miles E Vicuña, 4.XII. 1950 (Ross and Michelbacher), 1 specimen; Pisco Elqui, Elqui, 1300 m.s.n.m., 23.IX. 1973 (H. Vasquez), 1 specimen; Pangue, 19.X. 1957 (G. Kuschel), 1 specimen; 5 miles SW of Ovalle, 12.XII. 1950 (Ross and Michelbacher), 1 specimen; 10 km E Fray Jorge National Park, dry wash, 28.XII. 1966 (M. E. Irwin), 1 specimen; 30 km N Illapel, 5000 foot elevation, 30.XI. 1950 (Ross and Michelbacher), 2 specimens; 5 miles N of Illapel, 30.XI. 1950 (Ross and Michelbacher), 1 specimen; Hacienda Illapel, Rio Illapel, 600–900 m [m.s.n.m], 19.X. 1966 (E. I. Schlinger, M. E. Irwin) 1 specimen; Salamanca, 15.II. 79 (A. Gaete), en malezas, 1 specimen; 5 miles W of La Junta, 7.XII. 1950 (Ross and Michelbacher), 4 specimens; 10 miles W of La Junta, 7.XII. 1950 (Ross and Michelbacher), 6 specimens. VALPARAISO: Las Palmas, Ocoa, 10.XI. 1956 (N. Hichins), 1 specimen; E entrance to tunnel, Aconcagua, 90 km S Illapel, 28.XI. 1950 (Ross and Michelbacher), 3 specimens; [La] Ligua, IX. 1997 (Germain), 1 specimen; Los Andes, Aconcagua, en Romero, 14.II. 1975 (R. Ripa), 2 specimens; Los Andes, Santiago, 6.XII. 1944 (gift from G. Olalquiaga F.), 4 specimens; Quillota, XI. 1894 (Germain), 1 specimen; Cerro las Vizcaches [Las Vizcachas], 1840 m [m.s.n.m], 7.XII. 1951 (P. C. Hutchison), 1 specimen; Quebrada de Alvarado, 27.I. 1959 (N. Hichins), 5 specimens. METROPOLITAN REGION: Guardia Vieja, 8.XII. 2000 (M. Diéguez), 1 specimen; Farellones, Cord [Cordillera] Stgo [Santiago], 22.XI.1985, 1 specimen; Santiago, El Arrayan, 1.V.46, 3 specimens; Santiago, La Ermita, 16.XI. 1985 (G. González F.), 1 specimen; Santiago, La Reina, Precordillera, 5.X. 80 (G.González F), 1 specimen; Santiago, 30.IX. 1945 (P. G. Kuschel), 1 specimen; Santiago, 21.X. 1945 (P. G. Kuschel), 1 specimen; Santiago, Peñalolen, III. 1940 (R. Gutiérrez), 5 specimens; Santiago Prov., Que [Quebrada] La Plata 510 m [m.s.n.m], [La] Rinconada, Maipú, Malaise, 33 ° 31 'S 70 ° 47 'W, 1.I. 1967 (L. A. Stange), 1 specimen; Santiago Prov., El Canelo, on path near water, 33 ° 35 'S 70 ° 27 'W, 10.I. 1967 (E. I. Schlinger), 1 specimen; Stgo [Santiago], [El] Canelo, IX.1950, 1 specimen; Santiago, El Canelo, 18.X. 1951 (Hofmann), 4 specimens; Santiago, El Canelo, 15.XII.1954, 8 specimens; Santiago, S[an] Bernardo, I. 1940 (R. Gutiérrez), 1 specimen; Stgo [Santiago], Qbda [Quebrada] Macul, 3.IV.1913,1 specimen; Stgo [Santiago], Cajón del Maipo, El Melocotón, I. 1977 (G. González F.), 6 specimens; Stgo [Santiago], Cajón del Maipo, El Melocotón, 30.III. 1986 (G. González F.), 1 specimen; Lo Valdes, 800 m [m.s.n.m], 9.I. 1945 (Kuschel), 2 specimens. LIBERTADOR: Fundo Romeral, S. [San] Franscisco de Mostazal, XII. 1933 (Rafael Barros), 1 specimen; B. [Baños] de Cauquenes, Rancagua, 3 specimens. MAULE: Maule Prov., Rio Teno, 800 m [m.s.n.m], ca. 40 km E Curicó, 25–27.XI. 1981 (D. R. Davis), 1 specimen; Curicó, Los Queñes, VIII. 1944 (Monsalvez); Curicó, Los Queñes, 3.I. 1988 (Sergio Roitman), 3 specimens; Maule Pr., Forel Carrizalillo, 250 m [m.s.n.m], 30.I.– 5.II. 1981 (L. E. Pena), 1 specimen; Talca, R.N. Altos de Lircay, 6–7.I. 2001 (M. Diéguez), 3 specimens; BIO­BIO: Cord [Cordillera] Chillán, 1899 (Germain), 4 specimens; Ñuble, 15 Km E. Recinto, 31.I. 1968 (C.W.O’Brien), 1 specimen; Ñuble, NW Recinto, 1.XI. 1967 (C.W.O’Brien), 1 specimen; El Abanico, 30.XII. 1950 (Ross and Michelbacher), 1 specimen; Abanico, 800 m [m.s.n.m], 8.I. 1948 (Kuschel), 1 specimen; Los Ángeles, Huaqui, 22.I. 1944 (G. Kuschel), 9 specimens; Pemehue, altitude 1300 m [m.s.n.m], 14.I. 1946 (P. G. Kuschel), 1 specimen. ARAUCANÍA: Angol, 7.II. 1924 (D. S. Bullock), 1 specimen; Pemehue, 1894 (Germain), 6 specimens. LOS LAGOS: Valdivia, 4.III. 1945 (E. A. Chapin), 2 specimens; 8 miles E of Rio Bueno, Valdivia, 15.I. 1951 (Ross and Michelbacher), 1 specimen; Osorno, 2.III. 1945 (E. A. Chapin), 1 specimen; Llanquihue, Fresia, 7.II.1945, 1 specimen. (AAPC, AMNH, BMNH, GGPC, MNHN), Published as part of González, Guillermo & Vandenberg, Natalia J., 2006, Review of lady beetles in the Cycloneda germainii species complex (Coleoptera; Coccinellidae: Coccinellinae: Coccinellini) with descriptions of new and unusual species from Chile and surrounding countries, pp. 13-50 in Zootaxa 1311 on pages 36-41, DOI: 10.5281/zenodo.173868, {"references":["Mulsant, E. (1850) Species des coleopteres trimeres securipalpes. Annales des Sciences Physiques et Naturelles, d'Agriculture et d'Industrie, Lyon, ser. 2, vol 2, 1 - 1104.","Crotch, G. R. (1874) A revision of the coleopterous family Coccinellidae. University Press, London, 311 pp.","Brethes, J. (1925) Sur une collection de Coccinellidae (et un Phalacridae) du British Museum. Anales del Museo Nacional de Historia Natural, 33, 145 - 175.","Korschefsky, R. (1932) Coleopterorum Catalogus, Pars 120, Coccinellidae II. W. Junk, Berlin, 659 pp.","Blackwelder, R. E. (1945) Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 3. Bulletin of the United States National Museum, 185, 343 - 550.","Timberlake, P. H. (1943) The Coccinellidae or ladybeetles of the Koebele collection. The Hawaiian Planters' Record, 47 (1), 1 - 67.","Aguilera A. (1995) Contribucion al conocimiento de Coccinellina eryngii (Mulsant) (Coleoptera: Coccinellidae) en Chile. Acta Entomologica Chilena, 19, 99 - 104.","Vandenberg, N. J. (2002) The new world genus Cycloneda Crotch (Coleoptera: Coccinellidae: Coccinellini): Historical review, new diagnosis, new generic and specific synonyms, and an improved key to North American species. Proceedings of the Entomological Society of Washington, 104 (1), 221 - 236.","Gordon, R. D. (1987) A catalogue of the Crotch collection of Coccinellidae (Coleoptera). Occasional Papers on Systematic Entomology, 3, 1 - 46."]}

Cycloneda patagonica González & Vandenberg, new species (Figs. 1 C; 3 C; 5 A–D; 6 B; 8 C; 9 D; 10) Diagnosis: Distinguished from other Cycloneda species by the elliptical, somewhat depressed body form (Figs. 1 C, 3 C), narrow parallel­sided elytron with close deep punctation (separated by 1.0– 2.5 diameters) (Figs 5 A–D), epipleuron easily visible in lateral view (Fig 3 C), and short antenna, composed of 10 antennomeres with an abrupt club (Fig. 9 D). This species appears to be most closely related to C. germainii, and is most likely to be confused with the “ duplaris ” form (Fig. 4 K), which has very similar elytral maculation. It differs from the latter in the characters given above, and in the male genitalia, particularly the shape of the basal lobe which is shorter and apically less attenuate and more obtusely rounded. Description (Holotype male): Length 2.9 mm, width 1.7 mm. Form elliptical, parallel­sided, somewhat depressed (Fig. 3 C), apically rounded; elytral, pronotal margins very narrowly reflexed. Punctation on dorsal surfaces deep, close, with each puncture separated by 1.0– 2.5 X its diameter; surface between punctures shiny, strongly reticulate on head, pronotum, moderately reticulate on elytron. Dorsal color pattern as follows: Head black with two cream­colored spots, one at inner margin of each eye extending from eye canthus to just beyond level of upper 1 / 2 of eye; eye dark with hint of silvery reflections; labrum dark brown. Pronotum predominantly black; anterior, lateral margins with narrow variable cream­colored border, about 1 / 20 width of pronotum, narrower in anterior margin; anterior border medially infuscate; lateral border infuscate to interrupted on each side near basal 1 / 3, coincident with region of maximum pronotal width. Elytron predominantly black, with ocher­colored circumferential band, four maculae as shown (Fig. 1 C), all but basal macula slightly lighter than band; band narrow, even, entire, about 1 / 10 elytral width, apex fuscus. Elytral maculae arising from, broadly confluent with band, disposed as follows: irregular semicircular basal macula occupying median 2 / 5 of base, penetrating to basal 1 / 10; oblique irregular quadrangular macula near outer margin at middle of length; subapical irregular semicircular macula positioned in posterolateral 1 / 4; smaller parasutural circular macula just behind mid elytral length; union of dark, light areas somewhat irregular, suffused, reddish to ferrugineous. Each puncture of dorsal surfaces with pinpoint of brown at center, visible only in areas with light to medium background coloration; single to double staggered row of punctures at inner margin of sutural band, single to double staggered row along lateral margin beginning just outside of humeral bulge with more pronounced pigmentation. Anterior, lateral margins of pronotum narrowly yellowish amber, gradually darkening to onyx in posterior half of lateral margin; all margins of elytron narrowly amber, darker near apex of lateral margin, sutural margin more reddish amber. Ground color of ventral surfaces black; elytral epipleuron, triangular anterolateral half of pronotal hypomeron ocher; mesepimeron cream­colored; mouthparts amber brown; antenna with basal 2 antennomeres brown, remainder missing from specimen (see variation section below); leg black, with coxa dark reddish brown, tarsus brown. Ventral surfaces including appendages clothed in decumbent greyish white pubescence; hairs fine, long. Eyes finely facetted, separated by slightly more than 2 X eye diameter; inner orbits diverging toward top of head. Antenna broken (see variation below). Pronotum evenly convex except for very narrowly transparent lateral margin; margin distinctly reflexed in anterior half, not or weakly so in posterior half. Pronotum in outline with basal margin strongly arcuate in median half, flatter externally; lateral margin arcuate; anterior margin weakly emarginate, medially arcuately produced nearly as far as anterior angles; anterior angles subtriangulate, in dorsal view obscured by curvature of pronotum. Elytron narrow (Figs. 5 A–D), in dorsal view with humeral angle abruptly rounded, nearly linear from beyond humeral angle to apical two­fifths, arcuately tapered distally, rounded at extreme apex; epipleuron flat, ascending externally, visible in lateral view (Fig. 3 C). Prosternum T­shaped, with lateral arms strongly folded back from stem, in cross section abruptly raised at middle, not forming a simple arc; intercoxal process strongly convex with fine superficial median sulcus along most of length. Mesosternum elongate trapezoidal; anterior border approximately linear, with raised margin. Metasternum broad, with postmesocoxal line reaching lateral margin; surface transversely rugulose, discrimen shallow, somewhat obscured by rugulae. Abdomen elongate semi­oval, broadest in apical half of first ventrite; posterior margin of ventrites 1–4 linear, of 5 weakly arcuate; exposed portion of 6 spindle­shaped; postmetacoxal line of first abdominal ventrite curved posterolaterad, closely paralleling posterior margin for much of length, not attaining lateral margin. Tarsal claw with shallow subquadrate basal tooth. Male genitalia as shown (Fig. 6 B): basal lobe elongate, lateral margin weakly sinuate, roughly parallel­sided in basal two thirds, subapically slightly swollen with greatest width at apical one third; apex tapered, slightly attenuate; parameres slender, reaching three quarters distance to apex of basal lobe. Female: Similar to male except larger on average. Abdomen with posterior margin of ventrite 5 nearly linear, apex of ventrite 6 rounded. Female genitalia as in figure 8 C. Variation: Length 2.8 mm to 4.0 mm. Antenna very short (Fig. 9 D), length equal to distance between eyes, composed of 10 antennomeres; with third antennomere short, subequal to fourth; strongly clubbed. Some specimens with pale anterior, lateral margins of pronotum narrow but entire, forming an even band, or with portions of either or both infuscate, broken or obsolete. Elytral spots larger or smaller than in holotype (Figs. 5 A–D); in some specimens one or more of 3 posterior spots entirely surrounded by black; anterior spot same color as suture, paler cream as in remaining elytral spots, or of an intermediate color; ocher­colored circumferential band may be reddish at apex or absent. Legs may be entirely blackish; pronotal hypomeron with more or less extensive pale area; elytral epipleuron unicolorous or with apical 1 / 5 blackened. Type material: Holotype (male) “ Chile, I. Navarino, Pto. Williams, 1.2.57 / Coll. Kuschel ” (MNHN); Allotype (female) “ Magellanes, Canal Beagle, Isla Navarino / Pto. Williams, Feb. 1.1957, T.Cekalovik /ChpnSlide 58­369 [female genitalia and abdomen on separate slide mount]”(USNM); Paratypes (total= 10), 1 same data as Holotype but missing collector label, and with additional label “ Coccinella germaini Crotch,” and 1 “ Pto Williams, 1 feb 1962, Cekalovic Coll/Coleccion T. Cekalovic 1972 ”(MNHN), 1 “ Tierra del Fuego. Nose Peak, 14 –19.1.05, R.Crawshay. 1906 – 230 ” (BMNH), 3 “ S.America: Tierra del Fuego. Estancia Viamonte. P.W.Reynolds. B.M. 1931 – 273. XII. 1929 ” (BMNH), 1 “ Magellanes, Canal Beagle, Isla Navarino / Pto. Williams, Feb. 1.1957, T.Cekalovik ” (USNM), 1 same as preceding except with additional label “ Coccinellina sicardi (Brèthes), det Chpn 1958 ” (USNM), 1 “ CHILE, prov. Magalle[a]nes, 20 km E. Pto Percy arenales, 10 dic 1998, leg. J.E. Barriga /Coleccion J.E. BARRIGA CHILE 111686 ” (USNM), 1 “ Tierra del Fuego. Useless Bay, Dec. 1904, R.Crawshay, 1906 – 230 / 573 [black underline] FCC­ 683 " (BMNH) Etymology: From the collection locality of the specimens examined (Patagon + ica, adj., meaning "of Patagonia"). The name “Patagonia” was bestowed to the region by Magellan, and refers to that portion of South America which, to the east of the Andes, lies south of the Neuquén and Río Colorado rivers, and, to the west of the Andes, south of 42 °S. Remarks: This new species bears the distinction of being the most austral of any coccinelline reported. It is also, thus far, the most slender and darkly pigmented species in the genus Cycloneda. As with C. lacrimosa, the color pattern on the head is not genderspecific, and resembles the female color pattern found in the related C. germainii. Data from specimens examined (Map, Fig. 10): see “ Type material,” above., Published as part of González, Guillermo & Vandenberg, Natalia J., 2006, Review of lady beetles in the Cycloneda germainii species complex (Coleoptera; Coccinellidae: Coccinellinae: Coccinellini) with descriptions of new and unusual species from Chile and surrounding countries, pp. 13-50 in Zootaxa 1311 on pages 25-30, DOI: 10.5281/zenodo.173868

During an examination of type material of the New World endomychid genus Bystus Guérin-Méneville (Anamorphinae), the type series of Alexia hirtula Kirsch from Peru was found to contain a mixture of different taxa, none of which belong to the genus Bystus, the subfamily Anamorphinae, or even the family Endomychidae. Alexia hirtula is transferred to Delphastus Casey (Coccinellidae: Microweiseinae: Serangiini), establishing the new combination, Delphastus hirtulus (Kirsch), and a lectotype is designated. Of the three paralectotypes, one appears to be conspecific with the lectotype, one is identified as an undescribed species of Microscymnus Champion (Coccinellidae: Cryptognathini), and one, a partial specimen lacking the head, pronotum, and one elytron, is identified as a species of Leiodidae in the tribe Scotocryptini, probably Aglyptinus Cockerell. A diagnosis and redescription of D. hirtulus is provided, and Gordon’s (1994) key to Delphastus is modified to accommodate the newly transferred species. The historical classification of D. hirtulus is discussed along with characters justifying its revised placement.

Cycloneda pretiosa, new species is described from Lago Pirihueico in southern Chile The species is diagnosed and compared with both related and superficially similar species. Habitus and genitalic characters are illustrated. Geographical and temporal data are given. The new species is assigned to the Cycloneda germainii species complex and integrated into the key to species of this group (González and Vandenberg, 2006).