Insecta Mundi
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602 [korr. Vers.]
Species of Mortoniella are revised for the northern and Andean part of the South American continent, including the countries of Bolivia, Peru, Ecuador, Colombia, Venezuela, and Guyana. All previously described species from the region are reillustrated and redescribed, except for Mortoniella santiaga Sykora, 1999 and M. quinuas Harper and Turcotte, 1985, whose types could not be located, and M. tranquilla Martynov, 1912, whose type is based on a female specimen and thus is currently unidentifiable. Included in the revision are 35 described species and 59 new species. Mortoniella similis Sykora, 1999 is considered a junior synonym of M. roldani Flint, 1991, and M. macuta (Botosaneanu, 1998) is considered a junior synonym of M. limona (Flint, 1981). A new subgenus, Nanotrichia, is recognized to accommodate species previously referred to as members of the ormina and velasquezi groups. Mexitrichia pacuara Flint, 1974 is designated the type species for the subgenus. Species previously referred to as members of the bilineata and leroda species groups are retained in the nominate subgenus, along with additional taxa not previously placed to species group, and treated within a number of subgroups. Previously described species of M. (Mortoniella) which are redescribed and reillustrated include: M. angulata Flint, 1963; M. apiculata Flint, 1963; M. atenuata (Flint, 1963); M. bifurcata Sykora, 1999; M. bilineata Ulmer, 1906; M. bolivica (Schmid, 1958); M. chicana Sykora, 1999; M. denticulata Sykora, 1999; M. elongata (Flint, 1963); M. enchrysa Flint, 1991; M. flinti Sykora, 1999; M. foersteri (Schmid, 1964); M. hodgesi Flint, 1963; M. iridescens Flint, 1991; M. leei (Flint, 1974); M. limona (Flint, 1981); M. marini (Rueda Martín and Gibon, 2008); M. paralineata Sykora, 1999; M. paraenchrysa Sykora, 1999; M. pocita (Flint, 1983); M. punensis (Flint, 1983); M. roldani Flint, 1991; M. simla (Flint, 1974); M. spinulata (Flint, 1991); M. squamata Sykora, 1999; M. unilineata Sykora, 1999; and M. wygodzinskii (Schmid, 1958). New species described in M. (Mortoniella), followed by their respective areas of distribution, include: M. acutiterga (Ecuador); M. adamsae (Peru); M. akrogeneios (Ecuador); M. applanata (Peru); M. auricularis (Colombia); M. barinasi (Venezuela); M. biramosa (Venezuela); M. bothrops (Peru); M. brevis (Ecuador, Venezuela); M. bulbosa (Peru); M. catherinae (Peru); M. chalalan (Peru); M. cornuta (Peru); M. cressae (Venezuela); M. croca (Peru); M. curtispina (Venezuela); M. curvistylus (Ecuador); M. dentiterga (Ecuador); M. dinotes (Peru); M. draconis (Ecuador); M. emarginata (Ecuador, Colombia); M. esrossi (Colombia); M. flexuosa (Colombia); M. furcula (Ecuador); M. gilli (Ecuador); M. gracilis (Venezuela); M. grandiloba (Venezuela); M. guyanensis (Guyana); M. hamata (Colombia); M. langleyae (Ecuador); M. longiterga (Ecuador); M. membranacea (Bolivia); M. monopodis (Colombia, Ecuador); M. parameralda (Ecuador); M. pica (Ecuador); M. proakantha (Ecuador); M. prolata (Peru); M. quadrispina (Ecuador); M. rectiflexa (Ecuador); M. ruedae (Bolivia); M. schlingeri (Colombia); M. silacea (Colombia, Ecuador); M. sinuosa (Bolivia, Peru); M. spatulata (Venezuela); M. tanyrhabdos (Venezuela); M. tridens (Peru); M. triramosa (Bolivia); M. tusci (Venezuela); and M. variabilis (Venezuela, Colombia). Species assigned to the subgenus M. (Nanotrichia) which are redescribed and reillustrated include: M. aequalis (Flint, 1963); M. aries (Flint, 1963); M. collegarum (Rueda Martín and Gibon, 2008); M. eduardoi (Rueda Martín and Gibon, 2008); M. macarenica (Flint, 1974); M. pacuara (Flint, 1974); M. usseglioi (Rueda Martín and Gibon, 2008); and M. velasquezi (Flint, 1991). Previously described species of Mortoniella, outside the area of coverage, that are transferred to the subgenus M. (Nanotrichia) include: M. alicula Blahnik and Holzenthal, 2011; M. bocaina Blahnik and Holzenthal, 2011; M. catarinensis (Flint, 1974); M. froehlichi Blahnik and Holzenthal, 2011; M. ormina (Mosely, 1939); M. rodmani Blahnik and Holzenthal, 2008; and M. tripuiensis Blahnik and Holzenthal, 2011. New species in the subgenus M. (Nanotrichia), followed by their respective areas of distribution, include: Mortoniella cognata (Ecuador, Venezuela); M. coheni (Ecuador); M. licina (Ecuador); M. paucispina (Peru); M. quadridactyla (Venezuela); M. simplicis (Venezuela); M. spangleri (Ecuador); M. triangularis (Ecuador); M. venezuelensis (Venezuela); and M. zamora (Ecuador). A key to the males of species from the region is also provided, as well as a key to females for the major subgroups and a species key to females of the velasquezi group. Finally, a partially resolved phylogeny of the species is presented, along with a discussion of evolutionary trends within the genus.
602 [1. Version]
Species of Mortoniella are revised for the northern and Andean part of the South American continent, including the countries of Bolivia, Peru, Ecuador, Colombia, Venezuela, and Guyana. All previously described species from the region are reillustrated and redescribed, except for Mortoniella santiaga Sykora, 1999 and M. quinuas Harper and Turcotte, 1985, whose types could not be located, and M. tranquilla Martynov, 1912, whose type is based on a female specimen and thus is currently unidentifiable. Included in the revision are 35 described species and 59 new species. Mortoniella similis Sykora, 1999 is considered a junior synonym of M. roldani Flint, 1991, and M. macuta (Botosaneanu, 1998) is considered a junior synonym of M. limona (Flint, 1981). A new subgenus, Nanotrichia, is recognized to accommodate species previously referred to as members of the ormina and velasquezi groups. Mexitrichia pacuara Flint, 1974 is designated the type species for the subgenus. Species previously referred to as members of the bilineata and leroda species groups are retained in the nominate subgenus, along with additional taxa not previously placed to species group, and treated within a number of subgroups. Previously described species of M. (Mortoniella) which are redescribed and reillustrated include: M. angulata Flint, 1963; M. apiculata Flint, 1963; M. atenuata (Flint, 1963); M. bifurcata Sykora, 1999; M. bilineata Ulmer, 1906; M. bolivica (Schmid, 1958); M. chicana Sykora, 1999; M. denticulata Sykora, 1999; M. elongata (Flint, 1963); M. enchrysa Flint, 1991; M. flinti Sykora, 1999; M. foersteri (Schmid, 1964); M. hodgesi Flint, 1963; M. iridescens Flint, 1991; M. leei (Flint, 1974); M. limona (Flint, 1981); M. marini (Rueda Martín and Gibon, 2008); M. paralineata Sykora, 1999; M. paraenchrysa Sykora, 1999; M. pocita (Flint, 1983); M. punensis (Flint, 1983); M. roldani Flint, 1991; M. simla (Flint, 1974); M. spinulata (Flint, 1991); M. squamata Sykora, 1999; M. unilineata Sykora, 1999; and M. wygodzinskii (Schmid, 1958). New species described in M. (Mortoniella), followed by their respective areas of distribution, include: M. acutiterga (Ecuador); M. adamsae (Peru); M. akrogeneios (Ecuador); M. applanata (Peru); M. auricularis (Colombia); M. barinasi (Venezuela); M. biramosa (Venezuela); M. bothrops (Peru); M. brevis (Ecuador, Venezuela); M. bulbosa (Peru); M. catherinae (Peru); M. chalalan (Peru); M. cornuta (Peru); M. cressae (Venezuela); M. croca (Peru); M. curtispina (Venezuela); M. curvistylus (Ecuador); M. dentiterga (Ecuador); M. dinotes (Peru); M. draconis (Ecuador); M. emarginata (Ecuador, Colombia); M. esrossi (Colombia); M. flexuosa (Colombia); M. furcula (Ecuador); M. gilli (Ecuador); M. gracilis (Venezuela); M. grandiloba (Venezuela); M. guyanensis (Guyana); M. hamata (Colombia); M. langleyae (Ecuador); M. longiterga (Ecuador); M. membranacea (Bolivia); M. monopodis (Colombia, Ecuador); M. parameralda (Ecuador); M. pica (Ecuador); M. proakantha (Ecuador); M. prolata (Peru); M. quadrispina (Ecuador); M. rectiflexa (Ecuador); M. ruedae (Bolivia); M. schlingeri (Colombia); M. silacea (Colombia, Ecuador); M. sinuosa (Bolivia, Peru); M. spatulata (Venezuela); M. tanyrhabdos (Venezuela); M. tridens (Peru); M. triramosa (Bolivia); M. tusci (Venezuela); and M. variabilis (Venezuela, Colombia). Species assigned to the subgenus M. (Nanotrichia) which are redescribed and reillustrated include: M. aequalis (Flint, 1963); M. aries (Flint, 1963); M. collegarum (Rueda Martín and Gibon, 2008); M. eduardoi (Rueda Martín and Gibon, 2008); M. macarenica (Flint, 1974); M. pacuara (Flint, 1974); M. usseglioi (Rueda Martín and Gibon, 2008); and M. velasquezi (Flint, 1991). Previously described species of Mortoniella, outside the area of coverage, that are transferred to the subgenus M. (Nanotrichia) include: M. alicula Blahnik and Holzenthal, 2011; M. bocaina Blahnik and Holzenthal, 2011; M. catarinensis (Flint, 1974); M. froehlichi Blahnik and Holzenthal, 2011; M. ormina (Mosely, 1939); M. rodmani Blahnik and Holzenthal, 2008; and M. tripuiensis Blahnik and Holzenthal, 2011. New species in the subgenus M. (Nanotrichia), followed by their respective areas of distribution, include: Mortoniella cognata (Ecuador, Venezuela); M. coheni (Ecuador); M. licina (Ecuador); M. paucispina (Peru); M. quadridactyla (Venezuela); M. simplicis (Venezuela); M. spangleri (Ecuador); M. triangularis (Ecuador); M. venezuelensis (Venezuela); and M. zamora (Ecuador). A key to the males of species from the region is also provided, as well as a key to females for the major subgroups and a species key to females of the velasquezi group. Finally, a partially resolved phylogeny of the species is presented, along with a discussion of evolutionary trends within the genus.
601
Genera of Cephaloscymnini (Coleoptera: Coccinellidae: Coccidulinae) are discussed and a key to all recognized genera and species is provided. Succinctonotum, new genus, is proposed. Scymnus laboulbenii Mulsant and Prodilis maculata Weise are transferred to Neaporia as new combinations. Prodiloides bipunctata Weise, and Neaporia compta are transferred to Prodilis as new combinations. Neaporia cuprea Gorham is considered a junior synonym of Neaporia viridiscens Gorham and Cephaloscymnus bruchi Weise a junior synonym of Prodilis volgus Mulsant. New species described in Cephaloscymnus are C. beulah, C. candice, C. juanita. New species described in Neaporia are N. becky, N. bobbie, N. brandy, N. carole, N. cassandra, N. christy, N. daisy, N. deanna, N. dianne, N. felicia, N. gwendolyn, N. hilda, N. irma, N. jennie, N. jenny, N. kay, N. kayla, N. kristine, N. leah, N. lena, N. leona, N. longifrons, N. mabel, N. mae, N. margie, N. marsha, N. miriam, N. misty, N. myrtle, N. naomi, N. nina, N. nora, N. olga, N. opal, N. patsy, N. penny, N. priscilla, N. shelley, N. sonia, N. tracey, and N. violet. New species described in Succinctonotum is S. frosti. New species described in Prodilis are P. ada, P. alberta, P. alison, P. amelia, P. angie, P. araguaensis, P. bartletti, P. belinda, P. blanche, P. brandi, P. cecilia, P. claire, P. cora, P. dubitalis, P. erika, P. eunice, P. fannie, P. faye, P. flora, P. geneva, P. guadalupe, P. harriet, P. hattie, P. inez, P. iris, P. isabel, P. jan, P. janie, P. joanna, P. jodi, P. katrina, P. kristi, P. kristy, P. lindsey, P. lola, P. lula, P. lynda, P. madeline, P. maggie, P. mamie, P. margarita, P. maryann, P. melody, P. molly, P. monique, P. natasha, P. olivia, P. pecki, P. ramona, P. rosie, P. sabrina, P. sandy, P. shelley, P. sherri, P. sheryl, P. sonya, P. susie, and P. yvette. Ponaria, new genus of Coccidulini is proposed. Neaporia caerulea Gorham is transferred to Ponaria as a new combination. New species described in Ponaria are P. daviesi, P. hurtadoi, and P. paprzyckii. Lectotypes are here designated for Neaporia arrowi, N. coelestis, N. metallica, N. guatemalana, N. indagator, and Prodilis pallidifrons.
600
A new cryptic species, Aleochara (Xenochara) castaneimarmotae Klimaszewski, Webster, and Brunke, new species (Coleoptera: Staphylinidae: Aleocharinae), associated with Marmota monax (Linnaeus) burrows and caves in eastern North America, is described and illustrated. A key to Canadian species of subgenus Xenochara Mulsant and Rey and revised distributions of the taxonomically difficult fumata species group are provided. Aleochara quadrata Sharp is recorded from Washington and Oregon for the first time.
596
595
Many nomenclatural changes are implemented in the beetle families Georissidae, Histeridae, Hydraenidae, Hydrochidae, Hydrophilidae, Ptiliidae, Leiodidae and especially Staphylinidae, of the beetle series Staphyliniformia (Coleoptera), in preparation for making a world catalog of this group available online. Limited taxonomic changes are also made in the staphylinid subfamilies Osoriinae and Staphylininae.
At the level of family-group taxa, Article 29.4 of the current (1999) Zoological Code is reviewed and the original spellings of two tribal names, Nymphisterini Tishechkin (Histeridae) and Cryptonotopsisini Pace (Staphylinidae), are resurrected. The tribal name Stictocraniini Jakobson (Staphylinidae) is also resurrected as the valid name for its new synonym Fenderiini Scheerpeltz.
Changes at the genus-group level in Histeridae include placing Contipus Marseul as a new synonym of Hister Linnaeus due to the current placement of its validly designated type species C. subquadratus Marseul; proposal of Contipides Newton gen. nov. (type species Contipus digitatus Marseul) for the 10 species that had remained in Contipus of authors; and new designation of Idolia laevigata Lewis as type species of Idolia Lewis. In Ptiliidae, Rodwayia ovata Lea is newly designated as type species of Rodwayia Lea, and Throscidium germainii Matthews is newly designated as type species of Throscidium Matthews. In Staphylinidae, Paramichrotus Naomi is resurrected as a valid subgenus of Hesperosoma Scheerpeltz with Hemihesperosoma Hayashi placed as a new synonym of it; Sonoma corticina Casey is reaffi rmed as the type species of Sonoma Casey in place of Faronus tolulae LeConte; Stanosthetus Dejean is recognized as an available name and junior synonym of Euplectus Kirby; Taplandria Pace (type species T. guyanensis Pace) is recognized as a junior homonym and new synonym of Taplandria Pace (type species T. fl ava Pace); and Termitobiella Wasmann is resurrected as the valid name for the genus Felda Blackwelder. Replacement names for preoccupied generic or subgeneric names include in Histeridae Bellatricides Newton nom. nov. for Pachylister (Bellatrix) Mazur, junior homonym of Bellatrix Boie; and in Staphylinidae Foxiides Newton nom. nov. for Foxia Pace, junior homonym of Foxia Ashmead, and Xenasterides Newton nom. nov. for Xenaster Bierig, junior homonym of Xenaster Simonwitsch. Taxonomic changes at the generic level in Staphylinidae include proposal of Prolibia Newton gen. nov. (type species Lispinus californicus LeConte) for four Nearctic species recently placed in Clavilispinus Bernhauer; placement of Heterotrochinus Coiffait and its synonym Heterotrochus Coiffait as new synonyms of Eulibia Cameron; placement of the generic or subgeneric names Chapmaniella Bernhauer, Glenothorax Bierig, Euryolinus Bernhauer and Plesiolinus Bernhauer as new synonyms of Platydracus Thomson; and transfer of the subgenus Poikilodracus Scheerpeltz from Staphylinus Linnaeus to Platydracus. First reviser actions are used to select Georissites Ponomarenko (Georissidae) as the correct original spelling over the alternate original spelling Georyssites, and Kyrtusa Pace (Staphylinidae) as correct original spelling over Kirtusa.
Several hundred nomenclatural and taxonomic changes at the species group level are briefl y summarized here but are too numerous to list completely. Replacement names for preoccupied species or subspecies names in current use are proposed in Histeridae (3), Hydrochidae (1), Hydrophilidae (1), Leiodidae (2), Ptiliidae (3) and Staphylinidae (180); an additional staphylinid replacement name, Phloeopora nilgiriensis, is newly proposed by G. Paśnik. New or resurrected combinations are proposed for either nomenclatural or taxonomic reasons in the following genera (with indication of how many names in each genus): in Histeridae, Contipides Newton (10); in Staphylinidae, Abemus Mulsant and Rey (4), Allotrochus Fagel (6), Atheta Thomson (1), Cheilocolpus Solier (4), Eulibia Cameron (4), Foxiides Newton (1), Lispinus Erichson (3), Loncovilius Germain (2), Nacaeus Blackwelder (119), Naddia Fauvel (1), Neohypnus Coiffait and Sáiz (8), Neolosus Blackwelder (1), Ocypus Leach (2), Ontholestes Ganglbauer (1), Platydracus Thomson (59), Prolibia Newton (4) Termitobiella Wasmann (10), Thyreocephalus Guérin-Méneville (4), Xenasterides Newton (1), and Zeoleusis Steel (3). First reviser actions are used to resolve the correct original spellings (of two or more original spellings) of two species of Hydraena Kugelann (Hydraenidae) and 21 species of Staphylinidae. Changes in priority or availability of names are cited to establish the following names as valid over one or more new synonyms each: Acrotrichis rotundata (Haldeman) and Acrotrichis glabricollides Newton sp. nov. in Ptiliidae, Nemadiopsis franki Perreau in Leiodidae, and Gyrophaena nigra Kraatz, Heterothops fumigatus LeConte, Loncovilius germaini (Scheerpeltz), Philonthus upotovus Newton, sp. nov., Stenus fulviventris Rougemont, and nine species of Homalota Mannerheim in Staphylinidae. Finally, the species Eleusis lata Coiffait and Eleusis microlestiformis Coiffait are noted as not belonging to the genus Eleusis Laporte de Castelnau or to Staphylinidae, and are transferred without generic assignment to the subfamily Inopeplinae of the family Salpingidae.
594
The tribe Zolini (Carabidae: Trechinae) is revised for New Zealand. Two subtribes, fi ve genera, and fortyeight species are recognized. The presence of the subtribe Merizodina is confi rmed for New Zealand; it includes three genera (Maungazolus n. gen., Pseudoopterus Csiki, 1928, and Synteratus Broun, 1909). The subtribe Zolina contains two genera (Oopterus Guérin-Méneville, 1841 and Zolus Sharp, 1886).
One genus and twenty-fi ve species are described as new: Maungazolus n. gen.; Maungazolus acutus n. sp., Maungazolus priestleyensisn. sp., Maungazolus ranatungaen. sp., Maungazolus septempunctatus n. sp., Maungazolus tararuaensisn. sp., Oopterus anglemensisn. sp., Oopterus arthurensisn. sp., Oopterus corvinki n. sp., Oopterus discoideus n. sp., Oopterus garnerae n. sp., Oopterus marrisin. sp., Oopterus mohiensis n. sp., Oopterus monticola n. sp., Oopterus nanus n. sp., Oopterus nunni n. sp., Oopterus palmai n. sp., Oopterus punctatus n. sp., Oopterus quadripunctatus n. sp., Oopterus taieriensis n. sp., Oopterus trechoides n. sp., Pseudoopterus fi ordlandensis n. sp., Pseudoopterus otiraensis n. sp., Zolus kauriensis n. sp., Zolus unisetosus n. sp., Zolus wongi n. sp.
Lectotypes are designated for Zolus helmsi Sharp, 1886 and Oopterus parvulus Broun, 1903.
Seven taxa are reinstated as valid genera and previous combinations: Zolus Sharp, 1886 and Pseudoopterus Csiki, 1928 (previously synonymized with Oopterus Guérin-Méneville,1841); Oopterus carinatus Broun, 1882 = Zolus carinatus (Broun, 1882); Oopterus helmsi (Sharp, 1886) = Zolus helmsi Sharp, 1886; Oopterus latipennis Broun, 1903 = Pseudoopterus latipennis (Broun, 1903); Oopterus plicaticollis Blanchard, 1843 = Pseudoopterus plicaticollis (Blanchard, 1843); Oopterus subopacus (Broun, 1915) = Zolus subopacus Broun, 1915.
Three new combinations are established: Oopterus pallidipes Broun, 1893 = Maungazolus pallidipes (Broun, 1893); Oopterus parvulus Broun, 1903 = Maungazolus parvulus (Broun, 1903); Oopterus pygmeatus Broun, 1907 = Maungazolus pygmeatus (Broun, 1907).
Eleven new synonymies are established: Oopterus basalis Broun, 1915 and Oopterus minor Broun, 1917 = Maungazolus pygmeatus (Broun, 1907); Oopterus nigritulus Broun, 1908 and Oopterus laevigatus Broun, 1912 = Oopterus sobrinus Broun, 1886; Oopterus sculpturatus ovinotatus Broun, 1908 = Oopterus sculpturatus Broun, 1908; Oopterus probus Broun, 1903 and Oopterus latifossus Broun, 1917 = Pseudoopterus latipennis (Broun, 1903); Oopterus puncticeps Broun, 1893 = Oopterus laevicollis Bates, 1871; Zolus atratus Broun, 1893 and Zolus femoralis Broun, 1894 = Zolus carinatus (Broun, 1882); Zolus labralis Broun, 1921 = Zolus helmsi Sharp, 1886.
A revision of all taxa is provided. Descriptions, identifi cation keys, illustrations of male genitalia, habitus photos, distributional data and maps are given. Extensive information on ecology, biology, dispersal power, and collecting techniques is included for each species.
593
Species of Hydrosmecta C. G. Thomson, occurring in eastern Canada, are studied and reviewed. Six species are recognized, and two of them are described as new to science. The new species are: Hydrosmecta canadensis Webster and Klimaszewski, new species, and Hydrosmecta minutissimoides Webster and Klimaszewski, new species. Two species described in the USA are recorded from Canada and New Brunswick for the first time: Hydrosmecta caduca Casey and Hydrosmecta dulcis Casey. New distribution and collection data, diagnoses, a key for species identification, and images of habitus and genital structures are provided. Hydrosmecta newfoundlandica Klimaszewski and Langor, 2011, is transferred to the genus Atheta Thomson based on morphology of genital structures.
592
This paper revises the genus Ganelius Benesh, which is endemic to Madagascar, in the stag beetle tribe Figulini Burmeister (Coleoptera: Lucanidae: Lucaninae). Species in the genus are striking in their highly sexually dimorphic mandibles, a character that rarely occurs in the tribe. The genus was previously comprised of four species, but three of these names were problematic or questionably applied. A lectotype is designated for G. madagascariensis Laporte, a neotype is designated for Ganelius oberndorferi Nonfried, and Nigidius passaliformis Benesh is newly synonymized with G. oberndorferi. Two new species, G. gnamptus Paulsen and G. zombi Paulsen, are described from western Madagascar. The identity of the overlooked Ganelius nageli (Kriesche) is fixed through a neotype designation, and the species is moved to the new genus Agnelius, which is distinguished from Ganelius by a lack of sexual dimorphism, serrate protibiae, and a more flattened body.
591
New genera and species of mimetic Cleridae from Mexico and Central America (Coleoptera: Cleroidea)
(2017)
New genera and species of putatively mimetic Cleridae (Coleoptera: Cleroidea) are described from Mexico and Central America: Namba testacea, new genus and species from Nuevo León, Mexico; Aulicoides leavengoodi, new genus and species from Jalisco, Mexico; and Quadrophenia, new genus, which includes Q. townshendi, new species, from Mexico, Nicaragua, Costa Rica and Panama, Q. mooni, new species, from Costa Rica, Q. entwistlei, new species, from Guatemala and Honduras, Q. daltreyi, new species, from Costa Rica, and Q. sumidero new species, from Chiapas, Mexico. A key to the species of Quadrophenia is provided. All of the newly described species are hypothesized to be Batesian mimics of aposematically colored coleopteran models, some of which are known to be chemically protected.