VOL. 104 JANUARY 2002 NO. 1 (ISSN 0013-8797) PROCEEDINGS

of the ENTOMOLOGICAL SOCIETY

of WASHINGTON PUBLISHED QUARTERLY

ADAMSKI, DAVID—A new of Glyphidocera Walsingham (Lepidoptera: Gelechioidea:

Glyphidoceridae ) from Costa Rica 119

BRAILOVSKY, HARRY—A new species of Maevius Stal from Australia and some notes on the family Hyocephalidae (Hemiptera: Heteroptera) 41

BRAILOVSKY, HARRY—A new genus and a new species of Daladerini (Hemiptera: Heteroptera: Coreidae) from Madagascar Ill

BURKS, ROGER A. and JOHN D. PINTO—Reproductive and electrophoretic comparisons of

Trichogramma californicum Nagaraja and Nagarkatti with the T. minutum complex (Hyme- noptera: Trichogrammatidae) 33

CARROLL, J. F. —Notes on the responses of host-seeking nymphs and adults of the ticks Ixodes scapularis and Amblyomma americaniim (Acari: Ixodidae) to canine, avian, and deer-produced substances 73

DELLAPE, P M., M. del C. COSCARON, and B. F. AMARAL FILHO—Immature stages of Montina confusa (Stal) (Heteroptera: Reduviidae: Harpactorinae) 168

DeWALT, R. EDWARD, DONALD W. WEBB, and AMY M. SOLI—The Neoperia clymene (Newman) complex (Plecoptera: Perlidae) in Illinois, new state records, distributions, and an identification key 126

FERREIRA, PAULO SERGIO FIUZA and THOMAS J. HENRY—Descriptions of two new species of Fitlvius Stal (Heteroptera: Miridae: Cylapinae) from Brazil, with biological and biogeographic notes on the genus 56

GOEDEN, RICHARD D.—Descriptions of Tephritis footei and T. headricki, new species (Dip- tera: Tephritidae), with notes on their life histories in southern California 142

HALL, JASON P. W—A review of the new riodinid butterfly genus Panaropsis (Lepidoptera: Riodinidae: Symmachiini) 63

HARRIS, STEVEN C. and OLIVER S. FLINT, JR.—New Alisotrichia (Trichoptera: Hydroptili- dae) from Central and South America and the Greater Antilles 195

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PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 221-236 THE NEW WORLD GENUS CROTCH (COLEOPTERA: : COCCINELLINI): HISTORICAL REVIEW, NEW DIAGNOSIS, NEW GENERIC AND SPECIFIC SYNONYMS, AND AN IMPROVED KEY TO NORTH AMERICAN SPECIES

Natalia J. Vandenberg

Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560-0168, U.S.A. (e-mail: [email protected])

Abstract. —Taxonomic, nomenclatoral, and comparative moiphological studies of Cv- cloneda Crotch are reviewed. The genus is diagnosed and compared to both related and superficially similar taxa. Pseiidadonia Timberlake is recognized as a junior synonym (n. syn.). The species composition of the fauna is delineated, but the need to re-examine species limits and nomenclatoral priorities is stressed. New synonyms are proposed for two species of Cycloneda occurring in North America: Coccinella krikkeni lablokoff- Khnzorian, reported from India, and Cycloneda atra Casey (of unknown origin) are new junior synonyms of Cycloneda munda (Say); Cycloneda hondurasica Casey (Honduras) and Coccinella reflexa Germain (Chile) are new junior synonyms of Cycloneda sangui- nea (Linnaeus). Adalia galapagoensis Van Dyke is transferred to the genus Cycloneda (Cycloneda galapagoensis, n. comb.) and identified as a close relative of Cycloneda sanguinea. Cycloneda sanguinea cayniana Chapin is elevated to full species status {Cy- cloneda caymana, n. status). A new key to species of North American Cycloneda is provided.

Key Words: Cycloneda, Coccinellina, Pseudadonia, Synonychini, Coccinellini, lady , systematics, synonymy, Nearctic, Neotropical

Cycloneda Crotch is a New World genus without black spots, and a black pronotum of lady in the tribe Coccinellini. In with a white border design and pair of dis-

Central and South America (Blackwelder cal spots (Fig. 9). The basic components of

1945), the genus is a composite one, com- the white design are often coalesced to form prising an assemblage of two dozen or more a complete or broken ring in each lateral species which generally share a rounded third (Figs. 2, 5), and a median spur shaped convex body form, highly polished cuticle, mark is often present at anterior border and little else. In contrast, the three North (Figs. 2, 5, 8). America indigenes Cycloneda sanguinea The North American Cycloneda were (L.) (type species), C munda (Say), and C keyed and diagnosed most recently by Gor- polita Casey—appear to form a close-knit, don (1985), who followed Leng (1903) in monophyletic group, whose members often the use of pronotal color patterns and elytral have been confused in the literature (Gor- ground color in the separation of species don and Vandenberg 1993). These species (treated as varieties by Leng 1903). Unfor- are characterized by red to yellowish elytra tunately, these selected attributes are less —

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON than completely reliable, particularly in the elongated forms. Subsequent authors pro- identification of specimens from the west- vided keys and diagnoses for Cycloneda ern United States. In addition, both of the (Casey 1899, Leng 1920), but clearly con- above authors excluded the species Cyclo- cerned themselves only with an easily de- neda atra Casey ( 1 899) from their revisions finable subgroup consisting of the three on the assumption that the unique all-black North American species (C. sanguinea, C. type specimen from an unrecorded locality inunda, C polita) and synonyms thereof. was unlikely to be a member of the North Casey split off two new genera Olla American fauna. (1899) and Spiloneda (1908)—but this did

Here, I resolve the identity of Cycloneda little to improve the overall classification. atra and that of a second equally mysteri- With the transfer of additional species to ous specimen from Southern India de- Olla (Casey 1908), that genus soon proved scribed by lablokoff-Khnzorian (1982) as as heterogeneous and ill-defined as Cyclo- Coccinella krikkeni lablokoff-Khnzorian. neda (Vandenberg 1992). Spiloneda, on the The identities of Coccinella reflexa Ger- other hand, has remained monotypic be- main ( 1 854) from Chile and Cycloneda cause it was stipulated for "such forms as hondurasica Casey from Central America Gilardini, Muls., from Colombia and Cen- ." also are investigated. As a result, four new tral America . . (Casey 1908) without de- synonyms are proposed, two each for Cy- tailing additional species names or distin- cloneda mimda and Cycloneda scmguinea. guishing traits. The taxonomic history of the genus Cyclo- The examination of internal genital struc- neda is reviewed, the composition of the tures was an important advancement for

Neotropical Cycloneda fauna discussed, lady beetle , although it conferred and a new key to North American Cyclo- no immediate benefit to the classification of neda is proposed to make the identification Cycloneda species. Weise introduced the of species more reliable. use of male genitalia (shape of basal lobe) for distinguishing lady beetles of similar Historical Resume habitus (e.g., Weise 1904b), but did not use

The name Cycloneda was first applied by these structures for defining genera or de- Crotch (1871) as a replacement name for termining broader relationships. He contrib- Daidis Mulsant (1850) (preoccupied by uted a heterogeneous assortment of new Daulis, Erichson 1842). Although Crotch species to Cycloneda between 1898 and had restricted use of the name to the New 1922 (Weise 1898, 1902, 1904a, 1906,

World fauna (about 40 species), he still re- 1922). fered to Cycloneda as "an unsatisfactory Wilson (1926) conducted a morphologi- assemblage of species having but little in cal study on the male and female genitalia common" and lamented the lack of char- of selected representatives of the family acters to separate it into meaningful groups Coccinellidae. He found characteristics (Crotch 1874). The subsequent designation which suggested a close relationship be- of Coccinella sanguinea Linnaeus as the tween Cycloneda and Coccinella, and in- type species of Cycloneda (Crotch 1874) dicated that the male genitalia of Olla were only added to existing taxonomic confu- so extraordinary as to set it well apart from sion. While C. sanguinea clearly exempli- the other genera studied. Wilson's obser- fied the rounded convex body form (Figs. vations appear to have been overlooked by

8, 10) of most of its nominal congeners, it a subsequent generation of lady beetle tax- was not particularly closely related to the onomists who continued to stipulate minor majority of them. Conversely, many of its differences in external morphology, while true affiliates (Figs. 12, 13) were relegated ignoring the abundant generic characters to other genera because of their flatter, more provided by genital structures. —

VOLUME 104. NUMBER 1 223

Chapin (1941) was one of the first spe- inclined and descending externally or very cialists to question the value of external broad." characteristics used to separate Cyckmeda Despite an often noted resemblance be- from the Neotropical genera Neda Mulsant, tween Cycloneda species (sensu Timberlake Prociila Mulsant, and Olla, all of which had 1943) and C. emarginata (Mulsant 1850, been grouped together in the tribe Synon- Crotch 1874, Koebele in Timberlake 1943), ychini. He found that ranking a random se- Timberlake did not make a rigorous com- lection of 17 species by epipleural width or parison between Cycloneda and his new ge- depth of mesosternal emargination (at the nus Coccinellina, undoubtedly influenced time considered key differentiating charac- by the fact that members of these genera teristics), failed to segregate species accord- conventionally were placed in two distinct ing to their presumed generic placement or tribes (Synonychini and Coccinellini, re- to produce significant breaks between spectively) (Korchefsky 1932). Actually, groups of species in the series. In contrast, both nominal genera exhibit a wide range architecture he observed that three or more definite of epipleural with considerable overlap. types of genitalia could be found within the cross-generic Although Cycloneda sanguinea (type) may be said to have broad series, yet he made no attempt to revise ge- epipleura which are steeply inclined exter- neric assignments based on that observa- nally (Fig. 7), its congeners, polita (Fig. tion. C 4) and C. munda, (Fig. 1) have narrower Timberlake (1943) refined the definition epipleura which are nearly horizontal, as in of Cycloneda by restricting the name to Coccinellina emarginata. The epipleura of ""scniguinea and allies with immaculate el- C pulchella (placed in Coccinellina by ytra." He described three new genera for Timberlake) (Fig. 14) are broader than in some of the former Cycloneda species either Cycloneda polita or C munda. There based on material in the famous Koebele is also significant sexual dimorphism: in all Collection Panineda, Erythroneda, and of the aforementioned species, the male epi- Chloroneda. Unfortunately, he did not pleura tend to be broader and more steeply study types or material from South America inclined than the female epipleura. which might have allowed him to resolve Mader (1958) provided a key to Cyclo- some of the erroneous species synonymies neda species and former Cycloneda species proposed by Crotch (1874) and perpetuated useful for identification purposes, but based by other authors. He was uncertain, there- almost entirely on color patterns. He made fore, of the number of valid species which no attempt to validate or refute the genera in each of these should have been included proposed by Casey and Timberlake, and did new genera. not discuss male genital characters. Timberlake (1943) also described the ge- Chapin (1969) synonymized Pseudadon- nus Coccinellina (type species Coccinella ia cliiliana with Coccinella fulvipennis emarginata Mulsant) (Fig. 13) for Neotrop- (placed in Coccinellina by Timberlake). ical species formerly classified in Coccinel- Chapin continued to recognize Pseudadon- la, and the genus Pseudadonia (type species ia as a valid genus even though the distinc- Pseudadonia chiliana Timberlake) (Fig. 12) tive characteristic of the inflated basitarsi of based on a single male specimen from Chile the type specimen was confirmed as gender with dilated front and middle basitarsi specific, and no derived characteristic was (compare Figs. 15 and 16). Timberlake's identified for Coccinellina which would ex- generic key characterized the epipleura of clude C fulvipennis from membership. Coccinellina and Pseudadonia species as Several important regional works cover- "horizontal and never much expanded" and ing the genus Cycloneda were published in those of Cycloneda species as "more or less the next two decades. Gordon (1985) re- —

224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON vised the North American Cycloneda spe- neda and three other genera Coccinella, cies, but did not attempt to review non- Eiythroneda, and Neocalvia—was briefly North American species names and types discussed in a revision of the genus Ery- for possible synonymies. Gordon compared throneda (Vandenberg and Gordon 1988). the external morphologies of Cycloneda Vandenberg (1992) revised the genus Olla, and Olki and contrasted the male and fe- provided a key to major genitalic arche- male genitalia, but recognized the need to types occurring in the former Synonychini study the Neotropical species in order to as- and indicated the proper genus group affil- sess the significance of observed differenc- iation for each genitalic configuration. es and determine generic boundaries, lablo- These different archetypes undoubtedly koff-Khnzorian (1982) included some re- correspond to the several categories re- marks on New World Coccinellidae in his feiTed to by Chapin (1941 ) in his studies of revision of the Old World species. He con- the male genitalia of Cycloneda sensu lato. cluded that Cycloneda is most closely re- Although Chapin's work did not list the lated to Hannonia and Xanfhadalla, but he species examined, his handwritten records classified a newly discovered species (C and slide collection at the National Museum krikkeni lablokoff-Khnzorian ) with a re- of Natural History, Smithsonian Institution, markable resemblance to Cycloneda niunda Washington, DC (USNM), show that he in the genus Coccinella. dissected examples of each of the disparate Arioli (1985) contributed an analysis lineages classified in Cycloneda at that (similarity phenogram) of the Coccinellini time. Vandenberg's treatment of the new of Rio Grande do Sul, Brasil. She utilized genus Spilindolla (Vandenberg and Gordon some characters with good potential as in- 1996) provided additional information and dicators of phylogenetic relationships (e.g., genitalic illustrations for separating Neo- male genitalia, postmetacoxal line, pronotal tropical lady beetle genera and generic maculation), but the addition of too many groups. Using these improved criteria to de- trivial (labile) characters (e.g., coloration of termine relationships, the male and female certain structures) had the effect of obscur- genitalia of most '"Cycloneda^' species seg- ing some of the relationships otherwise regate with Neda, Olla, Spiloneda, Para- nicely revealed in her analysis, and in one neda, and allied genera, or with Egleis Mul- case yielded the lowest similarity coeffi- sant and allies, while those of the type spe- cient for a male and female of the same cies and a minority of others reveal a much species. closer relationship to Coccinella. Vanden- The idea of synonymizing Coccinellina berg (1992) also indicated differences be- with Cycloneda was first suggested publicly tween the larval dorsal armature of Cyclo- by lablokoff-Khnzorian in an informal cor- neda and superficially similar species allied respondence to Coccinella newsletter to Olla, and, in a later work (Gordon and

(1990), although it was proposed earlier in Vandenberg 1993), identified distinctive a format not intended as a permanent sci- features of the larval head capsule and mi- entific record (Vandenberg 1987). Vanden- crosculpturing for separating species of berg published a series of papers between Olla, Cycloneda, and Coccinella. 1988 and 1996 which refined and clarified Thus, Cycloneda has been a composite the relationships between Cycloneda and genus since its conception in 1871. A few other taxa, but which focused primarily on species have been removed through the cre- the revision (Erythroneda, Olla) or new de- ation of new genera, but taxonomic pro-

scription (Cirocolla Vandenberg, Spilindol- gress has been hindered severely by 1 ) the la Vandenberg) of genera whose species nearly exclusive use of superficial and had been improperly classified in Cyclone- poorly defined external characters to deter- da. The close relationship between Cyclo- mine generic assignments, 2) failure either VOLUME 104. NUMBER 1 225 to examine the male and female genitalia or Gordon and Vandenberg 1993: 302 (lar- to attach proper evolutionary significance to val descriptions, N. Amer. larval key, lar- their observed patterns of similarity, 3) a val generic comparisons); Vandenberg tendency for researchers to focus on taxa and Gordon 1996: 547 (generic compar- from only one region, one tribe or a single isons); Vandenberg 1997 (example of collection, and 4) the description of new clinal variation in C ancoralis). species based on unknown {Cycloneda Neda (Cycloneda): Chapuis 1876: 201. atra) or dubious (Coccinella krikkeni) type Daulis Mulsant 1850: 296 (Type species: localities, which were then omitted from not designated) (not Daulis Erichson later regional studies. 1842); Crotch 1874: 162 (as synonym of The genus Cycloneda (as recognized Cycloneda) (world revision); Berg 1874: here) is in need of a complete species level 290; Chapuis 1876: 201 (as synonym of revision. Many species exhibit both clinal Neda {Cycloneda) (diagnosis). variation (Vandenberg 1997) and discrete Coccinella {Cycloneda) Leng 1903: 202 (N. polymorphism; others appear poorly differ- Amer. key). entiated from their congeners, making the Coccinellina Timberlake 1943: 15 (Type delimitation of the different species partic- species: Coccinella emarginata Mulsant): ularly challenging. I hope that the present lablokoff-Khnzorian 1990: 59 (as syno- preliminary contribution will be of use to nym of Cycloneda). those who wish to pursue this interesting Pseudadonia Timberlake 1943: 53 (Type area of investigation. species: Pseudadonia chiliana Timber- lake) (preoccupied name, not Pseudadon- Genus Cycloneda Crotch ia Handlirsch 1906); Chapin 1969: 468. Cycloneda Crotch 1871: 6 (list of species) New synonym. (Type species: Coccinella sangidnea L., by subsequent designation of Crotch Diagnosis. —Distinguished from most 1874); 1873a: 371 (U.S. revision); other genera by the characteristic repertoire 1873b: 50 (N. Amer. checkHst) 1874: 162 of pronotal color patterns which vary as (world revision); Gorham 1892: 169 (C. shown (Figs. 2, 5, 8, 9, 10, 11, 12, 13, 14), Amer., distribution); Casey 1899: 84 or derive from Figs. 8 and 9 through loss (U.S., generic key); 1908: 404 (generic of discal spots, or from Figs. 2 and 5 limits); Leng 1920: 216 (N. Amer. cata- through extension of pale areas to form log); Wilson 1926: 63 (genital morphol- complete ring or solid white oval in each ogy, generic comparisons, higher classi- lateral third; head with gender specific color fication); Korschefsky 1932: 282 (world patterns as shown (Figs. 8, 9), ranging to catalog); Chapin 1941: 165 (generic com- all or mostly white in males and all or most- parisons); Timberlake 1943: 23 (generic ly black in females; elytra deep red to fla- limits); Wingo 1952: 24 (Mex., C. and S. vous or rarely ashen, with or without ad- Amer., W. Indies checklist); Mader 1958: ditional black and or white marks; venter 238 (Amer. key to species); Hatch 1961: black to dark brown marked with lighter

181 (N. Amer. generic key); J. Chapin patches. Postmetacoxal line incomplete, 1974: 62 (Louisiana revision); Belicek closely paralleling hind margin of first ab- 1976: 330 (W. Can., Alaskan revision); dominal sternite in outer third; oblique line Gordon 1985: 819 (N. Amer. revision, obsolete or represented by faint to moder- prey, distributions, genitalia); Vanden- ately distinct integumental wrinkle (as op- berg and Gordon 1988: 33 (generic com- posed to a sharply incised line). Middle and parison); lablokoff-Khnzorian 1990: 460 hind tibiae with pair of spurs at apex. In- (diagnosis, generic comparison); Vanden- fundibulum of female genitalia (Fig. 25) berg 1992: 372 (higher classification); well developed, flared at distal end and of- 126 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ten bearing a bulblike enlargement (some- ceptionally long antennae, coarsely facet- times obscure due to transparency) at prox- ted, often closely placed eyes (separated by imal (bursal) end; sipho of male genitalia less than two diameters) and the reddish ar- (Fig. 23) constricted before apex, terminat- eas of the pronotum and elytra entirely ing in a membranous area with imbedded transparent (see Vandenberg and Gordon spicules (reduced in some species); siphon- 1988 for a more detailed comparison). The al capsule well developed, often pigmented, Holarctic genus Coccinella (Figs. 17, 18) with inner arm arcuate to angulate, outer also is allied closely to Cycloneda. Cocci- arm often crested externally; basal lobe nella can be distinguished by the very dif- (Fig. 24) undivided; basal piece never high- ferent repertoire of pronotal color patterns ly elongated, more or less quadrate; para- which feature a large, subtrapezoidal or tri- meres well separated at base. angular white mark in each anterior pron- Remarks. —Lady beetles with similar otal angle, sometimes narrowly joined pronotal color patterns occur occasionally along the anterior border. Coccinella spe- in other coccinelline genera. Species of cies also have a more robust body and ap- Adalia with a very similar appearance can pendages, and the postmetacoxal line with be distinguished by the form of the post- a sharply incised oblique line. While Cyclo- metacoxal line which recurves toward the neda is restricted to the New World and has abdominal base. Olla and allied genera, its highest concentration in the Neotropics, which make up the bulk of the Neotropical Coccinella occurs in both the Eastern and coccinelline fauna {Olla, Cirocolla, Spilin- Western Hemispheres, but does not extend dolla, Spiloneda, Neda, Mononeda Crotch, south of Mexico. Neoharmonia Crotch, Procula, Paraneda, Species composition.—As defined here, Clynis Mulsant, Chloroneda). depart radi- Cycloneda includes species formerly as- cally from Cycloneda in the form of the signed to Coccinellina and Pseudadonia. genitalia of both sexes: female with infun- Timberlake (1943) provisionally transferred dibulum absent or rarely present as a simple the following species names from Cocci- tubular and weakly sclerotized sheath; si- nella when he proposed the genus Cocci- pho of male genitalia not constricted before nellina: C. ancoralis Germar, C areata apex, terminating in a simple rounded Mulsant, C emarginata Mulsant, C eryngii opening, preapical projections or lobes of- Mulsant, C. fulvipennis Mulsant, C. lucasii ten present; basal lobe often divided (Figs. Mulsant, C. petitii Mulsant, C. pulchella 20-22); basal piece generally longer than Klug. He also added two new species: Coc- wide, often highly elongate. Species related cinellina ecuadorica Timberlake and Coc- to Mulsantina Weise and Egleis can be sep- cinellina shannoni Timberlake. Most of the arated by the absence of tibial spurs as well older names have accumulated a number of as by the form of the male (Fig. 19) and synonyms as reported in the most recent female genitalia (see Vandenberg 1992, checklists and catalogs (Korchefsky 1932, Vandenberg and Gordon 1996 for further Blackwelder 1945), but Timberlake did not details). investigate the priority of existing names, Genera related to Cycloneda share the nor examine the type material to determine same major genitalic features outlined in which synonyms are justified. Gordon the diagnosis above. Most of these can be (1987) transferred additional species names separated easily by more obvious external to Coccinellina (C germainii Crotch, C. ar- characters. Cycloneda differs from the re- cula Erichson, C. ocelligera Crotch), but lated genera Erythroneda and Neocalvia by cited Timberlake's work as the source of the the presence of black to very dark brown change. Although Timberlake did not stip- pigmentation on the head, pronotum and ulate these additional names, he had made venter; the latter two genera also have ex- a general comment under his description of VOLUME 104. NUMBER 227

r 3

elytra, frontal Figs. 1-9. 1-3, C\cloneda munda. 1, Median cross-section, left elytron. 2, Head, pronotum and pronotum view, male. 3, Left hind leg. ventral view. 4-6, C. polita. 4, Median cross-section, left elytron. 5, Head, and elytra, frontal view. male. 6, Left hind leg, ventral view. 7-9. C. sangiiinea. 7, Median cross-section, left view, female. elytron. 8. Head, pronotum and elytra, frontal view, male. 9, Head, and pronotum, frontal

Coccinellina that "The neotropical species C. ocelligera does not belong here, but (except C. transversogiittata Fald.) which should be placed in or near Neda. have been referred to Coccinella are rather Pseudadonia Timberlake was always a different from the famihar holarctic species, monotypic genus. Following the synonymy ." Gordon proposed by Chapin (1969) it contributes . . (Timberlake 1943). Therefore, = transferred all the remaining Neotropical only Pseudadonia fidvipennis (Mulsant) ( species represented in the Crotch collection Pseudadonia chiliana Timberlake), a spe- which had not already been removed cies which Timberlake also had placed in through some other more recent revision. Coccinellina. This species fits well within This was appropriate in all but the last case: the generic concept of Cycloneda as diag- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON nosed above, furthermore the name Pseu- stcoxal line is more developed than in most dadonia is preoccupied by Pseudadonia members of Cycloneda. Handlirsch 1906, a fossil mycetophilid. Key to North American Cycloneda The list of species retained from those Species formerly classified in Cycloneda (Black-

1. (Fig. 10); suprahemi- welder 1945) is quite small. The three Form nearly circular spherical (Fig. 8); elytral epipleuron strongly North American species, C. sanguinea, C. concave, steeply descending externally (Fig. 7). C. polita, all belong here. The munda, and Female with white pronotal border strongly nar- more recently described subspecies Cyclo- rowed or interrupted anteromedially (Fig. 9); neda sanguinea caymana Chapin (1957) male with border entire, with median tapered prolongation extending posteriorly (Fig. 8), this also belongs in this genus, but is distinct mark lacking in female. Southern U.S., from from C. sanguinea. Moreover, it overlaps in North Carolina to Florida, west to Southern Cal- distribution with C. sanguinea limbifer ifornia, widespread in the Neotropics 2 (Chapin 1957) without forming intergrades. - Form oval, slightly elongate; subhemispherical Cycloneda caymana (Chapin) (new status) (Figs. 2, 5); elytral epipleuron weakly concave, subhorizontal (Figs. 1, 4). Both genders with resembles our west coast species, Cyclo- white pronotal border continuous across ante- polita (Fig. 2), because of the similar neda rior margin, with median tapered prolongation dorsal color pattern, small size, and oval, extending posteriorly (Figs. 2, 5). Widespread somewhat depressed body form, but a clos- in North America but absent from Florida and lower part of southern most states, extending er relationship to C sanguinea is suggested northward into southern Canada 3 by the shared attributes of an apically 2. Lateral border of elytron clear amber (Fig. 10) knobbed basal lobe (similar to Fig. 24), and sanguinea sanguinea (L.) steeply descending, concave elytral epipleu- - Lateral border of elytron narrowly darkened

(Fig. 1 ). West Indian subspecies, also re- ron (similar to Fig. 7). 1 A corded from southern Florida Adalia galapagoensis Van Dyke (1953) sanguinea limbifer Casey is a species from Charles Island of the Gal- 3. Hind leg with femoral apex, tibia, and tarsus apagoes Archipelago which also belongs to cream colored to reddish brown, rest of femur Cycloneda {Cycloneda galapagoensis Van black (Fig. 3). Elytron pale ; paler area on base of elytron, when evident, begins at scu- Dyke (new combination)). Its original tellum and continues as narrow band around placement in Adalia probably is due to a humeral angle (Fig. 2). Widely distributed in misinterpretation of the configuration of the the U.S. and southern Canada east of the postmetacoxal line, which can be used to Rocky Mountains munda (Say) - distinguish the two genera (see discussion Hind leg with femoral apex, small area at base and apex of tibia, and tarsus cream colored to under the diagnosis for Cycloneda and the reddish brown, rest of femur and tibia black ensuing remarks above). Cycloneda gala- (Fig. 6). Elytron pale orange to dark red; paler pagoensis is a close relative of Cycloneda area on base of elytron, when evident, restrict- sanguinea (widespread on the island chain). ed to semicircular spot adjacent to scutellum, not attaining humeral angle (Fig. 5). Widely The two species share a rounded convex distributed in the western U.S. and southern body form and similar structure to the basal Canada from the Pacific coast through the lobe of the male genitalia. Rocky Mountains polita Casey Cycloneda fryi Crotch and Cycloneda bioculata Korchefsky are two other species Cycloneda sanguinea sanguinea (L.) (Figs. 23-25) which belong in or near Cycloneda. Coc- 10, cinella no. 18 in the Biologia Centrali- Coccinella sanguinea Linnaeus 1763: 10. Americana (Gorham 1892) is an undescri- Daulis sanguinea: Mulsant 1850: 326. bed species which is better placed here than Cycloneda sanguinea: Crotch 1871: 6 elsewhere. The elytra of this species are an Crotch 1873a: 372; Crotch 1874: 164 unusual ashen color, and the oblique po- Blatchley 1910: 515; Korschefsky 1932 VOLUME 104, NUMBER 1 229

Figs. 10-16. 10-14, Habiti of Cycloneda species (male). 10, C. sanguinea sanguinea. 11, C. sanguinea limbifer. 12, C. fulvipennis (= Pseudadonia chiliana, type species of Pseudadonia). 13, C emarginata (type species of Coccinellina). 14, C. pulchella. 15-16, Middle legs of Cycloneda species. 15, Middle leg of C. emarginata (male) showing unmodified basitarsus typical of most Cycloneda species. 16, Middle leg of C. fulvipennis (male) showing swollen basitarsus (arrow). 230 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

286; Timberlake 1943: 23; Wingo 1952: assignment, descriptions of egg, larva, 46; Mader 1958: 241 (in key); J. Chapin pupa, habitats, prey). 1974: 62; Phuoc and Stehr 1974: 58 (pu- Coccinellina reflexa: Chapin 1969: 467 (re- pal description, key); Saini 1983: 392 moved from synonymy, generic reassign- (descriptions of larval color patterns); ment). 1985: 5 (colored photo 4th instar larva). Coccinella {Cycioneda) scuiguinea: Leng Cycioneda sanguinea sanguinea: Gordon 1903: 202. 1985: 820 (in key, diagnosis, distribution, Coccinella (Cycioneda) sanguinea van im- genitalia); Gordon and Vandenberg 1993: maculata: Leng 1903: 203. 307 (description of 4th instar larva, larval Coccinella (Cycioneda) sanguinea van rub- key). ripennis: Leng 1903: 203. Neda sanguinea: Bruch 1915: 388. Coccinella immaculata Fabricius 1792: Remarks.—Gordon (1985) published a bibliography for Cycioneda 267. synonymical foreign Daulis immaculata: Mulsant 1850: 327. scmguinea, but did not investigate material for possible unreported synonyms. Cycioneda immaculata: Casey 1899: 92 (in Examination of the female holotype of Cy- key); Gordon 1985: 820 (as synonym of hondurasica Casey ("Cycioneda sanguinea). cioneda hondurasica: Hond/CASEY bequest 1925/ Cycioneda munda ab. immaculata: Mader 35524 [red label]/Cycloneda 1958: 241. TYPE USNM hondurasica Csy[handwritten]/Casey de- Daulis steini Mulsant 1866: 222; Crotch sanguin-10'') reveals that it also be- 1874: 164 (as synonym of sanguinea) term sanguinea. Casey Korchefsky 1932: 286 (as synonym of C. longs to C. sanguinea sanguinea Mulsant); Gorham 1892: 170 had distinguished C. hondurasica in his key the presence of a short but distinct (as synonym of C. sanguinea Mulsant); by arc of Blackwelder 1945: 452 (as synonym of oblique line separated from the main C sanguinea Mulsant). the postmetacoxal line. In the case of the Cycioneda polonica Hampe 1850: 357; type specimen, and in other specimens of this condition Crotch 1874: 45 (as synonym of C. san- C. sanguinea which exhibit guinea Mulsant); Korchefsky 1932: 286 (less than 10%), it appears to be primarily slight (as synonym of C. sanguinea Mulsant); a postmortem artifact resulting from a Blackwelder 1945: 452 (as synonym of buckling of the abdomen upon drying. Live C. sanguinea Mulsant). specimens of C scmguinea often have a Cycioneda rubripennis Casey 1899: 92; faint integumental wrinkle in this same po- Korschefsky 1932: 285 (as synonym of sition which corresponds to the area where munda): Mader 1958: 241 (in key); Gor- the hind tarsus comes to rest when the ap- don 1985: 820 (as synonym of sangui- pendages are withdrawn during a death nea). feint. Rarely does this oblique feature have Cycioneda hondurasica Casey 1899: 92; a sharply incised appearance as it does in Mader 1958: 240 (in key) (examined). the related genus Coccinella. No significant New synonym. differences could be found from examining Coccinella reflexa Germain 1854: 333; Bre- the type of C. hondurasica which would thes 1923: 454 (as synonym of C. fulvi- justify maintaining a separate species for pennis Mulsant); Korchefsky 1932: 510 this minor and possibly artificial variant. (as synonym of C. fulvipennis Mulsant); Casey, himself, designated the specimen as Blackwelder 1945: 454 (as synonym of "sanguin-10" and placed it together with C. fulvipennis Mulsant) (examined). New other C. sanguinea in a single unit tray. synonym. Germain (1854) reported the common Neda reflexa: Rivera 1904: 16 (generic re- occurrence of adult and larval Coccinella VOLUME 104, NUMBER 231

17 19 20 21 22

Figs. 17-25. 17-22, Basal lobes of male genitalia, characteristic of different coccinelline genera (diagram- matic). 17, Coccinella transversoguttala Mulsant. 18, Coccinella monticola Mulsant. 19, Mulsantina sp. 20, Olla = sp. 21, Neda sp. 22, Paranedo sp. 23-24, Male genitalia of Cycloneda sanguinea sanguinea. 23, Sipho (sc siphonal capsule). 24, Phallobase (bp = basal piece). 25. Female genitalia of Cycloneda sanguinea sanguinea (inf = infundibulum).

reflexa on fennel {Foeuiculitm vulgare Mill- nis Mulsant, but provided no explanation er) in Santiago, Chile during the month of for his action. Chapin (1969) resurrected February. The consistency of this hostplant the species and transferred it to his new ge- association led Germain to speculate that nus Coccinellifia, indicating important dif- the species is phytophagous; however, his ferences in the size and dorsal color pat- description clearly indicates a glabrous hab- terns for the two nominate species. Al-

itus, and therefore not a member of the only though Chapin never examined the type phytophagous subfamily in the Neotropics: material, he based his concept of C. reflexa Epilachninae. Brethes (1923) placed C re- on specimens taken in Arica, Chile in 1966 flexa as a synonym of Coccinella fulvipen- by Alfonso Aguilera P., which compared in 2

2,1 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON detail with the type description except for a Cycloneda sanguinea limbifer Casey minor discrepancy in size (5 mm in length (Fig. 11) 6.3 These specimens as opposed to mm). Cycloneda limbifer Casey 1899: 92; Zeleny are not to be found in the pinned collection 1969: 333 (biology, toxicology); Hodek at the USNM; however, a slide mount of 1973: pi. XXVII, fig. 3 (4th instar larva the female genitalia of one of the specimens color illustration). was located in Chapin's slide cabinet. These Coccinella (Cycloneda) limbifer: Leng genitalia agree in all particulars with those 1903: 204. of Cycloneda sanguinea, a species which Cycloneda sanguinea ab. limbifera: Kor- ranges in length from 3.2 to 6.5 mm. schefsky 1932: 286 (unjustified emenda- The description of the larval and pupal tion of limbifer, ICZN 1999, Art. 31.2.1- stages of Coccinella reflexa made by Rivera 3.2.2); Mader 1958: 241 (in key). (1904) in Chile (as Neda reflexa (Germain)) Cycloneda sanguinea limbifer: Chapin is also in agreement with both published de- 1949: 23; 1957: 89; Gordon 1985: 823 scriptions (Saini 1985, Phuoc and Stehr (in key, distribution, diagnosis, genitalia, Vandenberg 1974, Gordon and Vandenberg 1993) and adult habitus); Gordon and museum specimens of Cycloneda sangui- 1993: 308 (description of 4th instar larva, larval key). nea. Cycloneda sanguinea is common in collections from Chile, and I do not know Remarks.—The name Cycloneda limbi- of any other very similar species from that fer was applied by Casey (1899) to speci- country. In southern California, where Cy- mens taken from the Bahamas (Egg Island), cloneda sanguinea is common, I often have that differ from most mainland examples of observed adults and larvae together on vol- C sanguinea by the presence of a narrow unteer growths of Foeniculum vulgare, the black border on the outer elytral margins. very habitat specified for C. reflexa in Ger- Authors have assigned variable rank to this main's original description. insular form, from aberration to full spe- The facts outlined above were compel- cies, a problem whose satisfactory solution ling enough for me to propose the synon- may require more sophisticated genetic ymy of C. reflexa even in the absence of studies and laboratory crosses. Chapin (1957) reports C. s. limbifer as widely dis- type material. I eventually had the oppor- tributed in the West Indian islands as far tunity to examine a syntype of Coccinella south as St. Lucia of the Lesser Antilles. reflexa ("CHILE [handwrittenj/Coccinella re- However, not all of the island forms are flexa Germ. [handwritten]/SINTIPO/CHILE strictly comparable. In specimens from M.N.H.N. Tipo No 2158'') from the Museo Cuba, Haiti, Dominican Republic, and Nacional de Historia Natural, Santiago, and Puerto Rico, the elytral border is always confirm my suspicions. It is a representative darkened, and the beetles are further distin- female of Cycloneda sanguinea, with head guished from their mainland relations by and pronotal markings much as in the ex- possessing a shorter body form, a broader ample illustrated (Fig. 9). The specimen is elytral base (relative to pronotal width) and card mounted, and the elytra are slightly a tendency towards dwarfism, particularly parted at the sutural apex, giving an initial in males (Fig. 1 1). If specimens from only impression of less convexity, which is dis- these islands were compared to the main- pelled upon closer examination. The spec- land (Fig. 10), one might conclude that lim- imen is at the upper range of body size for bifer is deserving of full species status. On the species (6.3 mm), but a number of ex- the other hand, specimens from the Baha- amples from Chile and Argentina boast mas are much like those from the mainland similar dimensions. except for the darkened border, and speci- VOLUME 104, NUMBER 1 233

mens from Jamaica appear somewhat inter- included species is nowhere more apparent mediate in body form and the elytral border than in his variable treatment of C. munda varies from black to clear amber. Various which he initially transferred from Cocci- intergrades also occur along the island nella to his newly established genus Cyclo- chains that form the Lesser Antilles, sug- neda (Crotch 1871), placed as a synonym gesting that a step cline or complex of of Cycloneda sanguinea in his revision of closely related species may provide a more the Coccinellidae of the United States accurate model of the Caribbean popula- (Crotch 1873a), omitted in a subsequent tions. checklist of the Coleoptera of America, Chapin (1949) reported two specimens of north of Mexico (Crotch 1873b), and then

C 5. limbifer from Key West, Florida, but resurrected as a valid species and returned it is not clear whether the subspecies is per- to the genus Coccinella in his World revi- manently established there. Although the sion (Crotch 1874). Crotch's vacillating darkened elytral border occurs indepen- perspective was probably due to the con- dently within the nominate subspecies, it flicting impressions provided by C. nmn- does so only rarely in North and Central da^s external color pattern (very like that of American populations. In Southern Brazil, C. sanguinea) and the external topology Chile, and Argentina the extreme outer bor- (more elongated and depressed than most of der is often either a dark reddish amber or its congeners). The similarity in the form of blackish. the genitalia of C. sanguinea and C munda (Gordon 1985) resolve this apparent dilem- Cycloneda munda (Say) ma and support the view of a very close (Figs. 1-3) systematic relationship. Casey (1899) described Cycloneda ater Coccinella munda Say 1835: 202; Crotch (later connected to atra) from an unlabeled 1874: 107. specimen found in the Levette cabinet. Daidis munda: Mulsant 1850: 324. While Casey later expressed some doubt re- Cycloneda munda: Crotch 1871: 6; 1873a: garding the proper generic placement of the 372 (as synonym of C. sanguinea Lin- unusual all black species (Casey 1908), he naeus); Casey 1899: 93; Leng 1920: 216; felt certain it was a member of the North Korschefsky 1932: 284; Timberlake American fauna because the cabinet con- 1943: 23; Wingo 1952: 46; Mader 1958: tained "little or no foreign material." Other 241 (in key); J. Chapin 1974: 63; Phuoc revisions, confined to the Nearctic fauna and Stehr 1974: 58 (pupal description, (Leng 1903, 1920; Gordon 1985), skirted key); Gordon 1985: 820 (in key, distri- the issue of classification by declaring the bution, diagnosis, genitalia, adult habi- specimen to be of probable foreign origin; tus); Gordon and Vandenberg 1993: 308 a not unreasonable assumption considering (description of 4th instar larva, larval the species failed to reappear in any other key). collected samples. Recent dissection and Coccinella {Cycloneda) sanguinea var. examination of the male holotype ("atra munda: Leng 1903: 203. Csy[handwritten]/bequest 1925/TYPE Cycloneda ater Casey 1899: 93; Gordon USNM 35528 [red label]") reveal it to be 1985: 820 (examined). New synonym. a wholly typical example of Cycloneda Cycloneda atra Casey 1908: 405; Leng munda in all respects except for the aber- 1920: 216. rant coloration. Not only are the chitinous Coccinella krikkeni lablokoff-Khnzorian external structures and genitalia deeply pig- 1982: 395 (examined). New synonym. mented, but the internal ligaments and fat Remarks.—Crotch's confusion over the body are sooty and oddly decomposed. taxonomic boundaries of Cycloneda and its Thus, it would seem that the black colora- PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tion is more likely due to some sort of post- Cycloneda nutnda ab. polita: Leng 1920: mortem treatment than an expression of ge- 216; Korschefsky 1932: 285. netics. Backlighting the elytra with a strong Cycloneda polita jiava Timberlake 1943: light reveals a ruddy glow, suggesting the 24. presence of the orange pigmentation found Remarks. Gordon synonymized the in typical examples of the species. — subspecies Cycloneda polita jiava Timber- lablokoff-Khnzorian described Coccinel- lake with the nominate subspecies because la krikkeni from a male specimen deposited it has identical genitalia, and differs only in in Naturalis, Nationaal Natuurhistorisch possessing a paler elytral coloration. Al- Museum, Leiden, and published the de- though I disagree with the reasons for the scription, along with illustrations of the synonymy, the action is still supportable. habitus and male genitalia, in a large vol- Specimens of Cycloneda polita with bright ume on Palearctic and Oriental Coccinelli- orange or scarlet elytra also occur within nae (lablokoff-Khnzorian 1982). While the the specified range of C polita fiava (Ala- genitalia of some CoccineUo species are not meda Co. and Santa Cruz Mountains, Cal- very different from those of Cycloneda, the ifornia); therefore, it is probably little more pronotal color patterns of C krikkeni are than an abenation which occurs at higher highly aberrant for Cocciuella, yet typical frequencies in some areas. of Cycloneda. The genitalia illustrations and description of leg coloration suggest Acknowledgments the species Cycloneda miinda, which is re- stricted to the eastern United States and I am grateful to the following individuals parts of Canada. Subsequent examination of and institutions for their kind assistance: G. the holotype ("Museum Leiden, S. INDIA House, Department of Entomology, Smith- madras State, Coimbatore, 1400 ft, X.1961 sonian Institution, National Museum of P.Susai Nathan/Holotypus Coccinella Krik- Natural History, Washington, DC (Casey keni Khnz [handwritten label]") confirmed collection) (USNM) for access to the ho- the suspected synonymy, but provided no lotypes of Cycloneda atra Casey and Cy- insight as to how it came to bear such an cloneda hondurasica Casey; R. de Jong and unexpected locality label. This unique re- J. van Tol, Naturalis, Nationaal Natuurhis- cord may be the result of accidental trans- torisch Museum, Leiden (RMNH) for loan port through commerce, or, more likely, an of the holotype of Coccinella krikkeni la- accident of mislabeling. blokoff-Khnzorian; M. Elgueta, Museo Na- cional de Historia Natural, Santiago for Cycloneda polita Casey loan of the syntype of Coccinella reflexa (Figs. 4-6) Germain; D. Kavanaugh, Department of Cycloneda polita Casey 1899: 93; Timber- Entomology, California Academy of Sci- lake 1943: 24; Mader 1958: 241 (in key); ences, California for access to type material Hatch 1961: 181; Belicek 1976: 330; of Adalia galapagoensis Van Dyke; R. Gordon 1985: 820 (in key, distribution, Pope, Department of Entomology, The Nat- diagnosis, genitalia, adult habitus); Gor- ural History Museum, London (BMNH) don and Vandenberg 1993: 309 (descrip- and Roger Booth, CAB International Insti- tion of 4th instar larva, larval key). tute of Entomology, London for access to Coccinella {Cycloneda) sanguinea van pol- type material of Cycloneda sanguinea cay-

ita: Leng 1903: 203. niana Casey; J. Obrycki, Department of En- Coccinella {Cycloneda) sanguinea: Palmer tomology, Iowa State University, and D.

1914: 232 (not sanguinea Linnaeus) (de- Nickle, J. Brown, E. Roberts, and R. Gor- scription larval instars, color habitus late don (emeritus). Systematic Entomology instar). Laboratory, Washington, DC for making VOLUME 104. NUMBER 1 235 corrections and supplying useful comments Chapius, F. 1876. Histoire naturelle des insectes. Gen- era des Coleopteres, Paris 12: 1-424. on earlier versions of the manuscript; E. Crotch, G. R. 1871. List of Coccinellidae. Cambridge, Roberts also prepared the genitalia draw- 8 pp. ings, as well as digital photographs and . 1873a. Revision of the Coccinellidae of the sketches used in making the illustrations. United States. Transactions of the American En- tomological Society 4: 363-382.

Literature Cited . 1 873b. Check List of the Coleoptera of Amer-

ica, North of Mexico. Salem, 136 pp. Arioli. M. C. S. 1985. Coccinellini no Rio Grande do . 1874. A Revision of the Coleopterous Family Sul, Brazil (Coleoptera, Coccinellidae). Revista Coccinellidae. London, 311 pp. do Centro de Ciencias Rurais, Brazil 15(1): 5-35. Erichson, W. F. 1842. Beitrag zur Insecten-Fauna von Belicek, J. 1976. Coccinellidae of western Canada and Vandiemensland, mit besonderer Beriicksichti- Alaska with analyses of the transmontane zoogeo- gung der geographischen Verbreitung der Insec- graphic relationships between the fauna of British ten, von Herausgeber. Archiv fur Naturgeschichte Columbia and Alberta (Insecta: Coleoptera; Coc- 8: 238-241. cinellidae). Quaestiones Entomologicae 12: 283- Fabricius, J. C. 1792. Entomologia Sy.stematica. Haf- 409. niae. Vol. 1, part 1, 330 pp., part 2, 538 Berg, F. G. C. 1874. Noticias criticas sobre algunas pp. P. 1854. Descripcion de 21 especies nuevas publicationes entomologicas. Boletin de la Aca- Germain, demia Nacional de Ciencias Exactas Esistente en de Coleopteros de Chile. Anales de la Universi- dad, Chile 11: 326-336. la Universidad de Cordova 1: 274-293. Blatchley, W. S. 1910. An illustrated catalogue of the Gordon, R. D. 1985. The Coccinellidae (Coleoptera) Coleoptera or beetles (exclusive of the Rhynco- of America north of Mexico. Journal of the New phora) known to occur in Indiana. Bulletin, Indi- York Entomological Society 93: 1-912.

ana Department of Geology and Natural Resourc- . 1987. A catalogue of the Crotch collection of

es I: 1-1386. Coccinellidae (Coleoptera). Occasional Papers on Blackwelder, R. E. 1945. Checklist of the coleopterous Systematic Entomology No. 3. London, 46 pp.

of Mexico, Central America, the West In- Gordon, R. D. and N. J. Vandenberg 1993. Larval sys-

dies, and South America. Part III. Bulletin of the tematics of North American Cycloneda Crotch United States National Museum 185: 343-550. (Coleoptera: Coccinellidae). Entomologica Scan- Brethes, J. 1923. Catalogue synonymique des Cocci- dinavica 24: 301-312.

nellides du Chili. Revista Chilena de Historia Nat- Gorham, H. S. 1892. Biologia Centrali- Americana, In-

ural. XXV. 1921(1923): 453-456. secta, Coleoptera, Coccinellidae 7: 161-176. Bruch, C. 1915. Catalogo sistematico de los coleop- Hampe, C. 1850. Beschreibung einiger neuen Kafer- teros de la Repiiblica Argentina. Revista del Mu- Arten. Stettiner Entomologische Zeitung 1 1 : 346— seo de La Plata, pars IX, 19: 346-400. 358. Casey, T. L. 1899. A revision of the American Coc- Hodek, I. et al. 1973. Biology of Coccinellidae. Aca- cinellidae. Entomologi- Journal of the New York demia, Czechoslovak Academy of Sciences, cal Society 7: 71-169. Prague, 260 pp. . 1908. Notes on the Coccinellidae. The Ca- Handlirsch, A. 1906-1908. Die fossilen Insecten und nadian Entomologist 40: 393-421. die Phylogenie der rezenten Formen. Ein Hand- Chapin, E. A. 1941. Lady-beetles belonging to the ge- buch fiir Paliiontologen und Zoologen. Engel- nus Procida Mulsant. Memorias de la Sociedad mann, Leipzig, 1430 pp. Cubana de Historia Natural "Felipe Poey" 15: Hatch, M. H. 1961. Beetles of the Pacific Northwest, 165-168. part III: Pselaphidae and Diversicornia. University . 1949. New beetle records for Florida. Cole- of Washington Press, Seattle, 503 pp. opterists' Bulletin 3: 23. lablokoff-Khnzorian, S. M. 1990. About the classifi- . 1957. Records of coccinellid beetles from the cation of the Coccinellini. Coccinella 2: 58-60. Cayman Islands, with descriptions of new species

. 1982. Les Coccinelles, Coleopteres-Coccinel- from the West Indies. The Entomologist's Month- lidae, Tribu Coccinellini des regions Palearctique ly Magazine 93: 89-91. et Orientale. Paris, 568 pp. . 1969. New synonymy and generic reassign- International ment in South American Coccinellina (Coleop- Commision of Zoological Nomenclature. tera: Coccinellidae). Proceedings of the Entomo- 1999. International code of zoological nomencla- logical Society of Washington 71(3): 467-469. ture, fourth edition. London, 306 pp.

Chapin, J. B. 1974. The Coccinellidae of Louisiana Korschefsky, R. 1932. Coleopterorum Catalogus, Pars (In.secta: Coleoptera). Bulletin of the Louisiana 120, Coccinellidae II. Berlin. 659 pp.

State University Experiment Station 682: 1-87. Leng, C. W. 1903. Notes on Coccinellidae. II. Journal :

236 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of the New York Entomological Society I 1: 193- . 1992. Revision of the New World lady beetles 213. of the genus OUa and description of a new allied

. 1920. Catalogue of the Coleoptera of Amer- genus (Coleoptera: Coccinellidae). Annals of the

ica, North of Mexico. Mount Vernon, New York, Entomological Society of America 85: 370-392.

470 pp. . 1997. Beetle economics. //; Hoagland, K. E. Linnaeus, C. 1763. Centuria In.sectorum Rariorum. and A. Y. Rossman. eds. Global Genetic Resourc- Uppsala, 32 pp. es: Access, Ownership, and Intellectual Property Mader, L. 1958. Die amerikanischen Coccinelliden der Rights. Association of Systematics Collections. Gruppe Synonychini. Annalen des Naturhisto- Washington, 348 pp. rischen Museums in Wien 62: 236-249. Vandenberg, N. and R. D. Gordon. 1988. The Cocci-

Mulsant, M. E. 1850. Species des Coleopteres trimeres nellidae (Coleoptera) of South America, part I. A securipalpes. Annales des Sciences Physiques et revision of the genus Erylhroueda Timberlake,

Naturelles, d" Agriculture et d'lndustrie, Lyon 2: 1943. Revista Brasileira de Entomologia 32: 31- 1-1104. 43.

. 1866. Monographic des coccinellides. Me- . 1996. A new genus of Neotropical Coccinel- moires de I'Academie Sciences, Belles-lettres et lini (Coleoptera: Coccinellidae) related to OUa Arts de Lyon 15: 1-112. Casey and allies. Proceedings of the Entomolog-

Palmer. M. A. 1914. Some notes on life history of ical Society of Washington 98(3): 541-550. lady-beetles. Annals of the Entomological Society Van Dyke, E. 1953. The Coleoptera of the Galapagos of America 7: 213-238. Islands. Occasional papers of the California Acad- Phuoc, D. T. and F W. Stehr. 1974. Morphology and emy of Sciences, XXII, 181 pp.

taxonomy of the known pupae of Coccinellidae Weise, J. 1898. Coccinellen aus Siidamerika. Deutsche (Coleoptera) of North America, with a discussion Entomologische Zeitschrift 1898: 125-126.

of phylogenetic relationships. Contributions of the . 1902. Coccinelliden aus Siidamerica. III. American Entomological Institute 10(6): 1-126. Deutsche Entomologische Zeitschrift 1902: 161-

Rivera, M. J. 1904. Biolojia des eos coleopteros Chi- 176.

lenos cuyas larvas atacan al trigo. Revista Chilena . 1904a. Coccinellidae in Argentina, Chili et

de Historia Natural 8: 241-254. Brasilia e collectione domini Caroli Bruchi. Re- Saini, E. D. 1983. Clave para estadios larvales de Coc- vista del Museo de La Plata 11: 193-198.

cinelidos. Revista de la Sociedad Entomologica . 1904b. Synonymische Bemerkungen zu Gor- Argentina 42: 397-403. ham, Biologia Centrali-Americana, vol. VII, Coc-

. 1985. Identification practica de "vaquitas" cinellidae. Deutsche Entomologische Zeitschrift

beneficas. Parte I. Departamento Publicaciones, 1904: 433-452.

Prensa y Difusion, Instituto Nacional de Tecnol- . 1906. Hispinae, Coccinellidae et Endomychi- ogia Agropecuaria, 22 pp. dae Argentinia et vecinitate e collectione Bruchi- Say, T. 1835. Descriptions of new American coleop- ana. Revista del Museo de La Plata 12: 219-231.

terous insects, and observations on some already . 1922. Coleoptera e collectione Bruchiana. An-

described. Boston Journal of Natural History 1 ales de la Sociedad Cientffica Argentina 94: 30- 151-203. 40.

Timberlake, P. H. 1943. The Coccinellidae or ladybee- Wilson, J. W. 1926. The genitalia of some of the Coc-

tles of the Koebele collection. Part I. Bulletin of cinellidae. Journal of the Elisha Mitchell Scientific the Experiment Station of the Hawaiian Sugar Society 42(1 & 2): 63-74. Planters' Association, Entomological Series 22: Wingo, C. W 1952. The Coccinellidae (Coleoptera) of 1-67. the upper Mississippi Basin. Iowa State Journal of

Vandenberg, N. J. 1987. A systematic study of Olio Science 27: 15-53.

Casey and allied genera of the New World (Co- Zeleny, J. 1969. A biological and toxicological study leoptera: Coccinellidae), Ph.D. dissertation. Uni- of Cycloneda liinbifer Casey (Coleoptera. Cocci- versity of California, Berkeley (unpublished for nellidae). Acta Entomologica Bohemoslovaca 66: nomenclatorial purposes). 333-344.