Phylogenetic Analysis of Cyanophrys Clench, a Synopsis of Its Species, and the Potentially Threatened C

Home , Callophrys, Cyanophrys, Cyanophrys herodotus, Cyanophrys miserabilis, Eumaeini, Incisalia

4 April 2005 PROC. ENTOMOL. SOG. WASH. 107(2), 2005, pp. 398-416

PHYLOGENETIC ANALYSIS OF CYANOPHRYS CLENCH, A SYNOPSIS OF ITS SPECIES, AND THE POTENTIALLY THREATENED C. BERTHA (JONES) (LYCAENIDAE: THECLINAE: EUMAEINI)

ROBERT K. ROBBINS AND MARCELO DUARTE

(RKR) Department of Entomology, RO. Box 37012, NHB Stop 127, Smithsonian In- stitution, Washington, DC 20013-7012 U.S.A. (e-mail: [email protected]); (MD) Coleçâo de Lepidoptera, Museu de Zoología, Universidade de Sao Paulo, Av. Nazaré 481, 04263-000 Sao Paulo SP Brasil (e-mail: [email protected])

Abstract.•A species level phylogenetic analysis of Cyanophrys Clench was performed using 14 characters of the frons, wing venation, wing shape, androconia, and genitalia. There were 15 most parsimonious cladograms, and a successive weighting iteration re- sulted in three of these 15. Cyanophrys is characterized in this paper by two hypothesized synapomorphies (an expanded hindwing anal lobe that is tan-brown and a pair of dorsal brush organs), and adults can be readily identified in the field. Cyanophrys has been divided into Plesiocyanophrys K. Johnson, Eisele and MacPherson, Apophrys K. Johnson and Le Crom, Antephrys K. Johnson, Eisele and MacPherson, Mesocyanophrys K. John- son, and Cyanophrys (as delimited in Johnson and Le Crom 1997a), but only the monotypic Antephrys is monophyletic on the most parsimonious cladograms. A synopsis of Cyanophrys species includes notes on their distribution, habitat, identification, nomenclature, larval food plants, and male behavior. Cyanophrys bertha, which occurs in moist evergreen and seasonally deciduous forests in the mountains of southern Brazil from 800 to 1,400 m elevation, has been proposed for threatened status. It is "cladistically distinct" {sensu Vane-Wright et al. 1991) in that its sister is a lineage of five montane species or seven primarily lowland species. Key Words: Callophrys, phylogeny, venation, genitalia, androconia, brush organs

Generic taxonomy of the primarily Neo- (Brown 1993, Brown and Freitas 2000 and tropical Cyanophrys Clench (Theclinae: references therein. Otero et al. 2000). More Eumaeini) has been unstable (Robbins recently, it has been listed as "almost 2004b). Clench (1961) described it as a threatened" (Mielke and Casagrande 2004). subgenus of the Holarctic Callophrys West- Little is known about C. bertha, and it is wood, but since 1993, Cyanophrys has been unclear which species, or group of species, divided into five genera and subgenera, pri- is its closest relative. Such information marily on the basis of differences in geni- might provide clues to its biology. Further, talia and color of the frons (Johnson et al. phylogenetic position is a factor to be con- 1993, Johnson and Le Crom 1997a). sidered in assessing the status of threatened While most of the 16 Cyanophrys species species (Vane-Wright et al 1991). are common and widespread, the Brazilian The first purpose of this paper is to assess C. bertha (Jones) is exceedingly rare and the monophyly of the genera and subgenera has been proposed for "vulnerable" status into which Cyanophrys has been parti- VOLUME 107, NUMBER 2 399

tioned. To accomplish this goal, we code though identification of Cyanophrys species interspecific morphological variation to in- was not a goal of this study, the coded char- fer phylogenetic relations among Cyano- acters can be used to identify most species. phrys species. Besides frons color and gen- And in the species synopsis, we note infor- italia, we code characters of the male fore- mation on the distribution, habitat, identi- wing venation, androconia, brush organs fication, nomenclature, behavior, larval {sensu Eliot 1973), and wing shape. food plants, and published illustrations of Johnson and co-authors have proposed adult Cyanophrys species. We summarize 433 new species names in the Eumaeini and assess what we know about C. bertha during the past three decades, including 31 in light of the phylogenetic results. in Cyanophrys (Robbins 2004a,b), but have not been careful. The adults and genitalia MATERIALS AND METHODS of many types are different sexes or species (Robbins and Lamas 2002). Data labels on Genitalia, androconia, and venation were types appear to have been switched (Rob- examined by standard techniques (Robbins bins and Nicolay 1999, Robbins and Lamas 1991) using the approximately 1,075 spec- 2002). In one case, the two adult types of imens of Cyanophrys in the National Mu- one species from different collections were seum of Natural History (USNM) as well similarly glued parts of species in different as others borrowed from the American Mu- genera (Robbins and Lamas 2002). Finally, seum of Natural History (AMNH) in New a number of types cannot be found (Rob- York, the Illinois Natural History Survey bins and Nicolay 2002, G. Lamas, personal (INHS) in Champaign-Urbana, The Natural communication). History Museum (BMNH) in London, and The publications of Johnson and co-au- the Universidade Federal do Paraná (UFPR) thors on Cyanophrys contain similar prob- in Curitiba. Wing venation illustrations lems. For example, Johnson and Le Crom were made by digital scanning of wing (1997b:23) designated a neotype for Papi- slides. We examined 82 male and 62 female lio amyntor Cramer 1775 from Surinam and genitalic preparations. Genitalic terms fol- deposited it in the Natural History Museum low those in Klots (1970), wing vein names (London). However, the labeled neotype de- follow Nicolay (1971, 1977), and andro- posited in this collection has data labels conial terminology follows Robbins (1991). from French Guiana. Another specimen The terminal taxa are C. goodsoni s with their neotype label in the American (Clench), C. argentinensis (Clench), C ber- Museum (New York) has a Surinam label, tha, C acaste (Prittwitz), C amyntor (Cra- but is a different species than the designated mer), C fiisius (Godman and Salvin), C. neotype in London. A second representative herodotus (Fabricius), C. miserabilis (Hew- example of their lack of care is outlined in itson), C. velezi K. Johnson and Kruse, C. the synopsis below under Cyanophrys ro- crethona (Hewitson), C. longula (Hewit- raimiensis K. Johnson and D. S. Smith. Al- son), C. pseudolongula (Clench), C agri- though we plan to address the neotype des- color (Butler and H. Druce), C banosensis ignation elsewhere, correcting all problems (Clench), and C remus (Hewitson). Cyano- A l^As^'^.d^ created by Johnson and co-authors in Cy- phrys roraimiensis K. Johnson and D.S. anophrys would be a Sisyphean task that Smith is omitted because it is known only could delay publication of our current re- from the holotype female, but we discuss sults for years. We follow the classification its likely phylogenetic placement on the ba- in Robbins (2004b). sis of structures in the female type. As out- The second purpose of this paper is to groups, we chose three species from three present a synopsis of Cyanophrys species, other subgenera of Callophrys recognized •X •'-.'•A,--.' with special emphasis on C. bertha. Al- by Clench (1961); Callophrys (Callophrys) 400 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Data matrix for Cyanophrys. The three Callophrys species are the outgroups, and the characters and their states are detailed in the text.

Characters Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14

Callophrys rubi 1 0 0 0 0 0 0 0 0 0 ? ? 7 ? C. gryneus 0 0 0 0 0 0 0 0 0 0 ? ? 7 ? C. niphon 0 0 0 0 0 0 0 0 1 0 ? ? 7 7 Cyanophrys goodsoni 1 0 0 0 0 0 0 0 0 0 0 0 C. argentinensis 1 0 1 0 0 0 0 0 0 0 0 C. bertha 0 0 0 0 0 2 0 0 1 0 0 C. acaste 1 0 1 1 0 2 0 0 1 0 0 C. amyntor 1 0 0 1 0 2 0 0 1 0 0 C. fusius 1 0 2 7 2 0 0 1 0 0 C. herodotus 1 0 1 1 2 1 0 0 0 0 C. miserabilis 0 0 1 1 2 1 0 0 0 0 C. velezi 0 0 1 1 2 1 0 0 0 0 C. crethona 0 0 1 1 2 1 0 0 0 0 C. longula 0 0 0 0 2 0 0 C. pseudolongula 0 0 0 0 2 0 0 C remus 0 0 0 0 2 0 0 C. banosensis 0 0 0 0 2 0 0 C. agricolor 0 0 0 0 1 0 0

rubi (L.), C. (Mitoura) gryneus (Hübner), Distributional information for the other spe- and C. (Incisalia) niphon (Hübner). cies is based on specimens from many mu- We coded 14 characters (Table 1) and seums and from the literature. Records of used a question mark (?) for inapplicable larval food plants without citation are based states and for one case of an intraspecific on Janzen and Hallwachs (2004) or data dimorphism. All multi-state characters were compiled by Robbins. Plant family names treated non-additively (unordered). We used follow those in Willis (1973). Records of the implicit enumeration option of Hen- "territorial" behavior are based on unpub- nig86 software to derive most parsimonious lished data compiled by Robbins. Type lo- cladograms. A strict consensus tree was de- calities are listed in Robbins (2004b), and termined. To test the assumption of equally citations for original descriptions can be weighted characters, a successive weighting found in Lamas (1995). iteration was performed (Farris 1969), and a consensus of the resulting trees was de- MORPHOLOGY AND CODED CHARACTERS termined. Mapping of characters on trees Head.•Presence or absence of iridescent was done with Winclada software (Nixon green scales on the frons is intraspecifically 2002) using the "unambiguous changes consistent. only" optimization option. Jackknife sup- Character 1: Iridescent green scales on ports were determined in Winclada using frons (0) absent, (1) present. Nona (1000 replications with mult* 10, This character needs to be used with cau- memory 1000 trees). tion because exposure to humidity, partic- The synopsis contains additional detail ularly during preparation, or to physical for C. bertha because of its conservation abrasion can change scale color from green interest. The distribution of C bertha is to brown. Iridescent butterfly scales return based on 13 museum specimens (Appen- to their original color after drying (Nijhout dix). Its morphology is based on 6 individ- 1991), but for reasons currently unknown, uals in the USNM collection (Appendix). this is generally untrue for hairstreaks VOLUME 107, NUMBER 2 401

Fig. 1. Dorsal (top) and ventral wings of male (left) and female Cyanophrys bertha. Scale 1 cm.

(Robbins, unpublished). Also, green scales and Ml (0) connected by vein UDC, which on the frons may be rubbed off of worn may be poorly developed (Figs. 3, 5-7), (1) individuals. fused at the origin of vein Ml (vein UDC Wing pattern.•We were unable to code absent) (Fig. 4). Males of C. pseudolongula wing pattern variation in Cyanophrys for a aie coded with a question mark because number of reasons. Presence of a postme- they are geographically variable for fore- dian line on the ventral forewing varies in- wing venation, with both states occurring in terspecifically and intraspecifically among males from Ecuador and northern Peru east species. A dark maroon band along the sub- of the Andes. margin of the ventral hindwing and dark Character 3: Male forewing veins R2 brown scales at the base of the ventral and R3 (0) arise from the discal cell (Figs. hindwing are conspicuous wing pattern el- 3-4, 6), (1) are stalked (Fig. 5). ements in some Cyanophrys species, but the Androconia.•There are three major an- dark maroon and brown scales occur in all droconial patterns. Males of some Cyano- Cyanophrys and the extent of their expres- phrys species have two clusters of andro- sion shows almost continuous variation conia; one at the base of veins R2, R3, and among species and sometimes within a spe- Ml and one along the base of vein M3 cies. (Figs. 4-5). Other species have androconia Male wing venation.•Male forewing only at the base of veins R2, R3, and Ml venation varies within Cyanophrys, and we (Figs. 3, 6-7), although this cluster is some- code it as two characters. times small, as in C. remus. Finally, males Character 2: Male forewing veins R3 of C. fusius are unique among Cyanophrys 402 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

60° 50° 40° 30° Fig. 2. Distribution of Cyanophrys bertha in southern Brazil from 13 museum specimens (Appendix). in having brown dorsal forewings (irides- cause it can vary between males and fe- cent blue in the others) that lack androcon- males (C. languid), geographically (C Her- ia. Because it lacks androconia, C. fiisius odotus, C. acaste), or at one locality (C. was coded ? (inapplicable) for characters amyntor, C. herodotus). We also did not 5-6. code wing shape because there appears to Character 4: Male dorsal forewing (0) be a continuum of variation from the round- with androconia only at the base of veins ed wings of C. goodsoni to the angular R2, R3, and Ml (Figs. 3, 6-7), (1) with wings of C agricolor. The character below, androconia at the base of veins R2, R3, and however, is distinct and unambiguously Ml and along the base of vein M3 (Figs. scored. 4•5), (2) lacking androconia. Character 7: Hindwing anal lobe (0) not Character 5: Dorsal forewing androcon- produced posteriorly (<0.1 mm beyond the ial cluster at the base of veins R2, R3, and outer margin of cell Cu2-2A) (Fig. 7), (1) Ml (0) extends into cell R2-R3 (Figs. 6-7), produced posteriorly, at least 0.25 mm be- (1) absent from cell R2-R3 (Fig. 3-5). yond the outer margin of cell Cu2-2A Character 6: Basal edge of dorsal fore- (Figs. 3, 6). wing androconial cluster at the base of Male genitalia.•We found considerable veins R2, R3, and Ml (0) located at, or just interspecific variation in the male genitalia basal to, the origin of vein R2 (Figs. 3, 5- and its associated dorsal brush organs. Var- 7), (1) extending basally well beyond the iation of some structures, such as length of origin of vein R2 to, or almost to, the origin the saccus (Table 2), phallobase (internal of vein Rl (Fig. 4). part of the penis), entire penis, and tegu- Wing shape and hindwing tails.-Despite men, was not phylogenetically useful be- variability in wing shape and number of cause differences in means between species hindwing tails, we code only one such char- were small (typically less than one standard acter We did not code number of tails be- deviation), which made it difficult to rec- VOLUME 107, NUMBER 2 403

R3 R2 RI Ml R3 R2 RI

Figs. 3-5. Forewing and hindwing venation witli detail of androconial clusters. 3, Cyanophrys bertha. Arrow at top points to the upper discocellular vein (UDC), which arises distal of the origin of vein R2. Arrow at bottom points to the posteriorly produced hindwing anal lobe. 4, C. herodotus. Detail shows anastomosis of veins R3 and Ml; there is no upper discocellular vein. Arrow points to androconia along the base of vein M3. 5, C. acaste. Veins R2 and R3 are stalked. Arrow points to androconia along the base of vein M3. The upper discocellular vein arises basal of the origin of vein R2. ognize distinct states. We were able to code Character 8: Dorsal male genitalia brush other aspects of interspecific variation. Be- organs (0) absent, (1) present (Figs. 8-10). cause they lack brush organs, the outgroup Character 9: Male genitalia valvae (0) species were coded? (inapplicable) for char- with a terminal thickening that extends acters 11-14. along the inner margin of the valva (Fig. 3 404 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Mean length in mm of the saccus mea- posterior ends of the dorsal male genitalia sured in ventral aspect along the sagittal plane. Species brush organs in lateral aspect (0) less than are listed in ascending rank order with differences be- 1.3 mm long (Figs. 9-10), (1) more than tween succeeding species typically less than a standard 1.4 mm long (Fig. 8). deviation. Character 13: Anterior end of the dorsal Standard male genitalia brush organs originates in Mean Length Deviation Sample Size Species lateral aspect (0) near or below the origin C. acaste 0.53 0.050 7 of the vinculum strut (Figs. 8-9), (1) pri- C. remus 0.54 0.036 5 marily above the origin of the vinculum 0.61 0.086 5 C. amyntor strut (Fig. 10). C. agricolor 0.64 0.042 4 C. pseudolongula 0.68 0.036 3 Character 14: Width of the dorsal male C. longula 0.69 0.035 4 genitalia brush organs (dorsal aspect) at its C. bertha 0.72 0 2 widest point (0) less than 0.5 mm (Figs. 8- C. herodotus 0.74 0.062 7 9), (1) more than 0.5 mm (Fig. 10). 0.051 2 C. velezi 0.79 Female genitalia.•Although length and C. goodsoni 0.79 0.204 2 C. fusius 0.81 0.091 4 width of the ductus bursae, development of C. argentinensis 0.82 0.062 4 sclerotized plates surrounding the ostium C. banosensis 0.83 0 1 bursae, and length of the spines of the signa C. miserabilis 0.86 0.030 5 vary in Cyanophrys, we did not code this 0.040 2 C. crethona 1.35 variation. For the ductus bursae, mean lengths and widths for species were similar, which made it difficult to delineate distinct in Wairen and Robbins 1993), (1) with a states. Similarly, we were unable to char- strongly demarcated terminal thickening acterize distinct states among species in de- that does not extend along the inner margin velopment of the lamellae vaginalis or of the valva (Fig. 10, Fig. 3 in Warren and spines of the signa. In brief, interspecific Robbins 1993), (2) with no terminal thick- variation in the female genitalia was not ening ("not capped" in Clench 1961) (Figs. phylogenetically informative even though it 8-9). was sometimes useful for distinguishing Clench (1961) reported valva tips with a species, such as C. herodotus and C. amyn- terminal thickening in subgenera Incisalia, tor. The bursa copulatrix of C. bertha is Sandia Clench and Ehrlich, and Xamia illustrated (Fig. 11). Clench. This valva tip was illustrated by PHYLOGENETIC ANALYSES AND RESULTS Warren and Robbins (1993), and also occurs in C. agricolor. A less prominent Analysis of the character matrix with the thickening that extends along the inner mar- Hennig86 "ie*" option, which searches ex- gin of the valvae occurs in subgenera Cal- haustively for the most parsimonious clad- lophrys and Mitoura (Warren and Robbins ograms, yielded 15 equally parsimonious 1993) as well as C. goodsoni and C. argen- 23-step trees with a consistency index of tinensis. 0.69 and retention index of 0.86. The equal- Character 10: Posterior-ventral process ly parsimonious trees differed primarily in of vinculum-tegumen (0) less than 0.15 mm the placement of C. bertha and C. fusius, in length (Figs. 8, 10), (1) with a long (> with C. bertha being the sister of either the 0.15 mm) pointed process (Fig. 9). C. longula group (C. longula, C pseudo- Character 11: Anterior end of the setae longula, C agricolor, C. banosensis, and C. comprising the dorsal male genitalia brush remus) or C. herodotus group (C acaste, organs oriented (0) dorso-posteriorly (Figs. C amyntor, C. fusius, C. herodotus, C. mis- 8, 9), (1) horizontally (Fig. 10). erabilis, C. velezi, and C. crethona). This Character 12: Distance from anterior to result is represented in the consensus tree VOLUME 107, NUMBER 2 405

("nelsen" option) as a trichotomy among MacPherson, Antephrys K. Johnson, Eisele C. bertha, the C. longula group, and the C. and MacPherson, Apophrys K. Johnson and herodotus group, but the latter group is not Le Crem, and Mesocyanophrys K. Johnson strongly supported (Fig. 12 with jackknife (Johnson et al. 1993, Johnson and Le Crom values). The consensus tree is not one of 1997a). The species that belong to each is the 15 most parsimonious trees. noted (Fig. 12). Subgenus Cyanoprhys con- A successive weighting iteration (Farris tains the C. longula group (as denoted 1969) produced three trees, each of which above), C. miserabilis, C. velezi, C. cre- was one of the 15 original most parsimo- thona, and C roraimiensis (the latter omit- nious ones. Wing venation and frons color ted from the phylogenetic analysis) (John- (characters 1-3) were weighted four or less son and Le Crom 1997a). This grouping is out of 10 while the other characters (4-14) polyphyletic in the 15 original most parsi- were fully weighted (10 out of 10) except monious cladograms, as summarized by the for "capped" valvae (character 9) and consensus tree (Fig. 12). Subgenus Plesio- brush organ length (character 12). In other cyanophrys contains C goodsoni and C. ar- words, androconia and male genitalia char- gentinensis (Johnson et al. 1993), a group- acters were less likely to be homoplastic ing that is paraphyletic in the 15 most par- than those of wing venation and frons color simonious trees. Subgenus Antephrys con- Carpenter (1988) argued that these trees are tains only C. fusius (Johnson et al. 1993, the best phylogenetic hypotheses. The con- Johnson and Le Crom 1991 SL, Johnson and sensus of the three successively weighted Amarillo 1997, Robbins 2004b), making it trees (24 steps) shows a sister relationship a monotypic genus. Subgenus Mesocyano- between C. bertha and the C. herodotus lin- phrys contains C acaste and C. bertha eage, and there is greater resolution within (Johnson and Le Crom 1997a), a grouping the C. herodotus group (Fig. 13) than in the that is paraphyletic or polyphyletic in the original consensus tree (Fig. 12). original 15 most parsimonious trees. Final- ly, subgenus Apophrys contains C. herod- MONOPHYLY OF GENERA AND SUBGENERA otus and C. amyntor (Johnson and Le Crom Hypothesized synapomorphies for Cy- 1997a), a grouping that is not monophyletic anophrys are (1) paired dorsal brush organs in the 15 most parsimonious cladograms. on the intersegmental membrane between the male genitalia vinculum and the 8* ab- SYNOPSIS OF SPECIES dominal tergum (Character 8, Figs. 8-10) Cyanophrys goodsoni (Clench, 1946) and (2) an elongate hindwing anal lobe with tan-brown coloration (Character 7, Figs. 3, Distribution, habitat, and abundance.• 6). These synapomorphies are unique in the Usually uncommon in seasonally dry hab- Callophrys Section (Robbins 2004b), but itats from southern Texas (United States) to occur in other sections of the Eumaeini. Guanacaste (Costa Rica). However, the green underside of the wings Larval food plants.•Blossom buds of coupled with the elongate hindwing anal Rivina (Phytolaccaceae) in Texas and on lobe allow adult Cyanophrys to be identi- Vernonia (Compositae) in Costa Rica. fied in the field. These results do not pro- Illustrations of adults.•Both sexes in vide data on whether the Holarctic mem- Scott (1986). bers of Callophrys {sensu Clench 1961) form a monophyletic group. Cyanophrys argentinensis (Clench, 1946) Cyanophrys as characterized in this paper Distribution, habitat, and abundance.• has been partitioned into subgenera (some- Uncommon to rare in both dry and wet for- times treated as genera) Cyanophrys, Ple- ests from sea level to over 1,000 m eleva- siocyanophrys K. Johnson, Eisele and tion from eastern Panama (Darien) and 406 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

s'^¿

\(f -y. "^ J

Figs. 6-7. Forewing and hindwing venation with detail of androconial clusters. 6, Cyanophrys goodsoni. Arrow at top points to the upper discocellular vein. Arrow at bottom points to the posteriorly produced hindwing anal lobe. 7, Callophrys rubi. Arrow at top points to the upper discocellular vein. Arrow at bottom points to the hindwing anal lobe that is not posteriorly produced.

northern Venezuela (Aragua) south along (Panama, Venezuela) to 0.25 mm (Peru) to the Andes to northwestern Argentina, then 0.20 mm (Argentina). eastward to Paraguay and southern Brazil. Male behavior.•A "territorial" male was Variation•Width of the ductus bursae in recorded at about 1400 hours on a hilltop, the middle in ventral aspect appears to vary Illustrations of adults.•Both sexes in geographically, from more than 0.30 mm D'Abrera (1995). VOLUME 107, NUMBER 2 407

Figs. 8-9. Male genitalia in lateral (left) and ventral aspect. 8, Cyanophrys bertha. B-brush organs, P-penis, S-saccus, T-tegumen, V-valvae, Vi-vinculum (which is fused with the tegumen in all eumaeines). 9, C. herodotus. The arrow points to the ventro-lateral process of the tegumen, which is foreshortened in this aspect.

Cyanophrys bertha (Jones, 1912) Distribution.•Coastal mountains of southern Brazil from about 800 to 1,400 m elevation in the states of Minas Gerais, Sao Paulo, Paraná, and Santa Catarina (Fig. 2, Appendix). There are literature records from Rio de Janeiro (Brown 1993) and Rio Grande do Sul (Draudt 1919-1920), but we Icnow of no extant specimens to verify the occurrence of C. bertha in these states. The elevation of the locality where one specimen was supposedly collected is 75 m, but it is probably mislabeled (discussed in Ap- pendix). Habitat.•The habitat of C. bertha is var- ied. Holdridge's (1947) ecological life zones of the localities in the Appendix include subtropical lower montane moist forest, subtropical moist forest, and warm temperate moist forest (Tosi 1983). Vegetation zones include dense evergreen forest, Fig. 10. Mdie gea&aXia of Cyanophrys agricolor in lateral (left) and ventral aspect. The arrow (right fig- mixed evergreen forest, and seasonally de- ure) points to terminal thickening of valvae. ViS-vin- ciduous forest (IBGE 1993). Although C. culum strut, which is an internal ridge. bertha appears to be primarily a resident of 408 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 11. Female genitalia bursa copulatrix of Cyanophrys bertha in ventral aspect. moist coastal mountains, three specimens Variation.•Variation of the ventral wing found at Barbacena (Minas Gerais) indicate pattern, particularly expression of the post- that it also occurs, at least on occasion, in median line and a white hindwing discal semi-deciduous forest and may possibly be bar, accounts for this species being named more widespread inland than is currently repeatedly, usually as a subspecies. Individ- realized. uals from the southern temperate parts of Identification.•The unique ventral wing the range are tailed (C. acastoides pheno- pattern of C. bertha (Fig. 1) has distin- type). guished this species since it was described. Larval food plants.•Chuquiraga (Com- Larval food plants.•The larval food positae) in Brazil. plant and immature stages of C. bertha are Male behavior.•Males "perch" on hill- unknown. Larvae of other Cyanophrys have tops from about 1100 to 1500 hours. been reared on flowers, fruits, or young Illustrations of adults.•Both sexes in leaves of plants in families Anacardiaceae, D'Abrera (1995). Boraginaceae, Dipsacaceae, Compositae, Leguminosae, Malvaceae, Phytolaccaceae, Cyanophrys amyntor (Cramer, 1775) Sambucaceae, Sterculiaceae, Ulmaceae, and Distribution, habitat, and abundance.• Verbenaceae. Male behavior.•Brown (1993) observed Widespread and common from sea level to males of C. bertha setting up mating terri- about 1,100 m from northern Mexico to tories in the crowns of trees on hilltops at southern Brazil. Recorded from Texas, Serra do Japi (cf. Appendix) in the early United States (Kendall and McGuire 1984), afternoon. Other than C. velezi, whose based on a female in the Illinois Natural males set up mating territories on hilltops History Survey. Johnson and Le Crom before 0930 hours, the afternoon male "ter- (1997b) stated that it was a misidentified ritorial" behavior of C. bertha is typical for female of C herodotus. Robbins (unpubl.) the genus confirmed their identification. Illustrations of adults.•Fig. 1. Variation.•Individuals in the southern parts of its range may have or lack tails, but Cyanophrys acaste (Prittwitz, 1865) do not differ otherwise. The name Cyano- Distribution, habitat, and abundance.•A phrys caramba (Clench) refers to tailless very common species in a variety of habi- individuals. tats and elevations from southern Brazil to Identification.•Slightly larger on aver- eastern Bolivia, south to central Argentina age than sympatric C. herodotus, but males and Uruguay. are distinguished by different androconial VOLUME 107, NUMBER 2 409

Callophrys (Callophrys) rubi

1 Callophrys (Mitoura) gryneus h-O- 0 • Callophrys (Incisalia) niphon 1 C. (Plesiocyanophrys) goodsoni 3 r-0- C. (Plesiocyanophrys) argentinensis 1 -•-• (75) 1 1 C. (Mesocyanophrys) bertha Cyanophrys 2 1•0 C. (Cyanophrys) longula (54) 0 C. (Cyanophrys) pseudolongula 2 11 13 14 • • • (64) C. (Cyanophrys) remus 1 1 1 1 C. (Cyanophrys) banosensis • C. (Cyanophrys) agricolor 3 1 r-O • C. (Mesocyanophrys) acaste C. (Apophrys) amyntor 2 4 C. (Antephrys) flisius 1 2 C. (Apophrys) herodotus 2 6 10 12 .«-•- • C. (Cyanophrys) miserabilis 1 1 1 L_0_ • C. (Cyanophrys) velezi 0 C. (Cyanophrys) crethona Fig. 12. Strict consensus of the fifteen equally most parsimonious cladograms (23 steps, ci = 0.69, ri = 0.86) for Cyanophrys species. Callophrys rubi, C. gryneus, and C niphon (top) are outgroups. Character numbers are placed above nodes and character state numbers below nodes. Open circles represent reversal or convergence of the character state at that node. Jackknife values are noted in parentheses. Subgeneric placements for Cyano- phrys from Johnson et al. (1993), Johnson and Le Crom (1997a), and Johnson and Amarillo (1997) patches on the dorsal fore wing. Females of Cyanophrys fiisius the two have different genitalia. (Godman and Salvin, 1887) Larval food plants.•Reared from Ul- Distribution, habitat, and abundance.• maceae {Celtis, Trema) in Mexico, Ecuador, Seasonally dry habitats from northern Mex- and Brazil (Kendall 1975, Hoffmann 1937), ico to Colombia and Venezuela (Clench and from Verbenaceae (Clerodendron) in 1946). Trinidad. Identification.•Males are brown above, Male behavior.•Males set up mating ter- not blue, and lack forewing androconia, ritories along trail edges and on hilltops both unique traits in Cyanophrys. Clench from about 1200 to 1600 hours. (1946) noted the similarity of females of Illustrations of adults.•Male in D'Abrera this and the previous species, but female C. (1995), but identity of the female that he fusius have a short white-tipped tail at the illustrated as this species is unclear. end of vein Cul while Central American 410 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

longula

pseudolongula

Fig. 13. Strict consensus of the three most parsimonious successively weighted trees for Cyanophrys species. Characters of wing venation and frons color were under-weighted because they were homoplastic. females of C. amyntor have only a black Variation.^Size and wing pattern of this stub at the end of vein Cul. species are highly variable, with the largest Larval food plants.•Reared from Arra- individuals being most frequent in the bidaea (Bignoniaceae) in Costa Rica. southern part of the range. Individuals from Male behavior.•A "territorial" male northwestern Peru, western Ecuador and was collected on a ridge top at 1500 hours. Colombia, and northern Colombia lack a Illustrations of adults.•Male in tail (to which the name C. detesta refers), D'Abrera (1995), but identity of the female but do not otherwise differ. Specimens with that he illustrated as this species is unclear. and without a tail occur sympatrically and synchronically in El Salvador and southern Cyanophrys herodotus (Fabricius, 1793) Brazil, but again, do not otherwise differ. Distribution, habitat, and abundance.• Despite substantive genitalic variation, in- The most common, widespread, variable, cluding clinal differences, we find no con- and weedy species in Cyanophrys, occur- vincing evidence of sibling species. ring in almost all habitats under 1,500 m Larval food plants.•A polyphagous spe- throughout the Neotropics. cies that has been reared from plants in the VOLUME 107, NUMBER 2 411

Anacardiaceae, Boraginaceae, Compositae, ing along the outer margin of the ventral Dipsacaceae, Malvaceae, Sambucaceae, and hindwing. Verbenaceae (Lima 1928, 1930, 1936; San- Male behavior.•Males "hilltop" in the tos 1933; Monte 1934; Biezanko et al. morning before 0930 hours, which is 1974; Robbins and Aiello 1982). unique in the genus, so far as is known, but Male behavior.•Males set up mating ter- which may be shared by the preceding and ritories on hilltops and ridges in the middle succeeding species. of the afternoon from about 1300 to 1600 Illustrations of adults.•This species has hours at slightly greater heights above the been illustrated, so far as we are aware, ground, on average, than males of C. amyn- only in the original description. tor. Illustrations of adults.•Male in Cyanophrys crethona (Hewitson, 1874) D'Abrera (1995), but identity of the female Distribution, habitat, and abundance.•A that he illustrated as this species is unclear. Jamaican endemic. Identification.•The male genitalia of C. Cyanophrys miserabilis (Clench, 1946) crethona is similar to the preceding two Distribution, habitat, and abundance.• species, but is distinguished by its larger Occurs in seasonally dry habitats from sea size (i.e.. Table 2) and a very lightly scler- level to about 2,000 m elevation from otized ventral cornutus. southern United States (Texas) to the Pacif- Nomenclature.•Cyanophrys hartii ic side of Costa Rica. Turner and J. Y. Miller differs from C. cre- Identification.•It is easily distinguished thona by ventral brown coloration. Because from the sympatric C. herodotus by the exposure to humidity or physical abrasion brown frons (green in C. herodotus) and by can change scale color from green to the greater amount of dark maroon scaling brown, as mentioned in the discussion of along the outer margin of the ventral Character 1, this name was synonymized hindwing. Cyanophrys miserabilis appears with C. crethona (Robbins 2004b). to form a superspecies (monophyletic line- Illustrations of adults.•A male is illus- age in which no two species are sympatric) trated in D'Abrera (1995). with the next two species (Figs. 12, 13). Cyanophrys roraimiensis Johnson and Besides the presumed evolution of a brown Smith, 1993 frons in the ancestor of these species, they also possess "thin, stringy" brush organs. Distribution, habitat, and abundance.• Larval food plants.•Recorded larval Known only from the holotype female, food plants are Compositae (Eupatorium) which was collected on the Brazilian side and Leguminosae (Caesalpinia, Parkinson- of Mt. Roraima. We have seen a picture of ia). a female collected in the adjoining tepui re- Illustrations of adults.•Both sexes in gion of Venezuela that may be this species, D'Abrera (1995). but have not had the opportunity to examine it. Cyanophrys velezi Identification.•The ventral hindwing Johnson and Kruse 1997 pattern and size of this species is exceedingly similar to that of C. crethona Distribution, habitat, and abundance.• (Huntington 1933, Comstock and Occurs in Panama, where it has been re- Huntington 1943), with which it also shares corded only during the dry season, to west- a brown frons, for which reason we suspect ern Ecuador in seasonally dry habitats. that it is closely related to the preceding Identification.•Unlike C miserabilis, it three species. lacks a hindwing tail and dark maroon scal- Nomenclature.•Johnson and Smith VOLUME 107, NUMBER 2 413

It probably occurs in Bolivia, but we know son and Amarillo 1997) is inconsistent with of no records. the phylogenetic results (Fig. 12). Four of Illustrations of adults.•A female is il- these subgenera are not monophyletic and lustrated in D'Abrera (1995). one is monotypic. In some cases, such as the aptly named Plesiocyanophrys, they ap- Cyanophrys remus (Hewitson, 1868) pear to have been characterized with sym- Distribution, habitat, and abundance.•A plesiomorphies. In others, such as the char- very common species in the mountains of acterization of Cyanophrys (a brown frons), southern Brazil, Argentina (Misiones), Uru- they were delimited by a homoplastic char- guay, and Paraguay. In the subtropical parts acter of its range, it occurs at lower elevations. The C. longula and C herodotus groups Identification.•Males have brilliant dor- are monophyletic on the 15 most parsimo- sal blue color, similar to that in C. longula nious trees. The C longula group consists and C. pseudolongula, but the underside of species with a brown frons (Character 1, wing pattern has extensive brown markings, which is homoplastic), brush organs wider similar to the preceding two species. than 0.05 mm (Character 14, Fig. 10), brush Larval food plants.•It has been reared organs that are oriented horizontally at the from Calliandra (Leguminosae) in Uruguay anterior end (Character 11, Fig. 10), and and from Abutilón and Pavonia (Malva- brush organs that arise primarily above the ceae) in Brazil (Zikán 1956, Biezanko et al. origin of the vinculum strut (Character 13, 1966, Silva et al. 1968). Fig. 10). The latter three character states are Male behavior•Males "hilltop" in the unique in the Callophrys Section. These early afternoon, with records from 1145 to species occur in montane habitats (above 1500 hours. 800 m) except in the subtropical parts of Illustrations of adults.•Both sexes in southern South America. The C. herodotus D'Abrera (1995). group consists of species with two androconial clusters on the dorsal forewing DISCUSSION (Character 4, Figs. 4-5, both lost in C fu- Building on the work of Clench (1961), sius according to our results). All seven Robbins (2004a) distinguished the Callo- species occur in the lowlands, unlike the phrys Section of the Eumaeini by valve tips previous lineage, but most are also found in that are ñattened and without setae, but montane habitats. sometimes with a terminal thickening (il- Cyanophrys bertha is the sister of the lustrated in Warren and Robbins 1993). The seven-species C herodotus lineage in the form of the anal lobe (Character 7) and successively weighted cladogram (Fig. 13), presence of brush organs (Character 8) dis- but in the equally weighted original most tinguish Cyanophrys from other members parsimonious trees, it was sometimes the of the Callophrys Section. Although the ge- sister of the five-species C. longula lineage. nus could be split into smaller genera on The genitalia of C. bertha are nearly iden- the basis of the phylogenetic results (Figs. tical to those of C. amyntor, but C. bertha 12 and 13), the anal lobe character allows shares frons color, androconial structure, individuals to be recognized in the field. It and restriction to montane habitats with C. is unclear whether the remaining Holarctic longula (Table 1). members of Callophrys, or the many genera Cyanophrys bertha has been proposed into which they have been divided, are for "vulnerable" status (Brown 1993, monophyletic. Brown and Freitas 2000 and references The subgeneric nomenclature used by therein. Otero et al. 2000) and listed as "al- Johnson and colleagues (Johnson et al. most threatened" (Mielke and Casagrande 1993, Johnson and Le Crom 1997a, John- 2004). Its relatively basal position in the 414 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON cladogram increases its priority as a species Biezanko, C. M. De, Ruffinelli, A. and D. Link. 1974. of conservation concern (Atkinson 1989, Plantas y otras sustancias alimenticias de las oru- gas de los lepidópteros uruguayos. Revista do Vane-Wright et al. 1991). Centro de Ciencias rurais 4: 107-148. ACKNOWLEDGMENTS Brown Jr, K. S. 1993. Selected Neotropical species, pp. 146-149. In New, T. R., ed. Conservation Bi- We are grateful to Keith Brown for his ology of Lycaenidae (Butterflies) lUCN, Gland, kind hospitality, for generously sharing his Switzerland. knowledge of C. bertha, and for providing Brown Jr, K. S. and A. V. L. Freitas. 2000. Diversidade us with study specimens. We thank Mary de Lepidoptera em Santa Teresa, Espirito Santo. Boletim do Museu de Biología Mello Leitâo (N. Sangrey and the NMNH Research Training Sér.) 11/12: 71-118. Program for providing an internship to Carpenter, J. M. 1988. Choosing among multiple Duarte; the Women's Committee of the equally parsimonious cladograms. Cladistics 4: Smithsonian Institution and the National 291-296. Science Foundation (award #BIR-9531331) Clench, H. K. 1946. Notes on the amyntor group of the genus Thecla (Lepidoptera, Lycaenidae). En- for supporting this program; Olaf Mielke tomologist 79: 152-157, 185-191. and Mirna Casagrande for obtaining CNPq . 1961. Tribe Thechni, pp. 177-220. In Ehrlich, support for Robbins to curate the UFPR P. R. and A. H. Ehrlich, eds. How to know the collection in Curitiba; Karl Ebert for allow- butterflies. Brown Company, Dubuque, Iowa, 261 ing us to examine his father's collection; the pp. Comstock, W. P. and E. I. Huntington. 1943. Lycaen- curators and collections managers at the idae of the Antilles. Annals of the New York AMNH, BMNH, UFPR, INHS for allowing Academy of Sciences 45: 49-130. us to examine specimens under their care; D'Abrera, B. L. 1995. Butterflies of the Neotropical the many biologists who shared larval food region. Part VII. Lycaenidae. Hill House, Black plant information with us; Carl Hansen for Rock, i-xi + 1098-1270 pp. technical help in digitizing wing venation Draudt, M. 1919-1920. Theclini F, pp. 744-812. In Seitz, A. ed. Macrolepidoptera of the World, Vol. directly from slides; George Venable for V, The American Rhopalocera, Alfred Kernen making the figures; Astrid Caldas for pro- Verlag, Stuttgart, 1139 pp., 194 pi. viding a smooth interface between English Eliot, J. N. 1973. The higher classification of the Ly- and Portuguese; and especially Andrew caenidae (Lepidoptera): A tentative arrangement. Brower, John Brown, Keith Brown, Astrid Bulletin of the British Museum of Natural History (Entomology) 28(6): 371-505. Caldas, Larry Gall, Jason Hall, Donald Har- Farris, J. S. 1969. A successive approximations ap- vey, Gerardo Lamas, James Miller, Carla proach to character weighting. Systematic Zoolo- Penz, J. B. Sullivan, and Andy Warren for gy 18: 374-385. commenting, sometimes extensively, on the Hoffmann, E 1937. Beiträge zur Naturgeschichte bras- manuscript. The junior author has been fi- ilianischer Schmetterlinge. Entomologisches Jahr- nancially supported by the State of Sao buch (Frankfurt) 1937: 126-139. Holdridge, L. R. 1947. Determination of world land Paulo Research Foundation•FAPESP (02/ formations from simple climatic data. Science 13898-0 and 03/05895-4). 105: 367-368. Huntington, E. I. 1933. Erycinidae and Lycaenidae LITERATURE CITED (Lepidoptera, Rhopalocera) from the regions of Atkinson, I. 1989. Introduced animals and extinctions, Mt. Roraima and Mt. Duida. American Museum pp. 54-69. In Warren, D. and M. Pearl, eds. Con- No vitales 611: 1-5. servation for the twenty-first century. Oxford Uni- IBGE, Diretoria De Geociências. 1993. Mapa de ve- versity Press, New York, xv + 365 pp. getaçâo do Brasil. Rio de Janeiro, Brazil. Bailowitz, R. A. and J. E Brock. 1991. Butternies of Janzen, D. H. and W. Hallwachs. 2004. http://janzen. Southeastern Arizona. Sonoran Arthropod Stud- sas.upenn.edu ies, Inc., Tucson, ix + 342 pp. Johnson, K. and A. R. Amarillo S. 1997. Review of Biezanko, C. M. De, Ruffinelli, A. and C. S. Carbonell. Antephrys with description of a new Colombian 1966. Lepidoptera del Uruguay. III. Notas com- species. Revista de Theclinae Colombianos, Vol. plementarias. Boletín Facultad de Agronomía 1, 16 pp. (Montevideo) 91, 53 pp. Johnson, K., R. C. Eisele, and B. N. MacPherson. VOLUME 107, NUMBER 2 415

1993. Genera for the small green hairstreak but- Central and South America (Lycaenidae, Thecli- terflies of the Neotropical realm (Lepidoptera, Ly- nae). Journal of the Lepidopterists' Society 25, caenidae, Theclinae). Reports of the Museum of Supplement 1, 39 pp. Natural History, University of Wisconsin (Stevens . 1977. Studies in the genera of American hair- Point) 39: 1-16. streaks. 4. A new genus of hairstreak from Central Johnson, K. and J. F. Le Crem. 1997a. The "brown- and South America (Lycaenidae: Eumaeini). Bul- frons" complex of Cyanophrys sensu lato and letin of the AUyn Museum 44: 1-24. new sister species from Colombia and Brazil. Re- Nijhout, H. F. 1991. The development and evolution vista de Theclinae Colombianos, Vol. 1, 28 pp. of butterfly wing patterns. Smithsonian Institution . 1997b. Clench's Cyanophrys "amyntor Press, Washington, DC, 297 pp. Group" in Colombia with descriptions of several Nixon, K. C. 2002. WinClada version 1.00.08. Pub- outlying endemics from elsewhere in South Amer- lished by the author, Ithaca, New York. ica. Revista de Theclinae Colombianos, Vol. 1, 21 Otero, L. S., K. S. Brown, O. H. H. Mielke, R. E pp. Monteiro, J. M. Costa, M. V. Macêdo, N. C. Ma- Johnson, K. and D. S. Smith. 1993. A remarkable new ciel, J. Becker, N. G. Salgado, S. B. Santos; G. E. butterfly species from Jamaica (Lepidoptera: Ly- Moya, J. M Almeida, and M. Duarte. 2000. In- caenidae), with notes on Jamaican endemics and vertebrados terrestres, pp. 53-62. In Bergallo, H. their sister species. Reports of the Museum of G., C. E D. Rocha, M. A. S. Alves, and M. Van- Natural History, University of Wisconsin (Stevens sluys, eds. A fauna ameaçada de extinçâo do es- Point) 24: 1-13. tado do Rio de Janeiro. Editora da Universidade Kendall, R. O. 1975. Larval foodplants for seven spe- do Estado do Rio de Janeiro, Rio de Janeiro. cies of hairstreaks (Lycaenidae) from Mexico. Robbins, R. K. 1991. Evolution, comparative mor- Bulletin of the AUyn Museum 24: 1-4. phology, and identification of the eumaeine but- Kendall, R. O. and W. W McGuire. 1984. Some new terfly genus Rekoa Kaye (Lycaenidae: Theclinae). and rare records of Lepidoptera found in Texas. Smithsonian Contributions to Zoology, No. 498, Bulletin of the AUyn Museum 86: 1-50. 64 pp. Klots, A. B. 1970. Lepidoptera, pp. 115-130. In Tux- . 2004a.Introduction to the checklist of Euma- en, S. L., ed. Taxonomist's Glossary of Genitalia eini (Lycaenidae), pp. xxiv-xxx. In Lamas, G., ed. in Insects. Munksgaard, Copenhagen, 359 pp. Checklist: Part 4A. Hesperioidea•Papilionoidea. Lamas, G., R. G. Robbins and W. D. Field. 1995. Bib- In Heppner, J. B., ed. Atlas of Neotropical Lepi- liography of butterflies. An Annotated Bibliogra- doptera, Vol. 5A. Association for Tropical Lepi- phy of the Neotropical Butterflies and Skippers doptera, Scientific Publishers, Gainesville, xxxvi (Lepidoptera: Papilionoidea and Hesperioidea). In + 439 pp. Heppner, J. B., ed. Atlas of Neotropical Lepidop- . 2004b. Lycaenidae. Theclinae. Tribe Euma- tera. Gainesville, Scientific Publishers, Vol. 124, eini, pp. 118-137. In Lamas, G., ed. Checklist: xiv + 463 pp. Part 4A. Hesperioidea•Papilionoidea. In He- Lima, A. M. C. 1928. Segundo catálogo systematico ppner, J. B., ed. Atlas of Neotropical Lepidoptera, dos insectos que vivem ñas plantas do Brasil e Vol. 5A. Association for Tropical Lepidoptera, ensaio de bibliografia entomológica brasileira. Ar- Scientific Publishers, Gainesville, xxxvi + 439 pp. chivos da Escola superior de Agricultura e Med- Robbins, R. K. and A. Aiello. 1982. Foodplant and icina veterinaria. (Niterói) 8: 69-301. oviposition records for Panamanian Lycaenidae . 1930. Suplemento ao 2° catálogo systemático and Riodinidae. Journal of the Lepidopterists' So- dos insectos que vivem ñas plantas do Brasil e ciety 36: 65-75. ensaio da bibliografia entomológica brasileira. O Robbins, R. K. and G. Lamas. 2002. Nomenclatural Campo (Rio de Janeiro) 8: 84-91. Changes in the Neotropical Eumaeini (Lepidop- . 1936. Terceiro catálogo dos insetos que vivem tera: Lycaenidae: Theclinae). Revista brasileira de ñas plantas do Brasil. Rio de Janeiro, Ministerio Zoología 19: 197-214. da Agricultura, 460 pp. Robbins, R.K. and S.S. Nicolay. 1999. Taxonomy of Mielke, O. H. H and M. M. Casagrande. 2004. Bor- Strymon toussainti, S. andrewi, S. amonensis, and boletas, pp. 713-739, In S. B. Mikich and R. S. S. rhaptos (Lycaenidae: Theclinae). Journal of the Bérnils, eds., Livro vermelho da fauna ameaçada Lepidopterists' Society 52: 328•334. no Estado do Paraná. Instituto Ambiental do Pa- . 2002. An overview of Strymon Hübner (Ly- rana, Curitiba, 763 pp. caenidae: Theclinae: Eumaeini). Journal of the Monte, O. 1934. Borboletas que vivem em plantas cul- Lepidopterists' Society 55: 85-100. tivadas. Publicaçôes. Secretaria da Agricultura. Santos, E. 1933. Inimigos e Doenças das Fruteiras. Rio Estado de Minas-Gerais (Serie agrícola) 21, 221 de Janeiro, Biblioteca Agrícola d'O Campo. 81 pp. pp. Nicolay, S. S. 1971. A new genus of hairstreak from Scott, J. A. 1986. The butterflies of North America. A 416 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Natural History and Field Guide. Stanford Uni- Brazil that we have examined are listed be- versity Press, Stanford, xv + 583p. low (museum acronyms listed in Methods). Silva, A. G. d'A. e, C. R. Gonçalves, D. M. Galvao, A. J. L. Gonçalves, J. Gomes, M. do N. Silva, and If the elevation was not recorded on a spec- L. de Simoni. 1968. Quarto catálogo dos insetos imen's data label, we parenthetically note que vivem ñas plantas do Brasil seus parásitos e the elevation for that site as it is Hsted in predadores. Ediçâo ampliada do "3° catálogo dos gazetteers. So far as we are aware, there are insetos que vivem ñas plantas do Brasil" de au- no other known specimens of C. bertha. toría do Prof. A. M. da Costa Lima. Parte II. In- setos, hospedeiros e inimigos naturals. índice de 1. Minas Gerais, Barbacana, 900 m. 1Ç insetos e índice de plantas. Rio de Janeiro, Min- (Private collection of Karl Ebert, Santa isterio da Agricultura. 1: xxvii -I- 622 pp.; 2: [viii] Clara, Sâo Paulo, Brazil); 12,11 May + 265 pp. 1969 (USNM); lî, 27 May 1986 TOSí, J. Jr. 1983. Provisional life zone map of Brazil at 1:5,000,000 scale. Publication of the Institute (USNM). of Tropical Forestry, Rio Piedras, Puerto Rico, 16 2. Minas Gérais, Poços de Caldas, 1,300- pp. + map. 1,400 m. 1 (J and 1 9, March (Ebert). Vane-Wright, R. I., Humphries, C. J. and R H. Wil- 3. Sâo Paulo, Serra do Japi, 1,050-1,250 liams. 1991. What to protect?•Systematics and m. 1 cî, 24 March 1990 (USNM); 1 S, the agony of choice. Biological Conservation 55: 235-254. 25 March 1990 (USNM); \ S, lA May Warren, A. D. and R. K. Robbins. 1993. A natural 1990 (USNM); 1 cî, 2 June 1990 hybrid between Callophrys (Callophrys) sherida- (USNM). nii and C. (Incisalia) augustinus (Lycaenidae). 4. Paraná, Curitiba, (934 m). IS, Decem- Journal of the Lepidopterists' Society 47: 236- ber 1945 (UFPR). 240. Willis, J. C. 1973. A Dictionary of the Flowering 5. Paraná, Ponta Grossa, (969 m). 1$, Plants and Ferns. 8th edition revised by H. K. March 1948 (UFPR). Airy Shaw. University Press, Cambridge, 1,245 6. Paraná, Castro, (999 m). 1 î (BMNH) pp. 7. Santa Catarina, Corupá, 19 October Zikán, J. E 1956. Beitrag zur Biologie von 12 Thecli- 1975. 1 ? (UFPR). (Corupá is located at nen-arten. Dusenia 7: 139-148. 75 m elevation at the base of the mountains on the road leading to Sâo Bento APPENDIX do Sul, a well-known butterfly locality The data for the 13 museum specimens above 800 m elevation that is the likely of C. bertha from 7 localities in southern collection place for this specimen.)