: a Sora as Wh right Sy zetene’ ner ne ee) cards USpits y eof en teapaees Pf be eee fet bd i rat Re arts i Sih tp be fstin) sb aay » Shane : neds retrain ce 4\ ‘ ns ' | } i } t » a ; 7 i : : 4 "i Wives | R tL hi rf \ ¥ ps Niel 1 { es -: ‘ ; a t y f ae th x i i ' ae E { / i h chin = tes “ i \ 4 ¢ @ ; { , Ro ar i ni ie f F ( 1 c ( $ 4 { , 1 5 ¥ 7 ch ’ 2 ned fieet + a 4 é ‘ f a , t i } t 5 j bs N ; , } ‘ ( } i i i “fi y t | J ri j D { ui ad - 4 e aioe Ag : } Laer i i ‘ 1 d Uy ans ) ; yi ‘ d ; ft oe > f i 3 fy ‘ { ; | . ‘ ; \ ey } } , «. " ‘ j iN I ' de 5 , ~ Mi ; } 3 Oi eee: rf eae = [ Volume 54 , 2000 Number 1 ISSN 0024-0966 JOURNAL of the LEPIDOPTERISTS’ SOCIETY Published quarterly by THE LEPIDOPTERISTS’ SOCIETY Publié par LA SOCIETE DES LEPIDOPTERISTES : mf Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN : Publicado por LA SOCIEDAD DE LOS LEPIDOPTEROLOGOS - [ ¥ s ‘ a = ze ré . ; P 36 we r 2 2 > ¥ 4 Z . bi - 5a és . . Wis ¥ as a Y * ve s u . * tag ves . a sg s ic “ 4 * ‘ * ~ Es Ww - > " > * z ~ -~ * » 4 we} iy 5 s x a > < e , % ‘ s . S . 5 i - © = Al - « » * ; ny > + “ * y's f he a a & - ‘ 2 = * - 7 ‘ = GREAT SOUTHERN WHITE Ascia monuste > . . =x + “hy AA PAA ALA LA? > SAARI AA a ‘ at ; : SI a eiSL J - hes ' * “ os i * a) ig > RED RIM Biblik hyperia - te + + NASR AAR einen A Ar b 5 « Dd , - ad as ha ? a! é . % 7 + é nd rE ' ~ E = + . % 5 + < “ * a a . * ‘ b, Hp; * - ~ POLYDAMAS SWALLOWTAIL Bottus polydomas NAA AALS AAAS » CARIBBEAN BUCK! Junonia genvecea: AAA ARAN AAA mA 6 November 2000 THE LEPIDOPTERISTS’ SOCIETY EXECUTIVE CouNCIL Joun W. Brown, President Susan S. Borxin, Vice President Micuaet J. Smiru, Immediate Past President Mirna M. Casacranpe, Vice President Manue A. Batcazar-Lara, Vice President Davin C. Irtner, Treasurer Ernest H. WixuiAMs, Secretary Members at large: Ronald L. Rutowski M. Deane Bowers George L. Balogh Felix A. H. Sperling Ron Leuschner Andrew V. Z. Brower Andrew D. Warren Michael Toliver Brian Scholtens E\prroriAL BoarpD Rosert K. Rossins (Chairman), Joan W. Brown (Member at large) M. Deane Bowers (Journal) WitiraM EF. Mitter (Memoirs) Puiu J. Scuarrert (News) Honorary Lire MEMBERS OF THE SOCIETY Cuartes L. Remincton (1966), E. G. Munroe (1973), Ian F. B. Common (1987), Joun G. Franc.emont (1988), Lincotn P. Brower (1990), Douctas C. Fercuson (1990), Hon. Miriam Roruscuiip (1991), CLaupe Lemaire (1992), Frepericx H. Rinpce (1997) The object of The Lepidopterists’ Society, which was formed in May 1947 and formally constituted in December 1950, is “to promote the science of lepidopterology in all its branches, . . . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards these aims. Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Journal and the News of The Lep- idopterists’ Society. Prospective members should send to the Assistant Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. Active members—annual dues $45.00 within the U.S., $50.00 outside the U.S. Affiliated members—annual dues $10.00 within the U.S., $15.00 outside the U.S. Student members—annual dues $20.00 within the U-S., $25.00 outside the U.S. Sustaining members—annual dues $60.00 within the U.S., $65.00 outside the U.S. Life members—single sum $1,800.00 Institutional subscriptions—annual $60.00 Airmail postage for the News $15.00 Send remittances, payable to The Lepidopterists’ Society, to: Kelly M. Richers, Asst. Treasurer, 9417 Carvalho Court, Bakersfield, CA 93311; and ad- dress changes to; Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. For information about the Soci- ety, contact: Ernest H. Williams, Department of Biology, Hamilton College, Clinton, NY 13323. To order back issues of the Memoirs, write for ayail- ability and prices to Kelly M. Richers, Asst. Treasurer, 9417 Carvalho Court, Bakersfield, CA 93311. The additional cost for members outside the U.S. is to cover mailing costs. Journal of The Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by The Lepidopterists’ Society, o Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA and at additional mailing offices. POSTMASTER: Send address changes to The Lepidopterists’ Society, “o Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Cover illustration: Butterfly stamps from the Commonwealth of Dominica; Great Southern White, Ascia monuste (Pieridae); Polydamas Swallowtail, _ Battus polydamas (Papilionidae); Red Rim, Biblis hyperia (Nymphalidae); The Flambeau, Dryas julia (Nymphalidae); Cassius Blue, Leptotes cassius (Ly- — caenidae); Monarch, Danaus plexippus (Nymphalidae); Godman’s Leaf, Anaea dominicana (Nymphalidae); Caribbean Buckeye, Junonia genoveva (Nymphalidae). J - SSE Coe ee ae ees Poet es or i JOURNAL OF Tue LeErPrIporptTeERiIstTs’ SOCIETY Volume 54 Journal of the Lepidopterists’ Society 54(1), 2000, 1-28 2000 Number 1 HESPERIIDAE OF RONDONIA, BRAZIL: “ANTIGONUS” GENUS GROUP (PYRGINAE), WITH TAXONOMIC COMMENTS AND DESCRIPTIONS OF NEW SPECIES FROM BRAZIL AND GUATEMALA GEORGE T. AUSTIN Nevada State Museum and Historical Society, 700 Twin Lakes Drive, Las Vegas, Nevada 89107, USA ABSTRACT. Some species of the pyrgine (Hesperiidae) “Antigonus” group (sensu Evans 1953) occurring in central Rondo6nia, Brazil, are discussed. These are six species of Mylon Godman & Salvin, 1894, including two new species Mylon simplex and Mylon argonautarum; four species of Carrhenes Godman & Salvin, 1895, including the new species Carrhenes recurva; four species of Clito Evans, 1953, including the new combination Clito clada, new status; one species of Xenophanes Godman & Salvin, 1895; three species of Antigonus Hiibner [1819]; and one species each of Timochreon Godman & Salvin, 1896, and Anisochoria Mabille [1877]. Taxonomic comments and illustrations (including male and female genitalia) are provided for these and for some related taxa from elsewhere. The identities of Leucochitonea jason Ehrmann, 1907, and Mylon ozema var. exstincta Mabille & Boullet, 1917, are clarified. A lectotype is designated for L. jason. The holotype of M. o. var. exstincta is identified and M. exstincta, new status, is raised from its current synonymy with M. jason. Mylon cristata, new species, is described from Guatemala. Carrhenes chaeremon (Mabille, 1891), revised status; Carrhenes leada (Butler, 1870), revised status; and Carrhenes lilloi Hay- ward, 1947, revised status, are raised to specific level taxa from their current subspecific or synonymic placements. Additional key words: genitalia, Neotropics, phenology, variation. Evans (1952, 1953) divided the Pyrginae (Hes- periidae) into several groups of genera. Although these groups may not be monophyletic and may contain mis- placed species (e.g., de Jong 1975, Burns 1990), they are convenient for discussions of species richness and taxonomy of local faunas. Near the municipality of Ca- caulandia in central Rond6nia, Brazil, an area of ongo- ing studies of the butterfly fauna (Emmel & Austin 1990), seven of the eleven genera of the “Antigonus” group have been found: Mylon Godman & Salvin, 1894; Carrhenes Godman & Salvin, 1895; Clito Evans, 1953; Xenophanes Godman & Salvin, 1895; Antigonus Hiibner, [1819]; Timochreon Godman & Salvin, 1896; and Anisochoria Mabille [1877]. In this paper, sixteen species previously described in these genera are dis- cussed, four new species are described, a new combi- nation is established, and a species considered by Evans (1953) to be a synonym is identified. STuDY AREA AND METHODS A detailed description of the study area near Cacaulandia is presented by Emmel and Austin (1990) and Emmel et al. (2000). In the Cacaulandia area, there is a pronounced seasonality of precipita- tion. A dry season extends from May through Septem- ber with practically no rainfall in June, July, and August. The wettest months are usually January and February. Forewing length was measured from the base to the apex. “GTA” numbers associated with specimens refer to genital vial numbers. Structures of the genitalia are those used by Austin and Mielke (1998). MyYLon GODMAN & SALVIN, 1894 Evans (1953) included eleven species and four sub- species in this Neotropical genus which ranges from Mexico to Argentina. Six species, including one con- sidered as a synonym by Evans (1953) and two unde- scribed, were recorded in central Rondénia. These are present in small numbers throughout the year, but are most prevalent during the early wet season (Figs. 107, 108). I propose that the species of Mylon be divided into three species groups based on wing pattern and the morphology of male and female genitalia. I also il- lustrate the genitalia of taxa from other areas, and de- scribe a new species from Guatemala. ‘lassia” Group Three species groups appear within Mylon, the first proposed being the “lassia” group including, in addi- tion to the species discussed below, Mylon zephus (Butler, 1870) and Mylon salvia Evans, 1953. This group is characterized by hyaline subapical macules on the forewing, a tibial tuft on the hindleg entering a thoracic pouch , a broadly triangular gnathos in ventral view, no style from the ampulla, one or two prominent spines on the aedeagus, and a membranous sac on the ventral side of the phallobase. Mylon lassia (Hewitson, 1868) (Figs. 1, 2, 55, 87) M. lassia is known from Mexico to northern South America (Evans 1953). Males from Costa Rica have genitalia (Fig. 55) as illustrated by Godman and Salvin (1879-1901) and Evans (1953). The female genitalia (Fig. 87) have a narrow lamella postvaginalis which is shallowly notched centrally on its caudal edge and a lamella antevaginalis with broad lateral plates. Mylon illineatus illineatus Mabille & Boullet, 1917 (Figs. 3, 56) The nominotypical subspecies of M. illineatus ranges from Ecuador to Peru. The genitalia of a male from Ecuador is illustrated here (Fig. 56) in more de- tail than by either Hayward (1947, 1948) or Evans (1953). Mylon orsa Evans, 1953 (Figs. 4, 57) M. orsa was known from the male and two females of Evans’ (1953) original description of this species from Costa Rica. The genitalia of an additional male from Costa Rica is illustrated here (Fig. 57) in more detail than previously. Mylon mestor Evans, 1953 (Figs. 5, 6, 58, 88) M. mestor was known only from the unique type from Ecuador (Evans 1953). An additional male and a female from Ecuador are illustrated here. The male genitalia are also shown (Fig. 58) and in more detail than by Evans (1953). The female genitalia (Fig. 88) JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY are similar to those of M. lassia, but have a narrower sterigma; the tubular ductus bursae leads to an oblong corpus bursae. Mylon ander ander Evans, 1953 (Figs. 7, 59) M. ander is known from Colombia to Bolivia and southern Brazil (Evans 1953). It is rare in the Ca- caulandia area with one record in October and three in November (Fig. 108). The male genitalia (Fig. 59) ap- pear as illustrated by Evans (1953). “menippus’ Group The two species of the here proposed “menippus” group, discussed below, differ from the “lassia” group by the lack of hyaline subapical macules on the fore- wing, no tibial tuft, a narrower gnathos, a short and blunt style caudad from the ampulla, and no spines and a generally smaller sack-like structure on the aedeagus. They do, however, have a dorsal process from the harpe as does the “lassia” group, similarly long arms of the un- cus (very long on M. cajus), and a very broad vinculum. Mylon menippus (Fabricius, 1777) (Figs. 9, 10, 60, 89) M. menippus is the most common Mylon in central Rond6nia with records for every month and a peak flight in the early wet season (Fig. 107). The male gen- italia (Fig. 60) are as illustrated by Godman and Salvin (1879-1901) and Hayward (1933, 1947, 1948) as My- lon melander (Cramer [1780]) and by Evans (1953). There appears to be no variation in the genitalia over the species’ broad distribution (Mexico to Argentina), but there is some individual variation in the shape of the harpe. The female genitalia (Fig. 89) have a rela- tively broad lamella postvaginalis (broader than on M. lassia and M. mestor), a lamella antevaginalis with broad lateral lobes, a long and thin ductus bursae, and an oblong corpus bursae. There is some individual (but not seasonal) variation in the extent and intensity of markings on the dorsal wings. Some individuals from Rondé6nia lack the pale-centered dark bar in the mid- discal cell of the forewing, the key character used by Evans (1953) to distinguish M. menippus from the fol- lowing species. => Fics. 1-8. Mylon (dorsal surface on left, ventral surface on right). 1. M. lassia male, COSTA RICA: San José Prov.; Finca E] Rodeo, 25 Mar. 1989; 2. M. lassia female, COSTA RICA: Alajuela Prov.; 6.8 km W of Atenas, 22 Dec. 1984; 3. M. illineatus male, ECUADOR: Pastaza Prov.; 25 km NE Puyo, 28 June 1980; 4. M. orsa male, COSTA RICA: Alajuela Prov.; 2.8 km S of Cinchona, 27 Sept. 1987; 5. M. mestor male, ECUADOR: Pichincha Proy.; Hotel Tinalandia, 2 July 1980; 6. M. mestor female, ECUADOR: Pichincha Prov.; 47 km E of Santa Domingo de los Colorados, 12 May 1988; 7. M. ander male, BRAZIL: Rondé6nia; 5 km S of Cacaulandia, 11 Nov. 1995; 8. M. cajus hera male, COSTA RICA: San José Prov.; cerro west of Patarra, 11 Oct. 1987. VOLUME 54, NUMBER 1 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY dorsal surface on left, ventral surface on right). 9. M. menippus male, BRAZIL: Rondénia; 5 km S of Cacaulandia, 11 nia; Fazenda Rancho Grande, 20 Oct. 1989; 11. M. pelopidas male, GUATEMALA: Petén; female, MEXICO: Pueblo; nr. Izucar de Matamoros, 11 Aug. 1962; 13. M. jason male lec- dénia: Linha C-20, off B-65 at Rio Canaa, 15 Nov. 1994; 16. M. Fics. 9-16. Mylon ( Nov. 1995; 10. M. menippus female, BRAZIL: Rond6é Parque Nacional Tikal, 4 Feb. 1992; 12. M. pelopidas totype; 14. M. jason female paralectotype; 15. M. exstincta male, BRAZIL: Ron exstincta female holotype. VOLUME 54, NUMBER 1 Fics. 17-21. Mylon (dorsal surface on left, ventral surface on right). 17. M. argonautarum male holotype; 18. M. argonautarum female, BRAZIL: Rondénia; 5 km S of Cacaulandia, 18 June 1994; 19. M. cristata male holotype; 20. M. cristata female, GUATEMALA: Petén; Par- que Nacional Tikal, 31 May 1993; 21. M. simplex male holotype. Mylon cajus hera Evans, 1953 (Figs. 8, 61) This subspecies of Mylon cajus (Pl6étz, 1884) was de- scribed from Panama and is known also from Guatemala and Costa Rica (Evans 1953). A male from Costa Rica is illustrated along with its genitalia (Fig. 61) in more detail than by Evans (1953). “pelopidas” Group The “pelopidas” species group, herein designated within Mylon, is characterized by the vinculum of the male genitalia extending conspicuously dorsad to en- velop most of the tegumen, being supported by a pair of flaps recurving outward from near the caudal end of the tegumen. The group is also distinguished from other Mylon by the short arms of the uncus, a compar- atively narrow vinculum, an elongate style from the ampulla, the lack of a dorsal process on the harpe (ex- cept for M. pelopidas), and a dextral hook near the caudal end of the aedeagus in dorsal view. The gnathos of the “pelopidas” group is narrow, there are no spines or a sac-like structure on the aedeagus, and, as on the “menippus’ group, there are no hyaline subapical mac- JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 22-29. Carrhenes; all from BRAZIL: Rondénia (dorsal surface on left, ventral surface on right). 22. C. chaeremon male, 3 km E of Fazenda Rancho Grande, 18 Nov. 1992; 23. C. chaeremon female, 5 km S of Cacaulandia, 10 May 1995; 24. C. bamba male, Fazenda Rancho Grande, 12 Noy. 1994; 25. C. bamba female, 5 km S of Cacaulandia, 7 Apr. 1995; 26. C. leada male, Fazenda Rancho Grande, 7 Nov. 1991; 27. C. leada female, Fazenda Rancho Grande, 13 June 1993; 28. C. recurva male holotype; 29. C. recurva female, B-65, 1 km N of Cacaulandia, 5 Nov. 1990. VOLUME 54, NUMBER | ules on the forewing or tibial tuft. The sterigma of the female genitalia is oval in shape and the lamella ante- vaginalis has a central process extending cephalad which divides a translucent area (“windows”) defined caudad and laterad by its lateral lobes. The shape and size of these “windows” and the shape of and the pat- tern of spiculation on the central process of the lamella antevaginalis are very useful for determining species. The anterior one-half to two-thirds of the sterigma is covered ventrally by a largely transparent membrane. The ductus bursae is long and very slender and leads to a globular corpus bursae. The group includes M. pelopidas, M. jason, M. exstincta (raised from syn- onymy below), and three new species, all of which are discussed in the following. M. pelopidas, M. exstincta, and two of the new species were encountered near Ca- caulandia. Mylon pelopidas (Fabricius, 1793) (Figs. 11, 12, 62, 90) M. pelopidas is a familar and widespread species (male forewing length = 19.9 mm [19.0-20.3, N = 5]; female forewing length = 21.4 mm [N = 1]; samples from Rond6nia) occurring from Mexico south to Paraguay and southern Brazil (Evans 1953). Although there is a certain amount of individual variation in the intensity of dorsal markings (some of this mediated by wear), there appears to be no geographical variation and the male genitalia (Fig. 62) are constant as more or less illustrated by Godman and Salvin (1895), Hol- land (1927), and Hayward (1933), all as Mylon ozema (Butler, 1870), and by Evans (1953). The female geni- talia (Fig. 90) have a lamella postvaginalis with a straight caudal end divided by a V-shaped central notch. The lamella antevaginalis is represented by broad lateral lobes extending nearly as far caudad as the caudal edge of the lamella postvaginalis and has a broad central process flared at its cephalad edge and densely spiculose on its lateral edges. The “windows” are broad and rectangular. M. pelopidas is uncommon in central Rondénia and is represented by records for January through May, July, and October through December (Fig. 108). Mylon jason (Ehrmann, 1907) (Figs. 13, 14, 63, 91) The concept of M. jason has been plagued with problems. The species was described by Ehrmann (1907) as Leucochitonea jason from specimens taken in Venezuela. Lindsey (1925) commented on and illus- trated the genitalia of a phenotype from South Amer- ica which resembled Mylon ozema (Butler, 1870), a name now synonymized with Hesperia pelopidas ~l Fabricius, 1793, but was apparently unaware of Ehrmann’s (1907) name. Holland (1927), in corre- spondence with Lindsey, resolved that two superfi- cially similar taxa with very different genitalia existed and that M. jason applied to the unplaced phenotype of Lindsey (1925). Holland (1927) went on to repro- duce Lindsey’s (1925) illustration of the genitalia of M. jason, as well as the Godman and Salvin (1895) figure of M. ozema, but took “the liberty of adding the termi- nal tuft of bristles, which we have found to be highly characteristic and to occur in every one of the numer- ous specimens which we have microscopically exam- ined .. .” to Lindsey’s figure. The “bristles” refer to those at the caudal end of the harpe. Liberty, however, is not always a good thing. The “type” of L. jason is a male and, although the genitalia had not been dis- sected, the harpes were observable beyond the ab- dominal integument and were clearly without bristles. Apparently, these were not examined by Holland (1927) although Lindsey (1925) may have seen this species as his figure showed no terminal bristles on the harpe. Lindsey, subsequently, also saw specimens with these bristles as noted in correspondence to Holland (Holland 1927), the latter apparently misled Lindsey on the concept of M. jason, and thus a confusion con- cerning the identity of M. jason was initiated. Neither Lindsey (1925) nor Holland (1927) seemed aware that in the years between the description of M. jason and their studies, Mabille and Boullet (1917) de- scribed another taxon, Mylon ozema var. exstincta which also had direct bearing on the problem in the identification of M. jason. That taxon was described from a single female from “Amazone sup.” in the Boul- let Collection at the Paris Museum. To complete this history, Hayward (1947, 1948) illustrated genitalia of Mylon “jason” showing bristles on the harpe and Evans (1953) synonymized M. exstincta with M. jason and illustrated genitalia without bristles. As if the complications in the identification and tax- onomy of M. jason perpetuated through time were not enough, three species of Mylon with this general su- perficial phenotype were found among material from central Rond6énia, two without bristles on the harpe and one with. It thus became critical to examine the types of the two applicable taxa for which names are available to establish their identity. As noted, the “type” of M. jason has no bristles on the caudal end of the harpe. Dissection of its genitalia revealed that the terminal end of the harpe is very robust, even in dorsal view, and completely unlike any of the three pheno- types from Rondénia. The two female “paratypes” of M. jason which exist (Holland 1927) were also dis- sected and proved to be of two species as thought by 8 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 36 Fics. 30-37. Carrhenes, Xenophanes, and Antigonus (dorsal surface on left, ventral surface on right). 30. C. lilloi male, ECUADOR: Rio Napo, Limoncocha, 10 July 1983; 31. C. canescens male (gray phenotype), GUATEMALA: Petén; El Remate, Cerro Cahul, 28 Sept. 1994; 32. C. canescens male (brown phenotype), MEXICO: San Luis Potosi; Cd. Valles, 12 July 1972; 33. C. canescens female (brown phenotype), MEX- ICO: San Luis Potosi; Cd. Valles, 12 July 1972; 34. X. tryxus male, BRAZIL: Rondénia; 5 km S of Cacaulandia, 8 July 1996; 35. X. tryxus fe- male, BRAZIL: Rondonia; 5 km S of Cacaulandia, 14 July 1995; 36. A. liborius male, BRAZIL: Rondonia; 5 km S of Cacaulandia, 15 July 1995; 37. A. liborius female, BRAZIL: Rondénia; 5 km S of Cacaulandia, 28 May 1994. VOLUME 54, NUMBER | Holland (1927). That female which he associated with M. jason (his Plate 1, fig. 2) matches well in the details of the superficial characters of the “type” male. The second female is a M. pelopidas. The “type” of M. o. var. exstincta likewise had not been dissected. Its gen- italia are different from those of both the “paratype” considered above as the female of M. jason and those of the associated female of M. pelopidas. Similarly, de- tails of the superficial markings of the M. o. var. exstincta “type” differed from those exhibited by each of those females. These markings, however, were nearly identical to those of males of one of the Rond6- nia phenotypes, this also without caudal bristles on the harpe (see below). To firmly establish the identity of Leucochitonea ja- son Ehrmann, 1907, the male “true type” (Holland 1927) is here firmly established as the lectotype, the female with similar superficial markings alluded to above is designated the paralectotype, and the wings and genitalia of both are illustrated herein. The lecto- type (forewing length = 21.7 mm) has labels as follows: red, handprinted - TYPE; white, printed - Ehrman [sic] Coll. / Carn. Mus. / Acc. 7815; white, handprinted - Ann. Carn. Mus. / vol. [not given] 1927. / PI. xxviii, fig. 1. 6; white, handwritten - L. jason Ehrmann / Type No. 555 / E. A. Klages Coll. / Suapure 10/28. 1899 / Venezuela; white, printed and handprinted - Genitalia Vial / GTA - 7307, with the following label added: red, printed - LECTOTYPE / Leucochitonea jason / Ehrmann, 1907 / designated by / G. T. Austin 1997. The paralectotype female (forewing length = 21.8 mm) has labels as follows: red, printed - Paratype; white, printed - Ehrman [sic] Coll. / Carn. Mus. / Acc. 7815; white, handprinted - Ann. Carn. Mus. / vol. [not given] 1927. / PI. xxviii, fig. 2. 9; white, handwritten - L. jason Ehr. / Type No. 555 / Edw. A. Klages C’ol. / Stia- pure 1/18/1900 / Venezuela; white, printed and hand- printed - Genitalia Vial / GTA - 7309, with the follow- ing label added: red, printed - PARALECTOTYPE / Leucochitonea jason / Ehrmann, 1907 / designated by /G. T. Austin 1997. As noted above, the second female “paratype” is a M. pelopidas with the following labels: red, handprinted - ParaType; white, handwritten - L. jason Ehr. / Type No. 555 / E. A. Klages Coll. / Stia- pure 1/9. 1900 / Venezuela; white, printed - Ehrman - [sic] Coll. / Carn. Mus. / Acc. 7815; white, handwritten - E. ozema Butl. / 2. Fixed by Holland, cf. / Ann. C. M., Vol XXII / Art. [not given], p. [not given].; white, handprinted - Ann. Carn. Mus. / vol. [not given] 1927. / Pl. XXVIII, fig. 7.2 white, printed and handprinted - Genitalia Vial / GTA - 7308. Superficially, M. jason is similar to M. pelopidas, but is larger. The mottling of the wings is as on M. pelopidas, but the ground color has more sheen, the postmedian line of the hindwing is less prominent than usual on M. pelopidas, and there is less gray- brown scaling between this and the margin. The end of the forewing discal cell, represented by darkened cross veins, is more or less parallel to the termen on M. jason and thus is directed towards the proximal edge of the postmedian macule in CuA,—2A whereas the end of the discal cell on M. pelopidas is more erect and directed towards the distal edge of this macule; this character was the one superficial char- acter used by Evans (1953) to distinguish the two species. The genitalia illustrated by Lindsey (1925) are probably of M. jason, those illustrated by Holland (1927) represent a mixed drawing of M. jason (after the figure by Lindsey 1925) with bristles of an unde- scribed species added, those represented by Evans (1953) may be of M. jason, but appear to be of M. exstincta or of an undescribed species. The genitalia represented as M. jason by Hayward (1948) have cau- dal bristles on the harpe and are obviously not of that species. The female genitalia of M. jason have a curved caudal edge to the lamella postvaginalis di- vided by a relatively broad V-shaped central notch and a lamella antevaginalis with wide and lobate lat- eral lobes and a moderately wide central process with a slightly broadened and convex cephalad end and lat- eral spicules. The “windows” are narrow and nearly round. Three additional females of M. jason were examined (forewing length = 20.6 mm [20.2-21.3]), all from Guatemala: Petén; Parque Nacional Tikal, 3-4 Feb. 1992 (GTA #7318, 7332, 7337). Thus its distribution extends at least from Guatemala to Venezuela. Records of M. jason in the literature must be treated as suspect until the specimens upon which they were based are critically reexamined. Mylon exstincta Mabille & Boullet, 1917, new status (Figs. 15, 16, 64, 92) Mylon ozema (Butler, 1870) var. exstincta Mabille & Boullet, 1917. Mylon jason Ehrmann, 1907: Evans, 1953. The identity and “type” of M. exstincta were dis- cussed above. This female type (called a male by Evans 1953, forewing length = 21.8 mm) with the fol- lowing labels: red, printed - TYPE; white, printed and handprinted - Amazone / Supérieur / 1905 / O. Staudinger / COLL BOULLET (in red letters on left side) / MUSEUM PARIS (in red letters on right side); green, handwritten - M. ozema, / var. Extincta [sic] / Mab. & Boull.; white, handwritten - Mylon / ozema var. / Exstincta Mab. Boull. / Bull. Soc. ent. France, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 10 VOLUME 54, NUMBER 1 1917, p. 55; white, printed and handprinted - Genitalia Vial / GTA - 5983, is here identified as the holotype of Mylon ozema var. exstincta Mabille & Boullet, 1917 and the following label has been added: red, printed - HOLOTYPE / Mylon ozema / var. exstincta / Mabille & Boullet, 1917 / identified by / G. T. Austin 1997. It and its genitalia are illustrated herein (Figs. 16, 92). A male associated with the holotype is a Mylon pelopidas and has the following labels: red, printed - TYPE; white, printed and handprinted - Amazone / Supérieur / 1905 / O. Staudinger / COLL BOULLET (in red let- ters on left side) / MUSEUM PARIS (in red letters on right side); Genitalia Vial / GTA - 5984; white, printed and handprinted - Mylon pelopidas / Fabricius (1793) / det. G. T. Austin 1997. M. exstincta resembles both M. pelopidas and M. ja- son superficially in overall wing shape and color. The species is larger than M. pelopidas and about the size of M. jason. M. exstincta is like M. jason in the orienta- tion of the distal end of the forewing discal cell. Al- though the dark forewing markings are similar on all three species, those of M. exstincta are much less con- trasting giving a noticably less mottled aspect. The hindwing of M. exstincta is largely unmarked with only a gray shade posteriorly and a darker gray submarginal line being apparent. The postmedian line is not trace- able anterior to vein M,. The genitalia of the holotype of M. exstincta are generally similar to those of M. jason, but differ in de- tail. The lamella postvaginalis is quadrate with a nar- row U-shaped notch on the caudal edge (angled with a broad V-shaped central indentation on M. jason) and the lamella antevaginalis is represented by a pair of rounded lateral lobes (narrower on M. jason), a cepha- lad directed central process which is relatively broadly expanded at and spiculose across its cephalad end (less expanded and with lateral spicules on M. jason). The “windows” are more elongate and ovate than they are on M. jason. The male from Rondonia, Brazil (Linha C-20, off B-65 at Rio Pardo, 15 Nov. 1990, GTA #830), considered to be M. exstincta (forewing length = 21.7 mm), also has genitalia generally similar to those of M. jason. The entire genital capsule of M. exstincta is less robust than is that of M. jason. The structure of the harpe is the key to identity. This is very robust, broad in dorsal view, heavily adorned with thorn-like spines, — 1] and recurved to about the level of the style from the ampulla on M. jason and much less robust, relatively thin in dorsal view, with fewer and smaller spines, and recurved but slightly on M. exstincta. Because M. o. var. exstincta differs from M. jason in superficial characters and in the genitalia of both sexes, it is here raised from Evans’ (1953) synonymy with M. jason to specific status. Mylon simplex Austin, new species (Figs. 21, 65) Description. Male - forewing length = 20.6 mm (19.7-21.8, N = 6); ground color of both dorsal wings shining off-white, forewing overscaled with gray and marked with darker gray macules giving mottled ap- pearance, these macules as submarginal, postmedian, postbasal, and basal series in addition to one in discal cell 2/3 distance from wing base to end of cell, sub- marginal and postmedian macules darkest (nearly black) towards costa; submarginal series sets off rela- tively distinct pale gray margin; pale ochre-brown be- tween postbasal and basal macules; veins black, those enclosing end of discal cell directed towards proximal edge of postmedian macule in CuA,—2A. Hindwing with extensive gray along anal margin, this broadest at base of wing, extending to vein M, or M, as post- median line; submarginal band of gray-brown decreas- ing in size and becoming macular (generally chevron- shaped) anteriorly; veins entirely black posteriorly, distally anteriorly. Venter with dorsal pattern very vague to obsolete except at forewing apex where most distinct. Head black on dorsum with white central patch, white beneath antennae and around eyes, antennae black with white at segments on venter and beneath club, nudum red-brown with 17 (N = 1) or 18 (N = 4) segments, palpi mixed white and dark gray; thorax brown with scattered white scales on dorsum, gray on venter, legs brown with white scales, mid-tibiae with single pair of spurs, hind tibiae with two pairs, no tib- ial tuft; abdomen gray on dorsum with white at seg- ments, white on venter. Genitalia - tegumen short, stout, with moderately broad recurved flaps caudad supporting anterior part of broad and enclosing vinculum; vinculum relatively straight; saccus broad, slightly upturned; uncus slightly Fics. 38-45. Antigonus; all from BRAZIL: Rondénia (dorsal surface on left, ventral surface on right). 38. A. nearchus male, 5 km S of Ca- caulandia, 27 Apr. 1995; 39. A. nearchus male, 5 km S$ of Cacaulandia, 4 July 1993; 40. A. nearchus male, 5 km S of Cacaulandia, 8 Feb. 1994: 41. A. nearchus female, B-80, between linhas C-10 and G-15, 19 Nov. 1991; 42. A. erosus male, Linha C-20 at Rio Pardo, 10 Dec. 1990; 43. A. erosus male, 5 km S of Cacaulandia, 18 Aug. 1993; 44. A. erosus female, 5 km S of Cacaulandia, 22 July 1995; 45. A. erosus female, 5 km S of Cacaulandia, 2 Oct. 1993. 12 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY Fics. 46-54. Clito, Anisochoria; and Timochreon, all from BRAZIL: Rondé6nia unless noted (dorsal surface on left, ventral surface on right). 46. C. clito male, Fazenda Rancho Grande, 16 June 1993; 47. C. clito female, 5 km S of Cacaulandia, 30 July 1997; 48. C. “aberrans” male, COSTA RICA: Guanacaste Proy.; La Pacifica, nr. Cafias, 17 Dec. 1984; 49. C. littera male, Fazenda Rancho Grande, 31 Oct. 1993; 50. C. zelotes male, Fazenda Rancho Grande, 20 Nov. 1991; 51. C. clada Fazenda Rancho Grande, 10 Oct. 1993; 52. A. pedaliodina pedaliodina male, Fazenda Rancho Grande, 20 Oct. 1989; 53. A. pedaliodina pedaliodina female, Fazenda Rancho Grande, 3 Nov. 1989; 54. T. satyrus male, Fazenda Rancho Grande, 3 Nov. 1989. => Fics. 55-62. Male genitalia of Mylon (from BRAZIL: Rondé6nia unless noted otherwise). Structures shown are lateral, dorsal, and ventral views of tegumen, uncus, gnathos, and associated structures; lateral internal view of right valva (second view of partial valva shows the same flat- tened); lateral and (usually) dorsal views of aedeagus and associated structures; and dorsal view of transtilla and juxta. 55. M. lassia, COSTA RICA (GTA #7354): 56. M. illineatus, ECUADOR (GTA #7355); 57. M. orsa, COSTA RICA (GTA #7359); 58. M. mestor, ECUADOR (GTA #7358); 59. M. ander (GTA #6394); 60. M. menippus (GTA #7349); 61. M. cajus hera, COSTA RICA (GTA #985); 62. M. pelopidas (GTA #2123). WES oO =
CVERREE
—
i
oOo
g
w
—
BR
>
wo
(7)
)
~)
q i) .
ow co iS
REET ERER
ee
i~ =)
oOo
=
>
BR
&
w
>
a
>
—
wee
co
o
=
BR
w
an an
~—
>
on
or
a
co
an
oOo
a
o
=
6
2
oe)
(7)
>
(=r)
>)
co
[=r]
oOo
=
[—)
oe
ad
ae
ease
rahe
(VEE
eared’
‘
th
1 72
3 14 16 7]
8 79 80 81 82
Fics. 23-141. Genitalia of 119 ¢ Pyrgus—in order of capture—from a single field at Portal, Chiricahua Mountains, 4800 ft [1465 m]J,
Cochise County, Arizona, USA, summer of 1974, J. M. & S. N. Burns (USNM): distal end of left valva in lateral view. 23-27, 11 July; 28, 29, 13
july; 30-43, 17 July; 4-46, 18 July, 47-49, 20 July; 50-52, 21 July; 53-67, 23 July; 68-87, 27 July; 88-111, 30 July; 112-141, 5 August. 23,
26, 30, 50, 68, 69, 75, 76, 83, 86-90, 93, 95-102, 105, 106, 108, 112-141, P. communis (N = 56). 24, 25, 27-29, 31-49, 51-67, 70-74,
77-82, 84, 85, 91, 92, 94, 103, 104, 107, 109-111, P. albescens (N = 63).
61
VOLUME 54, NUMBER 2
62
date from 22 May 1976 (Pensacola Beach, 1 4
[USNM]) and 7 September 1984 (Cantonment, 12 4, Je
M. Burns [USNM]). Those from farther east in
Florida as well as directly to the north in Alabama and
Georgia (34 d, J. V. Calhoun) date from 1989-95. An
eastward progression of albescens suggested by these
dates could be an artifact because the dates largely re-
flect times when I begged sympathetic, suitably situ-
ated, local collectors to sample Pyrgus. (Most collec-
tors find these ubiquitous skippers trashy and do not
stoop to collecting them, especially not in series). If
the presence of albescens along the Gulf Coast is none
too recent, then consider this: Grote (1872:69) de-
scribed P. communis from “central Alabama’—pre-
sumably the vicinity of Demopolis, which was his
home—so he may have come within a hundred miles
(160 km) of catching its look-alike, P. albescens, instead
(or besides).
However, it is clear that P. albescens is currently in-
creasing in numbers and spreading eastward and
southward in Florida at a rapid rate. In September and
October 1999, J. V. Calhoun caught a total of 70 male
albescens not only in most counties of the panhandle
but also at the top of the peninsula—in Alachua,
Gilchrist, and Levy counties—and nearly halfway
down the peninsula in Pasco County on the Gulf side
and in Volusia and Brevard counties on the Atlantic
side (Fig. 21). At the same time, he caught no P. com-
munis whatsoever. In Pasco County, he took 12 male
albescens in the very area (southeast of Dade City) in
which, 9 and 10 years earlier, at similar times of year,
he got only communis. In examining the Pyrgus mate-
rial of the Florida State Collection of Arthropods, I
found 33 male P. communis—taken between 1942 and
1977 in Alachua, Clay, Duval, and Liberty counties—
but no Florida examples of P. albescens. At least at this
evolutionary moment, albescens seems to be expand-
ing at the expense of communis and even displacing
it—a potentially instructive situation that should be
closely followed.
In the southwestern USA, where overlap is so ex-
tensive, some northern records of sympatry go further
back in time. For example, at Ft. Wingate, McKinley
County, New Mexico, P. albescens was taken on 18
June 1906 and P. communis on 19 July 1906 (AMNH);
and at St. George, 2800 ft (855 m), Washington
County, Utah, both species were taken on 10 June
1919 by T. Spalding (AMNH). Defying a strictly low-
land pattern, albescens (1 2) flew with communis (11 6
in all) at Loop Camp, 7400 ft (2255 m), 13 mi (21 km)
southwest of Grantsville, Tooele County, in northern
Utah between 16 & 20 July 1958, F., P., & J. Rindge
(AMNH). In a large majority of cases, the sympatry of
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
P. communis and P. albescens indicated by half-dots in
Figs. 21 and 22 has also involved synchrony.
Where P. communis and P. albescens coexist, their
mutual spatial relations are no doubt dynamic and,
therefore, rather unpredictable. Pyrgus communis has
long been known as a mobile species whose northern
distributional limit fluctuates considerably. Both
species regularly invade weedy, disturbed habitats.
Though predominantly austral, albescens shows up
high on the tops of various mountain islands (see Ma-
terials and Methods and Relevant Background for
some specific examples). Even the Portal, Arizona,
sample of Pyrgus (Figs. 23-141) shifts, over 26 days,
from mostly albescens to mostly communis.
GENITALIC VARIATION
Although the rampant variation in the male genitalia
of the Portal, Arizona, sample of Pyrgus (Figs. 23-141)
may look continuous at first, it clusters about two dis-
tinct modes. The variation around each of these modes
is extraordinary. In P. communis, with the higher, more
massive valve, the diagnostic valval process varies so
much in its length and width, and in the relative devel-
opment of its paired, terminal prongs, that no two in-
dividuals are exactly alike. In some of the more ex-
treme individuals, the lower terminal prong is weakly
developed (Figs. 76, 100, 123, 137) or vestigial (Figs.
50, 120, 124) or completely missing (Figs. 23, 87, 113,
129, 139)—yet even then the resulting, simpler, one-
prong process varies from narrow (Figs. 87, 129, 139)
to intermediate (Fig. 113) to wide (Fig. 23). Occasion-
ally, an extra terminal prong appears (Fig. 99). Most
individuals express the typical two-prong process
(Figs. 26, 30, 68, 69, 75, 83, 86, 88-90, 93, 95-98,
100-102, 105, 106, 108, 112, 114-119, 121, 122,
125-128, 130-136, 140, 141), but each in his own way;
and in one, this process is uncommonly short (Fig.
138). It is no wonder that Lindsey (1939), in measur-
ing four aspects of the valval process and its prongs in
100 males of P. communis, obtained such enormous
coefficients of variation.
Again, in P. albescens, with its lower, leaner look, the
variation in the valve is so great that every individual is
visibly unique—but none occupies the gap between
albescens and communis. The dorsodistal end of the
valve, which rarely is fairly even (Figs. 49, 62), usually
shows traces (Figs. 28, 32, 33, 38, 41, 42, 44, 47, 51, 52,
55, 61, 64, 70, 72, 73, 82, 84, 92, 94, 103, 104,
109-111) or real beginnings (Figs. 24, 27, 29, 31, 34,
45, 54, 56, 57, 60, 66, 74, 77, 79, 80, 85, 91, 107) or
clear expressions (Figs. 25, 36, 37, 39, 40, 43, 46, 48,
53, 58, 59, 63, 65, 67, 78, 81) of one or two teeth,
which rarely develop further into incipient but modest
VOLUME 54, NUMBER 2
<
142 143 144 145 146
147 148 149 150 151
152 153 154 155 156 157
158 ay 160 161 162
163 164 ah a 165 166 167
ee Ree,
168 169 170 171 172 173
174 175 176 171 178
179 180 181 182 183
Fics. 142-183. Genitalia of 42 6 Pyrgus communis from Galivants Ferry, Horry County, South Carolina, USA, 16 August 1957 to 27 Sep-
tember 1981, J. M. Burns (USNM)); distal end of left valva in lateral view.
64 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
185 188
S
190
196
01 9
2
1
SS. ee
oe
=o
So
197
So
oe
184 86
194
200 2
05
11
Sic gl ae
Fics. 184-220. Genitalia of 37 5 Pyrgus communis from Meade County, South Dakota, USA, 27 to 30 July 1975 (USNM)); distal end of left
valva in lateral view.
VOLUME 54, NUMBER 2 65
5 ae) 321
331 332 333
ow
i) EQ
o
339
w
w
€
w
w
co
343 344 345
w w
‘, :
—= iY
w
ai
4
w
a
—
362 363
367 368 369
373
Fics. 313-374. Genitalia of 62 6 Pyrgus albescens from the region of Laguna Chapala, Punta Prieta, Bahia de Los Angeles, Rancho Rosar-
ito, and Mission San Borja in southern Baja California Norte, MEXICO, 28 March to 2 April 1973, J. Donohoe, J. Doyen, D. Patterson, J. Pow-
ell (CAS): distal end of left valva in lateral view.
322 323 ae
“> 329 330
334 335 336
340 341 342
346 347 348
352 ) 2 354
358 359 360
364 365 366
370 371 372
374
68
}
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fic. 375. Detailed geographic distribution of Pyrgus communis and P. albescens in Louisiana and Mississippi, USA, based on male genitalia,
which appear (distal ends of left valvae) in lateral view.
289-312), and southern Baja California Norte (Figs.
313-374), both of which are well removed from any
contact with communis (Figs. 21, 22). And, finally, for
both species where they are in contact and narrowly
sympatric, these are Louisiana and Mississippi (Fig.
375).
Most extreme among P. communis genitalic variants
are those that have more or less lost the lower of the
two prongs on the long valval process. Because I en-
countered a number of these individuals (Figs. 23, 50,
87, 113, 120, 124, 129, 139) in the Portal sample of
mixed Pyrgus, it might be argued that loss of the prong
reflects genetic input from albescens. But nothing
would be further from the truth. Such one-prong vari-
ants have surfaced again and again in many different
populations of communis—including not only those
that are sympatric with albescens (as at Austin, Texas
[Figs. 225, 228, 248]) but also those that are decidedly
allopatric (as in South Carolina [Fig. 169] and, most
notably, South Dakota [Figs. 191, 192, 196, 197, 212,
915, 217).
Genitalic variants of P. albescens that tend most to-
ward P. communis are those whose two teeth at the
distal end of the valve are appreciably elevated as, for
example, at Portal, Arizona (Figs. 36, 39, 58, 59, 67),
and Harlingen, Texas (Figs. 269, 279)—both areas of
VOLUME 54, NUMBER 2
69
Fics. 376, 377. 246 Pyrgus albescens (left) and 24 3 P. communis (right) from the same time and place (midsummer 1974, Portal, Chiric-
ahua Mountains, 4800 ft [1465 m], Cochise County, Arizona, USA, J. M. & S. N. Burns [|USNM]) arranged in pinning units of the same size to
show the greater average wingspread of P. communis.
sympatry with communis and therefore of potential in-
fluence from it. But similar variants appear where
albescens is well separated from communis, as at San
Diego, California (Figs. 291, 294, 296, 311). Indeed,
the most extreme variants of this kind have turned up
in southern Baja California Norte (Figs. 325, 335, 344,
345, 348, 368, 373 and especially Figs. 331, 349, 366),
where albescens is about as far removed and isolated
from communis as it can be.
In light of this analysis, the picture of genitalic varia-
tion in Louisiana and Mississippi (Fig. 375) clearly
shows P. communis and P. albescens meeting and slightly
overlapping in space without genetically merging.
SIZE DIFFERENCE
It is always more satisfying to be able to bolster a
difficult species separation based on subtle genitalic
distinctions with evidence of another kind.
In his minimal original description of P. occidentalis
(=albescens), Skinner (1906a:96) said, “This is a
smaller . . . species than tessellata” (=communis); and,
soon after, he claimed (Skinner 1906b:278)—with no
70
TABLE 1. Length (mm) of right forewing in Pyrgus males from
Portal, Chiricahua Mountains, 4800 ft (1465 m), Cochise County,
Arizona, USA, July to August 1974, J. M. & S. N. Burns (USNM).
Species N Range Mean + SE SD CV
12.0-14.9 13.72+0.08 0.61 4.45
13.2-15.6 1458+0.07 0.54 3.70
albescens 63
communis 54
detail, explanation, or justification—“It expands in the
¢ 25 mm.; whereas tessellata expands 32 mm. This is an
average size for the two.” Tilden (1965:92) observed,
“In long series, P. communis appears a bit larger. . . . P.
albescens in series appears somewhat smaller. . . . The
smaller average size . . . of P. albescens might be ex-
pected of a desert population, as compared with a re-
lated population living in a more temperate climate.”
But, like Skinner, he offered no supporting data.
In truth, P. albescens really is a little smaller than P.
communis. The 1974 Portal, Arizona, sample is ideal
for comparing size in these species because both were
caught in numbers at the same time and place and
presumably had weathered similar environmental con-
ditions. A slight average difference in size becomes
readily perceptible when the mounted, genitalically
determined males of each species are segregated into
adjacent, identical pinning units (Figs. 376, 377).
Winglength measurements with a pair of vernier cali-
pers show an average difference of 0.86 mm (Table 1).
A quarter century later, on 15 and 16 August 1999,
Sarah and I caught 7 6 P. albescens and 21 6 P. com-
munis at an elevation of 4000 ft (1220 m) in Sycamore
Canyon, Santa Cruz County, Arizona. Mean forewing
lengths of these coexisting albescens and communis are
13.41 mm and 14.25 mm, respectively—for an equiva-
lent average difference of 0.84 mm.
ACKNOWLEDGMENTS
For lending museum material or letting me study it in situ, I
thank Frederick H. Rindge and the American Museum of Natural
History (AMNH), New York, New York, Eugene G. Munroe and the
Canadian National Collection, Ottawa, Ontario; Paul H. Arnaud Jr.
and C. Don MacNeill and the California Academy of Sciences
(CAS), San Francisco, California; Jerry A. Powell and the Essig Mu-
seum of Entomology, University of California, Berkeley (UCB), Cal-
ifornia; Julian P. Donahue and the Natural History Museum of Los
Angeles County (LACM), Los Angeles, California; John B. Heppner
and the Florida State Collection of Arthropods, Gainesville, Florida;
Richard L. Brown and the Mississippi Entomological Museum
(MEM), Mississippi State, Mississippi; Charles V. Covell Jr. and the
Lovell Insect Museum, University of Louisville, Louisville, Ken-
tucky; Brett C. Ratcliffe and the University of Nebraska State Mu-
seum, Lincoln, Nebraska; the Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts; and the National
Museum of Natural History (USNM), Smithsonian Institution,
Washington, D.C. For donating or lending personal material, I
thank Richard A. Anderson, Richard A. Bailowitz, John V. Calhoun,
Steven J. Cary, Ken Davenport, Richard Holland, Roy O. Kendall,
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Bryant Mather, William W. McGuire, Paul A. Opler, Jack R. Powers,
Stephen M. Spomer, and Gayle T. Strickland. For allowing access to
their alfalfa field, the Trollers. For KOH-dissecting genitalia, John P.
Basinger, Steven I. Cohen, Rogene G. Gillmor, Judy Gilmartin, and
Barbara L. Scott. For dry-dissecting many genitalia and drawing
these, as well as the KOH-dissected ones, in pencil on file cards,
Richard G. Robbins. For inking many of Robbins’s drawings, Young
T. Sohn (Figs. 142-375). For silhouetting Robbins’s drawings of the
Portal, Arizona, sample and for illustrating complete genitalia,
George L. Venable (Figs. 9-18, 23-141). For photographing geni-
talia and adults, Victor E. Krantz (Figs. 1-8, 19, 20, 376, 377). For
catching Pyrgus, producing computerized lists of specimens exam-
ined, and helping in many and various ways, Sarah N. Burns. For
providing financial support, the National Science Foundation (grants
GB 5935 and GB-37832) and the Smithsonian Institution (grants
from the Research Opportunities Fund in 1987, 1989, 1991, and
1999). For reviewing the manuscript, John W. Brown, Arthur M.
Shapiro, Felix A. H. Sperling, Andrew Mitchell, and Deane Bowers.
LITERATURE CITED
AusTIN, G. T. 1986. Pyrgus communis and P. albescens (Hesperi-
idae) in Nevada. J. Lepid. Soc. 40:55-58.
. 1998. Checklist of Nevada butterflies, pp. 837-844. In Em-
mel, T. C. (ed.), Systematics of western North American but-
terflies. Mariposa Press, Gainesville, Florida.
BaILowiTz, R. A. & J. P. BRocK. 1991. Butterflies of southeastern
Arizona. Sonoran Arthropod Studies, Inc., Tucson, Arizona. ix +
342 pp.
Barnes, W. & A. W. LINDSEY. 1921. On some species of Hesperia
(Lepid., Hesperidae). Entomol. News 32:78-80.
Brown, J. W., H. G. REAL & D. K. FAULKNER. 1992. Butterflies of
Baja California. Lepidoptera Research Foundation, Inc., Bev-
erly Hills, California. v + 129 pp., 8 pls.
Burns, J. M. 1964. Evolution in skipper butterflies of the genus
Erynnis. Univ. Calif. Publ. Entomol. 37:1-217.
. 1997. Presidential address 1996: On the beauties, uses, vari-
ation, and handling of genitalia. J. Lepid. Soc. 51:1-8.
Burns, J. M. & R. O. KENDALL. 1969. Ecologic and spatial distribu-
tion of Pyrgus oileus and Pyrgus philetas (Lepidoptera: Hes-
periidae) at their northern distributional limits. Psyche
76:41-53.
Cresson, E. T., JR. 1918. Entomological Section, The Academy of
Natural Sciences of Philadelphia [minutes of meeting]. Ento-
mol. News 29:37-38.
Dos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalo-
cera. Lepid. Soc. Mem. No. 1. v + 145 pp.
EMMEL, J. F., T. C. EMMEL & S. O. Mattoon. 1998. A checklist of
the butterflies and skippers of California, pp. 825-836. In Em-
mel, T. C. (ed.), Systematics of western North American but-
terflies. Mariposa Press, Gainesville, Florida.
Evans, W. H. 1953. A catalogue of the American Hesperiidae indi-
cating the classification and nomenclature adopted in the
British Museum (Natural History). Part II. Pyrginae. Section
2. British Museum, London. 246 pp., pls. 26-53.
Grote, A. R. 1872. On a new checkered Hesperia. Canad. Entomol.
4:69-70.
Kuots, A. B. 1951. A field guide to the butterflies of North America,
east of the Great Plains. Houghton Mifflin Co., Boston. xvi +
349 pp., 40 pls.
LayBERRY, R. A., P. W. HALL & J. D. LAFONTAINE. 1998. The butter-
flies of Canada. Univ. Toronto Press. 280 pp., 32 pls.
Linpsey, A. W. 1921. The Hesperioidea of America north of Mexico.
Univ. Iowa Stud. Nat. Hist. 9:1-114.
. 1939. Variations of insect genitalia. Ann. Entomol. Soc.
Amer, 32:173-176.
LinpsEy, A. W., E. L. BELL & R. C. WILLIAMS Jr. 1931. The Hespe-
rioidea of North America. Denison Univ. Bull., J. Sci. Lab.
26:1-142.
Mayr, E. 1942. Systematics and the origin of species. Columbia
Univ. Press, New York. xiv + 334 pp.
VOLUME 54, NUMBER 2
MiILier, L. D. & F. M. Brown. 1981. A catalogue/checklist of the
butterflies of America north of Mexico. Lepid. Soc. Mem. No.
2. vii + 280 pp.
. 1983. Hesperioidea, Hesperiidae, pp.42—48. In Hodges, R.
W. et al. (eds.), Check list of the Lepidoptera of America north
of Mexico. E. W. Classey Ltd. and The Wedge Entomol. Res.
Found., London.
PLOTZ, C. 1884. Analytische Tabellen der Hesperiinen-Gattungen
Pyrgus und Carcharodus. Mitth. nat. Ver. Vorpomm. 15:1—24.
POWELL, J. A. 1958. Additions to the knowledge of the butterfly
fauna of Baja California Norte. Lepid. News 12:26-32.
REVERDIN, J.-L. 1921. Hesperia tessellata Scudder. Var. occidentalis
Skinner. Pyrgus montivagus Reakirt. Syrichtus notatus Gay.
Bull. Soc. Lépid. Genéve 4:168-185, pls. 6, 7.
SKINNER, H. 1906a. New butterflies and moths with notes on some
species. Entomol. News 17:95-96.
. 1906b. Studies of Pyrgus syrichtus, tessellata, occidentalis,
and montivagus. Entomol. News 17:277-278, pl. XIL.
SKINNER, H. & R. C. WILLIAMS JR. 1923. On the male genitalia of
the Hesperiidae of North America, Paper II. Trans. Am. Ento-
mol. Soc. 48:283—306.
TILDEN, J. W. 1965. A note on Pyrgus communis and Pyrgus
albescens (Hesperiidae). J. Lepid. Soc. 19:91—94.
TILDEN, J. W. & A. C. Smiru. 1986. A field guide to western butter-
flies. Houghton Mifflin Co., Boston. xii + 370 pp., 48 pls.
Received for publication 5 August 1999; revised and accepted 30
March 2000.
————————————————————————
|
|
|
i}
I
i
i
i
t
Journal of the Lepidopterists’ Society
54(2), 2000, 72-75
A NEW SPECIES OF OMIODES GUENEE FROM SOUTH AMERICA (PYRALOIDEA: CRAMBIDAE)
M. ALMA SOLIS
Systematic Entomology Laboratory,
Plant Sciences Institute,
Agricultural Research Service, USDA
c/o Nat. Mus. Nat. Hist., MRC 168,
Washington, D.C. 20560-0168, USA
PATRICIA GENTILI
Dept. of Entomology, MRC 127,
Smithsonian Institution,
Washington, D.C. 20560-0127, USA
ABSTRACT. Omiodes pseudocuniculalis, new species, is described from South America where it is recorded from Peru, Ecuador, and
Bolivia. Five males were examined, and no females are yet known. The male and its genitalia are illustrated showing the diagnostic character of
short, protruding lobes located along the medial edges of the ventral groove of the uncus. The new species is most similar to O. cuniculalis
Guenée and externally resembles other members of the “cuniculalis group” of Omiodes by the cinnamon or brown color of the adults, the beige
or brown color of the ventral surface fo the the head, neck, thorax, and abdomen, and legs, and the brown color of the anal tuft. The key to the
“cuniculalis group” is revised to include the new species.
Additional key words: | Fabaceae, Western Hemisphere, genitalia morphology, parategumenal sclerites.
The genus Omiodes Guenée (Crambidae) was re-
cently redescribed within the context of a study of the
Pyraloidea of Costa Rica, and a checklist and key to
New World species were developed (Gentili & Solis
1998). It consists of 74 species worldwide, 33 species
in the Western Hemisphere, and includes Omiodes in-
dicata (F.), a worldwide pest on many crops in the
Fabaceae. Externally, the adults of this genus are di-
verse: “brown and noctuid-like, bluish and arctiid-like,
and yellow reddish and pyraloid-like”; the forewing
lengths vary from 7 to 26 mm long; the forewing is
elongated, the termen straight, but the hindwing is
usually round, in some species triangular; the thorax
and abdomen in some species with bright-yellow ar-
eas, but generally same background color as wings
(Gentili & Solis 1998).
During the study of Costa Rican Omiodes we also
discovered a new species from northern South
America, which we describe in this paper. The diagno-
sis and description of the new species is based on a
comparison with all Omiodes species of the Western
Hemisphere. The new species, Omiodes pseudocunic-
ulalis, belongs to a group of closely related species that
Gentili and Solis (1998) called the “cuniculalis group”.
This group includes O. cuniculalis Guenée, O. fulvi-
cauda (Hampson), O. anxiferalis (Schaus), O. al-
boanalis Amsel, and O. martini Amsel. They are dis-
tinguished from species in other Omiodes groups by
the cinnamon or brown color of the adults, the beige
or brown color of the ventral surface of the head, neck,
thorax, abdomen, and legs, the brown color of the anal
tuft, the uncus head with a crest with erect hairlike se-
tae and scale or spines in patches and lateral lobes
pendant.
Omiodes pseudocuniculalis Solis and Gentili,
new species
(Figs. 1, 4-8)
Diagnosis. Lobes along medial edges of ventral groove of uncus
neck short and protruding.
Description. Male. Head: light brown. Antenna light brown.
Labial palpus cream at base changing gradually to light brown at
tip. Patagium ventrally cream. Thorax: light brown. Tegula reaching
abdominal tergum II. Legs light brown, gradually lighter brown to-
ward and including tarsi. Forewing length: 16.7-18.3 mm (n = 5).
Forewing ground coloration light brown, slightly darker brown be-
yond postmedial line; wing pattern darker brown, postmedial line
zigzag; fringe with proximal row of scales same as ground color and
outer row lighter brown with area between 1A+2A and CuA,
cream. Hindwing same ground color as forewing, darker beyond
postmedial line; postmedial line zigzag with inner angles darker,
giving impression of dots; fringe as in forewing, outer row of scales
cream from apex to 1A+2A. Abdomen: light brown, ventrally lighter
and shinier. Posterior margin of male tergum VI convex; anal tuft
light brown. Male genitalia: uncus base slightly wider than neck;
neck with ventral groove present medial edges of apical half with
two well-delineated short, round, lobes with hairlike setae; head
with crest, two tufts of erect scales; apical tip upturned with brush
of hairlike setae; lateral lobes with hairlike setae. Ventral margin of
valva at end of sacculus not swollen; hooklike process as long as
sclerotized base along saccular margin; setose patch small.
Transtilla continuous. Cornutus spine short, about half length of
aedeagus; ductus ejaculatorius short. Parategumenal sclerites one-
fourth surface area of valva [The term “coremata” was incorrectly
used for this structure in Gentili & Solis (1998). To distinguish this
structure from the “coremata” of the Arctiidae, Clavijo (1990)
coined the term “parategumen sclerites” for these coremata-like
structures in the Pyraustinae and defined them as: “. . . A pair of lat-
eral sclerites located each in the basal region of the tegumen, typi-
cally containing long pencils or brushes of modified hair-like and/or
VOLUME 54, NUMBER 2
Fic. 1. Paratype adult male of Omiodes pseudocuniculalis, Ecuador, Environs de Loja. Wing length = 16 mm.
spatulate scale, presumably for scent production and/or distribu-
tion.” ] Female: unknown.
Types: Holotype: J, ECUADOR: “Environs de Loja,” “Equa-
teur,” “89-” “Dognin Collection,” genitalia slide number 105,812
[USNM]. Paratypes: 4 paratypes: ECUADOR: 1 4, “Environs de
Loja,” “Equateur,” “Dognin Collection,” genitalia slide number
106,832 [BMNH]. PERU: 1 4 , no label data, “Peru,” “Collection
Wm. Schaus,” genitalia slide number 105,811 [USNM]. BOLIVIA:
1 d, “Charuplaya,” “Juin 1901,” “Bolivie,” “Saison séche,” “Dognin
Collection,” genitalia slide number 105,809 [USNM]. UNKNOWN
LOCALITY: 1 4, “Dognin Collection,” genitalia slide number
105,810, head slide number 106,735, wings slide number 106,737,
legs slide number 106,736 [USNM]. Paratypes deposited in The
Natural History Museum, London, England [BMNH] and The Na-
tional Museum of Natural History, Smithsonian Institution, Wash-
ington, D.C., USA [USNM].
Remarks. Omiodes pseudocuniculalis is closely related to and
could be confused externally with O. cuniculalis, a species that oc-
curs from Mexico south to Brazil and Colombia, including the
Caribbean; it is difficult to determine whether these species are
sympatric because these areas are so poorly collected. Although we
have not seen specimens of O. cuniculalis from Ecuador, Peru, or
Bolivia, where O. pseudocuniculalis occurs.
We compared specimens with the types deposited at the BMNH
of O. cuniculalis, a male from French Guiana. Externally O. cwnicu-
lalis and O. pseudocuniculalis are very similar in color and size. In
both species the lobes on the medial edges of the ventral groove are
round with hairlike setae, but in O. cuniculalis they occur on the basal
half and are not well delineated while in O. pseudocuniculalis they oc-
cur on the apical half, are short, well delineated, and protruding.
Etymology. This species is named in reference to the close ex-
ternal resemblance to O. cuniculalis.
REVISED KEY TO SPECIES OF THE
“CUNICULALIS GROUP” OF OMIODES
1. Ventral surface of body and legs light brown, never bright
yellow, general body coloration cinnamon or brown, wing
pattern darker brown, postmedial line zigzag, fringe
ochraceous; uncus head with crest, medial edge of neck
ventral groove Withoutssenrateraneci) sienna een tenn neers 2
Ventral surface of body and legs bright yellow, some
specimens with terga VII and VIII yellow, general body
coloration dark brown, postmedial line not zigzag, lines
slightly darker, fringe white; uncus head without crest,
medial edge of neck ventral groove with serrate area
Fos aud Barat co Ned Gece are a eas ance Rea te fulvicauda
2. Cinnamon, homogeneous; uncus head with central
patch of two tufts of erect scales, medial edge of neck
ventral groove with lobes along edges...................- 3
Light to dark brown, sometimes forewing intermedial area
cinnamon; uncus head with two lateral patches of short
spines, medial edge of neck ventral groove without lobes ... . 4
3. Medial edges of uncus neck ventral groove lacking well-
delineated lobes along basal half (Figs. 2,3) ....... cuniculalis
Medial edges of uncus neck ventral groove with well-
delineated lobes along apical half (Figs. 4, 5)
NAPE ya a Potten ANS cth Tle. dee SrA ieee pseudocuniculalis
4. Wings unicolorous; medial edge of neck ventral groove
Huta lod GAO 5 os5ccceonoconscuvcuenssuoos alboanalis
74 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
uncus head
uncus neck
cornutus
fibula
transtilla
Is, Zt
Fics. 2-8. Male genitalia of O. cuniculalis and O. pseudocuniculalis. Scale line = 1.0 mm; 2, ventral view of O. cuniculalis uncus, USNM
slide 104,743; 3, lateral view of O. cuniculalis uncus, USNM slide108,814; 4, ventral view of O. pseudocuniculalis uncus, USNM slide106,832;
5, lateral view of O. pseudocuniculalis uncus, USNM slide106,832; 6, parategumenal sclerite of O. pseudocuniculalis, USNM slide106,832; 7,
valva and transtilla of O. pseudocuniculalis, USNM slide 106,832; 8, aedeagus of O. pseudocuniculalis, USNM slide106,832.
VOLUME 54, NUMBER 2
Wings not unicolorous, with intermedial area lighter in
color; medial edge of ventral groove of neck without
naimlikexsetacanwa cea sy ws aera ner e) & aa Ne leAncg ah aioe « 5
5. Uncus head with lateral patches of spines almost completely
meeting dorsomedially, crest conspicuous; transtilla con-
tinuous, same width throughout ................. anxiferalis
Uncus head with lateral patches of spines not meeting
dorsomedially, crest not conspicuous; transtilla divided,
oxoadpnite dially arpa amrn os dein crosman nna ae martini
ACKNOWLEDGMENTS
We thank Susan Escher for most of the illustrations, and Linda
Lawrence for the plates. We thank Michael Shaffer at The Natural
History Museum, London, England, for his hospitality. A. Konstan-
tinov and D. Smith of the Systematic Entomology Laboratory, PSI,
ARS, USDA, Washington, D.C. and J. Shaffer of George Mason
University, VA, USA, for helpful comments on the manuscript.
LITERATURE CITED
Ciavyo, J. A. 1990. Systematics of black and white species of the
genus Diaphania Hiibner (1818) (Lepidoptera: Pyralidae:
Pyraustinae). 215 pp. Dissertation, McGill University, Montreal.
GENTILI, P. & M. A. Souts. 1998. Checklist and key of New World
species of Omiodes Guenée with descriptions of four new Costa
Rican species (Lepidoptera: Crambidae). Entomol. Scand. 28:
471-492.
Received for publication 14 October 1998; revised and accepted 30
March 2000.
sasausilessnasessaeesiteeeesaeaeeeseenenesee
BSE ROS TS Oe ee
BOOK REVIEWS
Journal of the Lepidopterists’ Society
54(2), 2000, 76
REVISTA DE THECLINAE COLOMBIANOS/ REVIEW OF COLOMBIAN
THECLINAE, by Museo de Historia de National de la Universidad de
Caldas, Manizales, Museo de la Universidad Pedagégica Nacional,
Santefé de Bogoté, Museo De La Salle de Ciencias Naturales, San-
tefé de Bogota. Volumes 1 & 2, no actual date for publication is
given, but the mailing date is given as 15 August 1997. Overall di-
mensions of both publications, 5 1/2 by 8 1/2 inches. Volume 1 con-
tains 179 pages of text with numerous line drawings, 2 color plates,
and 4 plates of black and white photos; volume 2 contains 178 pages
of text with numerous line drawings, 6 color plates, and 6 plates of
black and white photos. Text in English and Spanish. Soft covers.
ISSN 0123-1677, free, available from Dr. Kurt Johnson, Environ-
mental Affairs, The Ethical Culture Society, 53 Prospect Park West,
Brooklyn, NY USA 11215.
According to the authors these two volumes are but the begin-
ning of a series of volumes that will cover the entire spectrum of the
Theclinae of Colombia with the goal of producing at least one vol-
ume per year. The title is a bit misleading in that both volumes are a
collection of research works and papers, some previously unpub-
lished, containing many newly described genera and species. Very
little time or space is devoted to the many species previously named
by others that occur in Colombia. Some of the described species in
the many newly described genera do not occur in Colombia, but be-
cause they are a part of the original papers involved, appear in these
volumes. One may readily determine the titles, the authors, and a
brief of the contents of each paper on the outer back cover of each
volume. There are 8 papers listed in volume 1, and 10 in volume 2.
A number of typographical errors and the incredibly disjointed
and complex format of the entire text and illustrations make these
volumes difficult to read and to follow logically and easily. To com-
pound the entire matter, the size of the font is reduced in many in-
stances for reasons that escape this reviewer.
It would be presumptuous of this reviewer to comment specifi-
cally on the array of new species and, more particularly on the many
new genera that have been erected in this publication. I have col-
lected rather extensively in South America and Panama (and yes, in
Colombia) since 1962, and I am surprised to note that most, if not
all, of the many specimens figured in these volumes with which I am
familiar, now sport new specific names and even more amazing, be-
long to new and different genera, in many cases a new genus for
each species!
As a case in point, the first work in volume 1 is the description of
Strephonina, a “New Infratribe” of the Eumaenini. This is a new
term to this reviewer and I can find no definition for it in this work.
The Strephonina contains fourteen new genera. The only genus
listed as containing more than a single species is Strephonota which
contains 4 species. Some of the remainder contain more than a
single species, but one has to search the text carefully in order to as-
certain this fact.
In volume 2 another new infratribe carries the title of Macusiina.
It appears to be a collection of very different and disparate species,
especially when one looks at the array of genitalia sketches repre-
senting the various species. Again, one must search the text carefully
in order to determine the species contained in each of these newly
described genera. This is as good a place to mention this: there is no
complete index of species or genera in either volume. If one wishes
to locate a particular species or genus in either volume, one must
search the entire volume, page by page. It is time consuming and
frustrating.
Finally in volume 2, the species of Strymon illustrated and de-
scribed as “new” left me shaking my head and wondering if I had
read and viewed the descriptions and plates correctly. Strymon meli-
nus (Hiibner, 1813) herein named and figured as Strymon caldasen-
sis is a dead ringer for any number of specimens in the collections of
various individuals and museums I have observed and studied in the
past 50 years. It is one of the most common and widespread eu-
maeines in the Western Hemisphere. Although not as common and
widespread, three other examples of this distortion of names of fa-
miliar species are contained on the same color plate: Strymon rufo-
fusca (Hewitson, 1877), named herein as S. guanensis; Strymon
bubastus (Cramer, 1782), herein named S. vividus, and Strymon
gabatha (Hewitson, 1870), herein named S. alicia. I have studied
the descriptions and photos of these “newly described” species and
can only surmise that the widely held and trusted concept of varia-
tion within a species is no longer a valid concept.
There are a number of disturbing factors in these two volumes,
but the most glaring error noted in this cursory reading of the 2 vol-
umes is to be found on photoplate XIV, figure H which bears the
name Strymon carmencitae Le Crom & Johnson. The taxon figured
here and elaborated upon in the text is not a Strymon and it does
have a name — Hewitson named this species in 1868. This species
does not bear the characteristic Strymon characters of the genital
ring, valvae, aedeagus, or the scent pad on the forewing of the male.
It is not a common species, but I have studied specimens from
Brazil as well as Colombia
Finally, if one wishes to be overwhelmed by a profusion of new
names for familiar species, then these two volumes will fit the bill.
The appropriate text is difficult to read, the plates and line drawings
are not in such a sequence as to be easily followed and related to the
text, and there are no legend numbers or notes on the plate figures
that allow easy cross reference to the appropriate text. These fea-
tures, added to the lack of any index to the species and/or genera in
either volume, make them difficult to use.
S. S. Nicoxay, 1500 Wakefield Drive, Virginia Beach, Virginia,
23455, USA.
Date of Issue (Vol. 54, No. 2): 15 November 2000
|. EDITORIAL STAFF OF THE JOURNAL
M. Deane Bowens, Editor
- Entomology Section
University of Colorado Museum
Campus Box 218
_. University of Colorado
Chee Boulder, Colorado 80309, U.S.A.
pide ‘ ees os 2: email: bowers@spot. colorado.edu
Sphere ere ee its : Associate Editors:
ei me ai Roesiiae ice peal KENELM W. Pure (USA), Moaeer K. Rospins (USA),
Feu A, H. SPERLING (UB), ‘Davip L, Wacner (USA), Origeren WIKLUND evades)
2 be sn om _ NOTICE TO. CONTRIBUTORS
eature eee aoa and Cover Illustrations. Dottie: must be eibaeed by the Presidzavolthe Society. Requirements
empl: ae Cover Ilustrations a are » stated on page 11lin Volume 44(2). Journal submissions amoeld be sent to the editor at the above
ca v witl sade margins, on one ae mate of white, eke ea paper. fare manuscripts aie to the iit
cee ubmit them flat, not fol ed. perligts are’ eppeted to Broads a ' diskette cenys of ae ble accepted see in'a standard PC or
d word processing format.
> abstract chould orale He Ser a ee and Profiles. at:
ad to aye ae words or terms not in, a ioe should ay ay Biel Profiles, General Notes, and Technical Com-
NE ae i as s adults ith wings ee should be illustrated: apiee hole oe is crucial. Pho-
ounted on stiff, white backing, arranged i in the desired format, allowing (with particular regard to lettering) for re-
ustrations larger than letter-size are not acceptable : and should be reduced photographically to that size or smaller. The
numbers as ‘the te ould be printed on the back of each illustration. Figures, both line drawings and photographs,
n amb od cons utively in Arabic numerals; “plate” should not be employed. Figure legends must be typewritten, double-spaced, on a sep-
not “att shed to ill ations), he led ExrLanarion or Figures, with a separate as devoted to each page of esd Color illus-
: contact the. editor for submission OO) ae and cost. °
7 ; that anole eae siclude life hisiede act associations, immature One and ear enquiries.
ee edited ee and cee ir will be aa to the author for correction of —- errors. Excessive author's changes at this
ENT
~~ Volume 54 2001 Number 3
ISSN 0024-0966
JOURNAL
of the
EPIDOPTERISTS’ SOCIETY
Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTEROLOGOS
Sw THSON A
OCT 04 2001
LIBRARIES
ty
by both the professional worker and da amateur in the field; to secure cooperation in all measures” directed t owards these aims
and special lepidopterological interests. In alternate years a list of members of the Society i is eke wth, addresses
Au me ae ae em a _ Student members—annual dues $20.00 within the U.S., $25.00 outside the U.S.
fe sites to: Julian’ P. Denwhee. Natural a Miaseuth. ‘900 oe Bhd, i anes t cA 90007-4057. For
a POSTMASTER: Send daddies chats to The oon Society, To Neteat eis! Museum, 900. ee Bed, , Los ‘ng :
“Execunve Courcn,
Ta Va igen Wy. Brown, Peder? : Pama wae None Sean's. ae Vice P
Micuaet J. Srru, Immediate Past President Vee Casncnanne, Vi
~Manue A. Barcazan-Lara, Vice President
Envest H, WILLIas, Meee
STEMS Reon Seine ieee euint Mente tees
tg Le Hutowsle «MM. Deane Bowers —
Felix A, Hi Sperling <* Ron Leuschner
»Andrew.D. Warren’ Michael Toliver i on ‘
anon Boarp
_ Ropert K. se (Chairman), Joun: W. BAG (Member at large)
4 M. Deane Bowers (Journal) Be
Witiiam E. MiLver (Memoirs) _ SaaS a A
Vee ds Scuaprenr (News)
"Honorary Lire Mempers OF THE eee
Giant ee REMINGTON (1966), LE G3 Munrox (1973), Tan F. B. Chinon (1987), 3
-Joun G:; FRANcLeMont (1988), LINCOLN P. Brower (1990), Doucias C. ‘FERGUSON. (1990), -
Hon. Mina Rornsouitp (1991), Craupe eee ‘Pnepenck H. Rinpge See,
aa
y
- The object Ee The. Lepidopterists Society, whieh. was s formed i in ‘May 1947 and formally constitute’ i Daeenibey 1950 i
of lepidopterology in all its branches, . . . to issue a periodical and other publications on. Lepidoptera, to facilitate the exchange
ia
SS
fe
Soe
5.
oF
Membership in the Society is open to all persons interested in the study of Lepidoptera. All members recei
idopterists’ Society. Prospective members should send to the Assistant Treasurer full dues for the current year, together with
Dota BA a Cy auee $45, 00 within the U:S., $50. 00 patside the ws ak
Affiliated members—annnal dues $10.00 within the U.S., $15.00 outside the. U. *
Sustaining members—annual dues $60. 00 within: the a S.. gers 00 outside se U. 8.
Life members—single sum $1,800.00 + tay %
Institutional subseriptions—annual $60.00.
Airmail rors for the News Rue By
ety, contact: Ernest H. Williams, Department of Biology, Hamilton College, Clinton, NY 13323. To order back i issues of the Memoirs, y
ee and prices to Kelly M. Richers, Asst. ‘Treasurer, 9417. Sale Court, ‘Bakersfield, ( CA 93311. GOLAN,
The hte cost for members outside the U.S. is to cover mailing Bee
Natural ioe 900 a Blvd., Los Andee “CA 90007-4057. Periodicals eae an at Los eee CA ae at ae
5
Cover ater Last instar larva of the Tomato Hornet ioaduae sexta (Sphingidae). Ink ns bes iene
JOURNAL OF
THE LEPIDOSTEMI mS: SOCIETY
Volume 54
Journal of the Lepidopterists’ Society
54(3), 2001, 77-82
2001 Number 3
ONTOGENETIC CHANGES IN LEAF SHELTER CONSTRUCTION BY LARVAE OF
EPARGYREUS CLARUS (HESPERIIDAE), THE SILVER-SPOTTED SKIPPER
Eric M. Linp,’ MEc T. JONES, JEREMY D. LONG? AND MARTHA R. WEISS®
Department of Biology, Georgetown University, Washington, D.C. 20057, USA
ABSTRACT. In this paper we examine patterns of shelter construction by larvae of the Silver-spotted Skipper, Epargyreus clarus
Cramer (Hesperiidae). Through observations of field and laboratory populations we characterize 1) the types of shelters constructed over
larval ontogeny and their relationship to larval size and instar, and 2) the location of shelters on the host plant. We also describe various as-
pects of larval feeding behavior. Each larva builds and inhabits its own shelter, successively abandoning shelters and constructing new ones
approximately five times across five instars. On kudzu (Pueraria lobata; Fabaceae), larvae produce shelters in four distinct styles that change
predictably as the insects grow. Ontogenetic changes in style of shelter construction are likely to be related to larval size, needs, and physi-
cal capability.
Additional key words: leaf folder, caterpillar behavior.
Lepidopteran larvae in at least 18 families construct
shelters from leaves that are rolled, folded, or tied and
sealed with silk (Scoble 1992). These shelters are
thought to provide a variety of advantages to the larvae,
including protection from natural enemies (Damman
1987; Ruehlmann et al. 1988), creation of a favorable
microhabitat (Henson 1958), increased leaf nutritional
quality (Sagers 1992), or protection from phytotoxicity
(Sandberg and Berenbaum 1989). Very little is known,
however, about the pattern or process of leaf shelter
construction (Clark 1936, Fraenkel and Fallil 1981,
Ruehlman et al. 1988, Fitzgerald and Clark 1994).
The vast majority of skippers (Hesperiidae) live
singly in a shelter constructed of host leaf material and
silk (Moss 1949, Scoble 1992). Shelter styles vary
among species, and also across larval ontogeny within a
single species (Scudder 1889, Clark 1936, Moss 1949).
The diversity of shelter styles includes leaf rolls, folds,
peaked tents and perforated pockets (Scudder 1889,
Moss 1949). Shelter construction may be initiated at
the leaf margin or in the center of the leaf, and may in-
volve a small portion of a leaflet, an entire leaf, or mul-
‘Current address: The Nature Conservancy, 4225 N. Fairfax Dr.,
Arlington, Virginia 22203.
* Current address: School of Biology, Georgia Institute of Tech-
nology, 310 Ferst Dr., Atlanta, Georgia 30332.
°To whom correspondence should be addressed.
tiple leaves. For certain species on a given host plant,
shelter size, style, and placement on the leaf can be di-
agnostic (H. Greeney pers. comm., J. Brock pers.
comm.).
In this paper we describe the pattern of shelter con-
struction by larvae of the Silver-spotted Skipper, Epar-
gyreus clarus Cramer (Hesperiidae). Through observa-
tions of field and laboratory populations we
characterize 1) the types of shelters constructed over
larval ontogeny and their relationship to larval size and
instar, and 2) the location of shelters on the host plant.
MATERIALS AND METHODS
Study organism. The Silver-spotted Skipper,
Epargyreus clarus, ranges throughout North America
from Saskatchewan in the north through Baja
California, Texas, and Florida in the south (Scott,
1986). In the Washington, D.C. area these large
skippers fly from mid-April through October, and
commonly use black locust trees (Robinia pseudo-
acacia) and kudzu (Pueraria lobata) (both Fabaceae)
as hosts (Clark & Clark 1951). In this study we used
kudzu as our host plant because of its abundance,
accessible vining growth form, and the longevity of cut
leaves in the laboratory.
Caterpillars inhabit leaf shelters throughout their
larval lives, leaving only to feed or to build a new shel-
TABLE 1. Summary of Epargyreus clarus larval and shelter characteristics across 5 instars. Values are given as the mean ( SE (sample size).
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
9
Instar
Larval length, mm* 5.5 + 0.1 8.4+0.1
(29) (25)
Shelter length x width, 7.1+01x 13.1+0.4x
mm §.1 + 0.2 14.7 +0.5
(shelter type)? {1} (69) {1} (31)
Shelter size / Larval length‘ 1.4 1L7/
Distance to subsequent 5.4 + 0.5 8.9+0.8
shelter, cm (55) (47)
Duration of feeding 3.4 + 0.18 5.2 + 1.2
bout, minutes (97) (8)
% daytime spent 4.3+0.4 3.7 + 1.7
feeding (61) (14)
Maximum feeding 11+0.2 1.9+0.1
distance, cm (50) (20)
Location of feeding site 100% sll 100% sll
relative to shelter® (40) (37)
3 4 5
14.3 + 0.4 26.4 + 0.5 36.2 + 1.2
(20) (21) (20)
18.8 + 1.x 36.7 + 1.6x 46.5 + 1.6x
15.9 + 0.8 18.4 + 1.2 26.3 + 2.2
{1} (15) {2, 3, 4} (25) {2, 3, 4} (15)
NBD, 1.0 1.0
114413 24.6 + 11.2 =
(20) (5)
5.1 + 1.4 2.7+ 0.2 4+ 0.6
(3) (73) (11)
1.3 + 0.7 2.1+0.6 1941.1
(13) (11) (16)
91+0.2 7.0 + 0.6 18.3 + 1.9
(20) (15) (15)
100% sll 78% sll, 14% all,
(45) 8% alf (36)
4% sll, 20% all,
76% alf (25)
“Measurements taken of mid-instar larvae
> Measurements taken of only those shelter types indicated
© Ratio of (mean shelter width + length)/2 to mean larval length
‘sll = same leaflet; all = adjacent leaflet, same leaf; alf = adjacent leaf
ter. Larvae do not feed within the shelter, although
early instars feed close by (Table 1). Based on observa-
tions of many larvae in the field, we determined that
during the day, caterpillars spend approximately 3% of
their time feeding, and that the average feeding bout
(time elapsed between a caterpillars departure from
and return to its shelter) lasts about 4 minutes (Table
1). Young larvae feed on the same leaflet that their
shelter occupies, while older larvae may venture to an
adjacent leaflet or leaf (Table 1).
Collection and care of larvae
Larvae were obtained from eggs of adults caught on
the Georgetown University campus and from meadow
habitats on the Eastern Shore of Maryland from June
through October 1997 and 1998. After the first gen-
eration, the colony contained both lab-reared and
field-collected adults. Butterflies were kept outdoors
in 2 m’ mesh cages, and were fed flowers of Trifolium
pratense (Fabaceae), Buddleja davidii (Buddlejaceae),
and Lantana camara (Verbenaceae), supplemented
with 10% sucrose solution. Freshly cut kudzu leaves
were provided for oviposition. Kudzu leaves contain-
ing eggs were collected from the outdoor cage each
morning, brought into the lab, and placed in 13” x 7.5”
x 4.25” clear plastic boxes with opaque lids. Larvae
were housed in these boxes and were given fresh cut
kudzu leaves as needed until pupation.
1) Types of shelters constructed over larval
ontogeny
To characterize patterns of shelter construction, we
collected and examined over 600 leaf shelters con-
structed by larvae in field and lab populations, and
identified the instar of the inhabitant. Based on our
observations of shelter characteristics, we developed a
classification of shelter types. We also measured the
dimensions of a subset of shelters of each type, and de-
termined the length of their larval inhabitants, using a
Mitutoyo Absolute Digimatic caliper (£0.01 mm).
To determine how patterns in the size and shape of
shelters relate to the timing of a molt to a later instar,
fifteen newly hatched larvae were placed on fresh
kudzu leaves in individual plastic boxes in the labora-
tory. Every other day, the boxes were opened and the
larval instar and house type recorded. Care was taken
not to touch or manipulate the larvae themselves.
2) Location of shelters on the host plant
In the field, we measured the height above ground of
the shelter-bearing leaf for 36 first instar shelters. Height
of first instar shelters is a good indication of height at
which eggs are laid, because empty egg shells and first
shelters are generally found on same leaflet (pers. obs.).
We also divided the leaflet into four quadrants (quadrant
1 = right apex; 2 = left apex; 3 = left base; 4 = right base)
and noted in which one a shelter was located, for 356
VOLUME 54, NUMBER 3
cb
Type 1: Two cut fold
ev
Type 2: One cut fold
79
Shelter Types
Ls
Type 3: Leaf roll
Type 4: Two leaf pocket
Fic. 1. E. clarus larvae construct 4 types of shelters over larval ontogeny. Lines connected to shelters represent silk guy wires.
shelters made by first, second and third instar larvae. We
often encountered leaflets or leaves that contained three
or four shelters which we inferred were constructed by a
single larva, based on the predictable progression of
shelter size and style. We used a ruler to measure the
shortest linear distance between sequential shelters.
RESULTS
1) Types of shelters constructed over larval
ontogeny
We found that Epargyreus clarus larvae construct
shelters in four distinct styles (Fig. 1), designated types
1, 2, 3, and 4 for the approximate order in which they
appear in larval life. For shelter type 1 (‘two-cut
folds’), the larva makes two cuts of precise length and
orientation in from the margin of the leaflet, applies
multiple strands of silk at corner ‘hinges’ to pull the tri-
angular to rectangular flap over towards the center of
the leaflet, and secures it to the leaf surface with silken
‘guy-wires’. A peaked roof is formed by the tight silk-
ing of a small cut perpendicular to one of the main
cuts. Type 2 shelters (‘one-cut folds’) are similar, but
entail only one cut in from the leaflet margin. Type 3
shelters (‘leaf folds’) have no cuts; caterpillars fold the
margin of some or all of a leaflet towards the center
and secure it to the surface with long guy wires. Type
4 shelters (‘two-leaf pockets’) consist of two leaflets
pulled together by silk strands to form a pocket.
80
ee 1st instar
ag (n=114)
% 50-4
255
(0) T 1 T
1 2 3 4
ad 3rd instar |
78 (n=1113) |
%
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
2nd instar
| (n=147)
a = T
2 3 4
5th instar
(n=203)
Ath instar
(n=109)
Shelter type
Fic. 2. Distribution of shelter types varies predictably across larval instars.
The style of shelter constructed varies predictably
over larval ontogeny (Fig. 2). First and second instar
caterpillars build type 1 shelters almost exclusively,
while third instar larvae construct mostly type 1, some
type 2, and rarely type 4 shelters. Fourth and fifth in-
stars never build type | shelters; instead, they con-
struct mostly shelter types 3 and 4.
Larger larvae build larger shelters, both across and
within shelter types. That is, each successive shelter
type is generally larger than the previous, and for a
given shelter type, shelter size is positively correlated
with larval size (Table 1). The relationship between
larval length and shelter size is relatively constant
across all five instars: the size of the shelter (approxi-
mated as (length + width)/2) ranges between 1.0 and
1.7 times the length of the larva (Table 1).
Laboratory-reared larvae allowed to change shelters
at will built a mean 5.1 shelters (40.24 SE: N = 15) over
five instars. Larvae generally constructed a new shelter
one to several days after molting within the previous
shelter (mean + SE = 1.7 + 0.2 days after first molt; 1.6
+ 0.2 days after second molt; 2.8 + 0.3 days after third
molt; 4.0 + 0.8 days after fourth molt). Only three out of
60 observed molts took place outside of a shelter.
2) Location of shelters on the host plant
The height above the ground of leaves bearing first
instar shelters ranged from 0.25—1.25 m, with a mean
+ SE of 0.65 + 0.03 m. The distribution of shelters
across the 4 quadrants differed significantly from ran-
dom (X? = 109.86, 3 df, p < 0.001). Almost 50% of the
shelters were located in quadrant 2, and together
quadrants 1 and 2 (the apical half of the leaf) con-
tained three-quarters of all shelters. As larvae grow,
the distance between shelters increases. First, second,
and third instar shelters are generally located on the
same leaflet, while shelters constructed by fourth and
fifth instar larvae are often constructed on an adjacent
leaflet or leaf (Table 1).
DISCUSSION
Although larvae of many lepidopteran species fold, roll
or tie leaves into shelters, it is not uncommon for these
taxa to begin their larval lives with a radically different
feeding habit, such as leaf-mining or boring (Gaston et al.
1991). Presumably, this is related to the small size of
early-instar larvae. For example, Caloptilia serotinella
(Gracillariidae), the cherry leaf-roller, is a leaf-miner in
VOLUME 54, NUMBER 3
its early stages, while fourth and fifth instar larvae are
leaf-rollers (Fitzgerald & Clark 1994). In a range of other
taxa, hatchling and early instar larvae use a shelter made
by another species, construct a communal shelter or silk
canopy, or hide in a cranny, while late instar larvae build
their own shelters (Doerksen & Neunzig 1976, Damman
1987, Cappuccino 1993, Loeffler 1994).
Like E. clarus, larvae in several other diverse taxa
construct leaf shelters throughout larval life, and ex-
hibit ontogenetic changes in style of shelter construc-
tion. The golden-banded skipper, Rhabdoides (=Au-
tochton) cellus, builds shelters in a progression of
styles very similar to that of E. clarus (Clark 1936).
Shelters made by larvae of the skipper Staphylus hay-
hurstii are also similar to those of E. clarus, although
the shelters lack a notch in the second cut and thus do
not have a peaked roof (pers. obs). Larvae of the
pyralid moth Herpetogramma aeglealis sequentially
construct and inhabit shelters of three distinct types on
fronds of Christmas fern (Ruehlmann et al. 1988).
Early-stage Nephopterix celtidella (Pyralidae) larvae
web two leaves flatly together, while last-stage larvae
web a dead, curled leaf to the surface of a living leaf
(Doerksen & Neunzig 1976).
Ontogenetic changes in shelter size and style may
be due to the biological needs and/or physical capabil-
ities of the larva. Certainly, to be fully enclosed and
hidden from predators, larger larvae require larger
shelters, so that while a small folded section of leaflet
is sufficient to cover a first instar caterpillar, larger lar-
vae need the increased area that a folded leaflet or two
leaves silked together can provide. Perhaps, in addi-
tion, a fixed relationship between larval size and shel-
ter size is necessary to maintain a particular internal
microclimate, or to restrict access to predators.
Changes in shelter size and style may also reflect
changes in the physical abilities of the larvae. A 3-mm-
long E. clarus hatchling may not be able, even utilizing
the axial retraction forces of stretched silk (Fitzgerald
et al. 1991) to fold a large flap over itself or pull two
leaflets together. It can, however, cut a small flap of leaf
tissue and fashion it into a shelter. As the larva in-
creases in size, it is able to manipulate larger pieces of
leaves, and cutting eventually becomes unnecessary.
Indeed, by the time larvae reach the fifth instar, they
rarely cut leaves prior to constructing their shelters,
and either fold over the entire edge of a leaflet or join
two leaflets together with silk. Cuts made in leaves by
late instar larvae might be counter-productive as well as
unnecessary, as the weight of a fifth instar larva (~700
mg) could pull down or tear a leaf flap. Cutting leaf tis-
sue may also cause the release of volatile compounds
that could attract parasitoids (Turlings et al. 1995).
$1
The types of shelters built by each larval instar may
also reflect selection for speed and efficiency of con-
struction, as leaf-rolling or leaf-tying insects are gener-
ally palatable to natural enemies (Bernays & Cornelius
1989), and exposed larvae are much more likely to be
eaten than are sheltered ones (Damman 1987, Cap-
puccino 1993).
Little is known about the ontogenetic patterns of
shelter construction for most taxa that fold, tie, or roll
leaves to make a shelter. Some species, like E. clarus
and Herpetogramma aeglealis (Pyralidae) (Ruehlmann
et al. 1988), build very regular structures that change
predictably in size and style over larval ontogeny, while
others produce more variable shelters (pers. obs.). The
relationship between insect size and shelter size is also
likely to vary across taxa. Larvae that feed inside the
shelter may make relatively larger shelters than those
that venture out to eat, and those that retain frass in the
shelter may also make larger shelters than those that
eject their frass. Comparative studies of shelter-building
taxa will help to elucidate the relative importance of var-
ious factors, including larval physical ability, feeding and
defecation behavior, vulnerability to predators, leaf
toughness, and internal microclimate, that may be in-
volved in determining patterns of shelter construction.
The innate behavior patterns underlying the con-
struction of different shelter styles are also worthy of
further study. We have determined that the almost in-
variant size and shape of first instar shelters results from
a prescribed pattern of larval movements and behaviors,
in which larvae use their body length as a ‘ruler’, and lay
down a silk ‘blueprint’ on the leaf surface prior to initi-
ating cuts (Weiss et al. in prep.). We are currently inves-
tigating the behavior of E. clarus on leaves of different
sizes and morphologies to determine the degree of plas-
ticity in these seemingly hard-wired behaviors.
ACKNOWLEDGMENTS
We thank Mike Singer and Liz Bernays of the University of
Arizona, Tucson, Deane Bowers of the University of Col-
orado, Boulder, Jennifer Maupin of Georgetown University,
and an anonymous reviewer for helpful comments on the
manuscript. We are grateful to Stephanie Brooks, Sharon Ko-
marow, and Dhara Levers for capable help in the field. This
work was supported in part by USDA NRI grant # 97-35311-
5151 to MRW.
LITERATURE CITED
Bernays, E. A. & M. L. CorNneELtus. 1989. Generalist caterpillar
prey are more palatable than specialists for the generalist
predator Iridomyrmex humilis. Oecologia 79:427-430.
Cappuccino, N. 1993. Mutual use of leaf-shelters by lepidopteran
larvae on paper birch. Ecol. Entomol. 18:287—292.
CiarK, A. H. 1936. The Gold-banded skipper (Rhabdoides cellus).
Smithsonian Miscellaneous Collections, vol. 95 (Publication
3386); Washington, D.C.
NSS 00
me we ee
Ciark, A. H. & L. F. Ciark. 1951. The butterflies of Virginia.
Smithsonian Institution, Washington, D.C.
DamMan, H. 1987. Leaf quality and enemy avoidance by the larvae
of a pyralid moth. Ecology 68:88-97.
DOERKSEN, G. P. & H. H. Neunzic. 1976. Biology of some im-
mature Nephopterix in the Eastern United States (Lepi-
doptera: Pyralidae: Phycitinae). Ann. Entomol. Soc. Amer.
69:423-431.
FRAENKEL, G. & FALLIL, F. 1981. The spinning (stitching) behaviour
of the rice leaf folder, Cnaphalocrocis medinalis. Ent. Exp. &
Appl. 29:138-146.
FITZGERALD, T. D. & K. L. CiarK. 1994. Analysis of leaf-rolling be-
havior of Caloptilia serotinella (Lepidoptera: Gracillariidae). J.
Insect Behav. 7:859-872.
FITZGERALD, T. D., K. L. CLARK, R. VANDERPOOL & C. PHILLIPS.
1991. Leaf shelter-building caterpillars harness forces gener-
ated by axial retraction of stretched and wetted silk. J. Insect
Behav. 4:21-32.
Gaston, K. J., D. REAVEY & G. R. VALLADARES. 1991. Changes in
feeding habit as caterpillars grow. Ecol. Entomol. 16:339-334.
HENSON, W. R. 1958. Some ecological implications of the leaf-
rolling habit in Compsolechia niveopulvella Chamb. Can. J. of
Zool. 36:809-8158.
LOEFFLER, C. C. 1994. Natural history of leaf-folding caterpillars
Dichomeris spp. (Gelechiidae), on goldenrods and asters.
J. New York Entomol. Soc. 102:405-428.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Moss, A. M. 1949. Biological notes of some Hesperiidae of Para and
the Amazon. Acta Zool. Lilloana 7:27—79.
RUEHLMANN, T. E., R. W. MaTrHEews & J. R. MatrHews. 1988.
Roles for structural and temporal shelter-changing by fern-
feeding lepidopteran larvae. Oecologia 75:228-232.
SAGERS, C. L. 1992. Manipulation of host plant quality: herbivores
keep leaves in the dark. Funct. Ecol. 6:741-743.
SANDBERG, S. L. & M. R. BERENBAUM. 1989. Leaf-tying by tortricid
larvae as an adaptation for feeding on phototoxic Hypericum
perforatum. J. Chem. Ecol, 15:875-885.
SCOBLE, M. J. 1992. The Lepidoptera: form, function and diversity.
Oxford University Press, Oxford, 404 pp.
Scort, J. A. 1986. The butterflies of North America; a natural his-
tory and field guide. Stanford University Press, Stanford, Cali-
fornia.
SCUDDER, S. H. 1889. Butterflies of the Eastern United States and
Canada. 3 vol. Cambridge, Massachusetts.
TURLINGS, T. J. C., J. H. Loucurin, P. J. MCCALL, U.S. R. RosE, &
W. J. Lewis. 1995. How caterpillar-damaged plants protect
themselves by attracting parastic wasps. Proc. Natl. Acad. Sci.
USA 92:4169-4174.
Received for publication 7 September 1999; revised and accepted 2
October 2000.
Journal of the Lepidopterists’ Society
54(3), 2001, 83-87
REPRODUCTIVE OUTPUT AND EGG MATURATION
IN RELATION TO MATE-AVOIDANCE IN MONANDROUS FEMALES
OF THE SMALL COPPER, LYCAENA PHLAEAS (LYCAENIDAE)
MAMORU WATANABE AND MIYUKI NISHIMURA
Department of Biology, Faculty of Education, Mie University, Tsu, Mie 514-8507, Japan
ABSTRACT Females of the small copper, Lycaena phlaeas, were captured in the field and dissected to investigate their mating frequency
and reproductive output. Spermatophore counts showed that most females were monandrous. A single spermatophore usually occupied about
half of the bursa copulatrix, though the size of the spermatophore decreased with the female’s age. Young females had 250 immature eggs in
their ovaries and laid an estimated 150 eggs throughout the course of their lives. Mate avoidance behavior was frequently observed in both
mated and virgin females. In laboratory experiments, virgin females contained a few mature eggs immediately after eclosion, and the number
of mature eggs gradually increased with age. Ovaries of females accepting copulation contained significantly larger numbers of mature eggs than
ovaries of females avoiding mating. Copula duration was about 16 min (at 30°C) irrespective of male age. Egg maturation in the ovaries is thus
seen as important in the mating behavior of such monandrous species.
Additional key words: _ bursa copulatrix, egg load, Lycaena phlaeas, mate-avoidance, monogamous, spermatophore.
In butterflies, ejaculates from males during copula-
tion are used in female egg production and/or somatic
maintenance. Females may, therefore, benefit from
mating more than once (Boggs 1981, Boggs & Gilbert
1979, Watanabe, 1988). One potential benefit of re-
peated matings for the female butterfly is the substan-
tial increase of sperm supply (Lederhouse 1981). As a
rule, however, males transfer a number of sperm in ex-
cess of that needed to inseminate all eggs at a single
mating (Watanabe et al. 1998). Consequently, in cases
in which a spermatophore is used only as a reservoir of
sperm to fertilize eggs, females should benefit from
mating only once. Re-mating may decrease the time
available to females for egg-laying and foraging activi-
ties, and may increase the risk of predation while in
copula. In the evolution of such species, therefore,
natural selection should favor females who avoid males
after mating. The mate refusal posture displayed by
unreceptive females has been reported in many but-
terfly species (Shapiro 1970, Watanabe et al. 1997).
The small copper, Lycaena phlaeas (Linnaeus), is a
multi-voltine butterfly that flies from spring to autumn
in central Japan. Adults inhabit sunny areas such as for-
est margins, clearings, open fields with low vegetation,
banks and roadsides (Suzuki 1976). Suzuki (1978) re-
ported mate-avoidance behavior in females and sug-
gested that females generally mate only once. However,
there are few data on spermatophore counts in the
bursa copulatrix of wild females, and few studies have
been conducted on oviposition behavior, nor have any
data on lifetime reproductive schedules been gathered.
In this paper, we clarify the mating frequency and
fecundity of L. phlaeas in the field. Lifetime egg pro-
duction and reproductive success are discussed. Since
female monandrous butterflies may solicit courtship
before copulation and actively avoid males after mat-
ing in order to allow time for oviposition (Wiklund
1982), we examined the relationship between mate-
avoidance behavior and fecundity in the laboratory.
MATERIALS AND METHODS
The data presented in this paper were obtained pri-
marily from summer generations of the small copper, L.
phlaeas, in Shirouma in Nagano Prefecture, which is lo-
cated in a cool-temperate zone of Japan. The major
habitat consists of rice-fields and margins of deciduous
forests where adults may feed on nectar plants, and lar-
vae on host plants (primarily Rumex japonicus, Polygo-
naceae), during the summer. During the day, large
numbers of males fly near the ground to gather nectar
and search for mates. Chasing behavior between males
and females was sometimes observed over the course of
the summer. Females engaged in various activities such
as feeding, roosting, flying, copulating and ovipositing,
were collected from late July to early August of both
1995 and 1996. To examine mating frequency of fe-
males flying in natural populations, sampling was done
on windless, sunny days (a total of 13 days). When fe-
males were captured, their abdomens were amputated
and immersed in a solution of 50% ethyl alcohol, and
their wing condition and forewing length were
recorded. We classified individuals into 5 age groups, on
the basis of the degree of wing damage, using a ranking
from O to IV, following Watanabe and Andos method
(1993). The five age groups were as follows: O, newly
emerged females with wings having lustrous scales and
no visible damage; I, fine tears and fewer lustrous
scales; II, tears and frayed scales; III, notched tears and
frayed scales; and IV, broken or extensive tears and
frayed scales. All females were examined for the num-
ber of spermatophores in their bursa copulatrix and the
number of eggs stored in their ovaries.
84
The individuals of L. phlaeas used in the mating ex-
periments were the offspring of females collected in
the wild. They laid eggs in cages, and the hatching lar-
vae were reared on the host plant, R. japonicus, in
small chambers at 25°C, with 18 hours of light per day.
A newly emerged virgin female was placed in a mating
cage (30 x 40 x 45 cm) along with 10 virgin males (less
than 10 days old) to allow matings. The cage was set in
a greenhouse for two hours around noon (at ca 30°C).
In the present study, 451 females emerged in the lab-
oratory. Their average forewing length was 16.5 + 0.6
mm (SD), which is not significantly different from that
of the captured wild females (16.3 + 0.9 mm, U-test, Z
= (0.825, n.s.). Out of the 451 females, 147 were ran-
domly selected for mating experiments.
The mating pairs were kept in small cages (24 x 30
x 38 cm) until they separated. The duration of copula-
tion was recorded. After copulation terminated, fe-
males were dissected to examine the spermatophore
and to count the number of eggs in ovaries. As a con-
trol, virgin females of various ages in stock culture
were also dissected.
In the present study, many females introduced into
the cages avoided or refused to copulate with males
displaying courtship behavior, although the males re-
peatedly approached females in the course of two
hours. These females were taken out of the cage and
supplied with sugar solution in another small cage at ca.
25°C. On the following day, they were then placed back
in the mating cage. In this way, the copulation trials for
virgin females were repeated every day until the female
engaged in copulation.
Since both the spermatophore and the bursa copula-
trix are oval, their respective volumes were calculated as
an ellipsoid. Although eggs in the ovaries could be classi-
fied into three groups (mature, submature, immature),
those in the oviducts were primarily of the mature type.
These oviduct eggs were large, pale green and suffi-
ciently well-formed (with a semi-spherical shape and
pronounced ribbing) to be ready for oviposition. The
vitellarium of the ovaries contained mainly submature
eggs; these were large but more lightly colored than the
mature eggs. Immature eggs, which included oocytes,
were found in the terminal filament, the germarium and
part of the vitellarium of the ovarioles. Eggs in the ovar-
ioles decreased in size toward the tip of the terminal fil-
ament filled with oocytes. Because oocytes do not con-
tain yolk, they appear white, and we were able to count
them using tweezers and a light microscope (40x).
RESULTS
Over the course of the two-year study period, we ex-
amined a total of 99 wild females from the study area.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
TABLE 1. The frequency of spermatophore distribution in the
bursa copulatrix of female Lycaena phaeas caught in the wild for 5
age classes (0 ~ IV) in the summer generation of 1995 and 1996.
Number of Age class
Years spermatophores
present 0 I Il II IV
0 5 2 0 0 0
1995 1 29 16 i 1 0
2 2 2 0 0 0
0 2 1 0 0 0
1996 1 3 9 6 8 4
2) 0 0 IL 0 1
In 1995, more than half of the females captured were
young (ages O and I), and no females of the oldest age
class (age IV) were captured (Table 1). Five of 36 age-
O and 2 of 20 age-I females were virgins, and 2 addi-
tional virgin females were captured at age I. In 1996, 3
age-I females were virgin. In both years, there were a
few females that contained both an intact full-sized
spermatophore and a spermatophore fragment, thus
showing evidence of having mated twice as shown in
Table 1. No females contained mating plugs blocking
their genital openings.
In mated females, the spermatophore has been ejac-
ulated into the bursa copulatrix and is filled with white
secretion. The sperm sac is an elongated cone which oc-
cupies the bursal duct and has its opening at the end of
the duct near the seminal duct. There is no appendix
bursae in the female reproductive system.
In age-O females, the average volume of the sper-
matophore was about 0.05 mm, which occupied about
31% of the volume of the bursa copulatrix (Fig. 1).
There was no significant difference in the size of the
spermatophores among females of age O to III (F =
1.610, n.s.). Among females of these four age classes,
the spermatophore occupied about 40% of the bursa
copulatrix. In age-IV females, the spermatophore size
significantly decreased (p < 0.01), compared with the
spermatophore size of age-I (U = 14), age-II (U = 6) and
age-III (U = 3) females, but still occupied about 45% of
the bursa copulatrix.
The mean number of eggs (+SE) in wild-caught vir-
gin females of age O was 41.7 + 13.3 (n = 7), 19.14 3.8
(n = 7) and 242.1 + 39.3 (n = 7) for mature, submature
and immature eggs, respectively. The mated females
of age O contained means (+SE) of 261 + 15 immature
eggs (n = 29), which number was similar to that for vir-
gin females. Therefore, the potential fecundity of L.
phlaeas was estimated at about 300. Figure 2 shows
the lifetime changes in the number of immature eggs
in monandrous females. The number of immature
eggs decreased with age. Since there were less than 20
VOLUME 54, NUMBER 3
ied by
=
Proportion
p
spermatophore
)
iSal
a
eee See ee ae
r
bursa copulatrix > 5
27 |
| |-spermatophore
10
occu
S
ee)
1 f
B)
20
0.1-
Volume of bursa copulatrix (mm
Age class
Fic. 1. Change in the volume of spermatophore in the bursa
copulatrix and its proportion in wild monandrous females of Ly-
caena phlaeas for 5 age classes (O, I, II, If and IV). Each bar repre-
sents SE. The numbers above the bars show sample size.
mature eggs in the ovaries of females of age IV, the de-
crease in immature eggs resulted from resorption or
oviposition. However, no fused eggs were observed in
the ovaries, suggesting that few eggs were consumed
for somatic maintenance. In such cases, in which there
is no addition of immature eggs during the adult stage,
the decreasing number of immature eggs in the
ovaries is thought to be due to oviposition. A monoga-
mous female of age IV may thus have laid 150 eggs.
In the mating cages, all L. phlaeas males were seen
perching, and would presumably mate with the females.
We observed a total of 605 unsuccessful courtships,
which can be described in terms of five distinct behav-
ioral events. After approaching a female, the male tried
to make initial contact. In 43% of unsuccessful
courtships, females remained perched and ignored ap-
proaching males. Walking or flying away from males was
85
300
an
rt)
iT)
oe
2
3 200°
S
E
£
So
°o
nw 100-
)
2
E
5
Zz.
ot
l ! ! !
0 I I sit IV
Age class
Fic. 2. Changes in the number of immature eggs of age O, I,
IL, IM and IV wild monandrous females of Lycaena phlaeas in 1995
(circle) and 1996 (triangle). Each bar represents SE. The numbers
above the symbols show sample size.
seen in 29% and 14% of approaches, respectively. In
some cases, females tumed face-to-face with the male
(9%) or fluttered (4%) until the male flew away. The lat-
ter two behaviors were considered to constitute mate-re-
fusal behavior rather than mate-avoidance, although no
females showed the pierid mate-refusal posture in which
the wings are spread and the abdomen elevated
(Shapiro, 1970). There was no significant difference be-
tween the forewing length of males failing to copulate
(wing length = 15.2 + 0.2 [SE] mm, n = 24) and that of
males succeeding in copulation (wing length = 15.6 + 0.2
mm [SE], n = 9) (U-test, Z = 0.788, n.s.).
Out of 77 virgin females aged 2-days at the first trial,
only 22 engaged in copulation (28.6%). The other fe-
males were re-introduced to the mating experiment on
the following day and 35.0% of them (7/20) copulated.
In the third trial, 28.6% of the virgin females (2/7) cop-
ulated. The remaining virgin females did not copulate
with males.
The copulation duration was about 16-17 min, re-
gardless of the number of previous trials (Table 2).
When the copulation terminated, most females were
TABLE 2, The number of eggs loaded in accepting and avoiding females during male courtship behavior (mean + SE).
Females of 1st trial Accepting
Number of females dissected 22
Copula duration (min) 15’ 33” + 41”
Number of mature eggs 62.6 + 8.9
Number of submature eggs 39.1 + 3.8
Number of immature eggs 254.7 + 8.6
Females of 2nd and 3rd trials Accepting
Number of females dissected 9
Copula duration (min) 16’ 45” + 1’ 22”
Number of mature eggs 60.9 + 12.8
Number of submature eggs 32.8 + 5.0
Number of immature eggs 297.3 + 12.3
Avoiding U-test
6
2.8 + 2.6 Z=3.191,p<0.01
20.8 + 4.4 Z = 2.715, p < 0.01
228.7 + 20.1 LZi= ie 2255 nes:
Avoiding U-test
6
30.3 + 11.9 U = 8, 0.05 > p > 0.01
21.5 + 5.0 U = 13.5, nis.
24).7 + 7.8 U= 18, ns.
|
|
|
|
|
|
86
dissected, and their bursa copulatrix and the number of
eggs in their ovaries were examined. Every female had a
single spermatophore in the bursa copulatrix, though it
was not completely solidified in any of the females. The
average volume of the bursa copulatrix among mated fe-
males was 0.195 + 0.017 [SE] mm® (n = 25), which was
significantly larger than that of wild females of age O (U-
test, Z = 3.182, p < 0.01). The average volume of the
spermatophore was 0.145 + 0.015 [SE] mm? (n = 20),
which occupied 74% of the bursa copulatrix compared
with 40% in wild females. Spermatophore volume was
also significantly larger than that of wild-caught females
of age O (U-test, U = 37, p < 0.01).
After the trial, a number of virgin females who had
consistently avoided mating were also examined to de-
termine the number of eggs in the ovaries. As shown in
Table 2, females accepting mating at the first trial con-
tained significantly more mature and submature eggs
than those who avoided mating at the first trial. The
number of immature eggs did not differ among female
mating outcomes. At the second and the third trials, fe-
males with immature eggs were older and must there-
fore have further developed their eggs in the ovaries.
However, mated females loaded about 60 mature eggs,
which was significantly greater than that of the females
avoiding mating (30 mature eggs) in Table 2. There was
no significant difference in the numbers of submature
or immature eggs between females accepting and
avoiding mating in either the second or third trials.
Figure 3 shows the change in the number of mature
eggs in females reared in the laboratory. Very few ma-
ture eggs were found in the ovaries of newly emerged
females (0-day-old). It follows that maturation of eggs
occurs with aging. About 50 mature eggs had accumu-
lated one week after emergence. Figure 3 also shows
that the number of mature eggs in mating females was
above the mean number of mature eggs of virgin fe-
males, and that the number of females avoiding mating
was below it. An increase in the number of mature eggs
thus appears to facilitate female willingness to mate.
DISCUSSION
The mating frequency of L. phlaeas females was ex-
amined in terms of the number of spermatophores in
the bursa copulatrix, because each mating generally re-
sults in the deposition of a single spermatophore. It is
well known that the number of spermatophores in sey-
eral butterfly species increases with female age (Leder-
house 1981, Watanabe & Nozato 1986, Watanabe &
Ando 1993). However, there is little information on the
mating frequency of female lycaenid butterflies (Burns
1968, Suzuki 1978). In the present study, most females
(94%) of the small copper, L. phlacas, had a single
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
200 -
a: virgin
e : mated
©: avoiding males
ol
fo)
I
e
Number of mature eggs
an =
fo) oO
0 5 10 15
Days after emergence
Fic. 3. _ Daily change in the number of mature eggs of virgin
Lycaena phlaeas females (squares, + SD) and the number of mature
eggs of females accepting copulation (closed circles) and of those
avoiding males (open circles) in laboratory experiments.
spermatophore in the bursa copulatrix irrespective of
their age class. We found a few polyandrous females
(6%) that had two spermatophores, one of normal size
lying near to the site of sperm migration in the bursa
copulatrix and the other being very small or consisting
of fragments. Although we did not find any signs that
the spermatophore had been absorbed by the female,
the spermatophore just after copulation of laboratory-
reared young females was larger than that of females
of age class O, suggesting that females quickly ab-
sorbed the spermatophore. This means that wild-
caught females had retained the remnants of the
spermatophore. Therefore, the fragments of sper-
matophore must be derived from a small sper-
matophore, which might be transferred by old males
(e.g., Lederhouse et al. 1989) or by recently mated
males (Watanabe et al. 1997). When the first sper-
matophore is too small for females to avoid re-copula-
tion, females must be exceptionally polyandrous.
Female reproductive success will depend on the
amount of three potential resources available to fe-
male somatic maintenance or offspring. One is the ni-
trogenous reserves accumulated during their own lar-
val feeding and stored in the fat body, which is usually
depleted with egg development (e.g., Dunlap-Pianka
et al. 1977). Another is nectar feeding during the adult
stage (e.g., Boggs 1986, Watanabe 1992), though the
nectar contains little nitrogen. The third resource is
the contributions of the male ejaculate during copula-
tion (e.g., Boggs & Watt 1981). Boggs and Gilbert (1979)
showed that the ejaculates were used for egg develop-
ment by females. In the present study, however, fe-
males seemed not to use the ejaculates to increase
their fecundity. Bissoondath and Wiklund (1995) stated
that both relative ejaculate mass and protein content
VOLUME 54, NUMBER 3
in the spermatophore are low in monandrous species.
In this study, the small copper females had few ma-
ture eggs following eclosion, and they used their own
energy or nectar to develop eggs without ejaculates
from males. The lifetime reproductive output was esti-
mated at about 150 eggs in the present study. Females
feed on nectar as an energy source. On the basis of our
experiments in which virgin females were kept in a
flight cage and allowed to feed freely on nectar, such
females live more than 3 weeks without mating. Boggs
(1986) reported that females of the nymphalid butter-
fly, Speyeria mormonia, which is monandrous, sur-
vived for an average of 19 days in captivity, laying eggs
for 14 days on average.
The present study showed that the ratio of sper-
matophore volume to the volume of the bursa copulatrix
was stable, regardless of the spermatophore size. Sug-
awara (1979) showed that a certain volume of sper-
matophore is perceived by the stretch receptors which
stimulate the mate refusal posture of the female white
butterfly, Pieris rapae. The stretch receptors in the bursa
copulatrix may also operate in the small copper to induce
mate-avoidance behavior over the course of the insect’s
life, and thereby maintain female monandry.
The present study suggests that the number of ma-
ture eggs determines the mating behavior of the fe-
male butterflies. This means that there is a period of
sexual immaturity and avoidance or refusal of males af-
ter eclosion. Therefore, females of this species have a
pre-mating adult period, during which they mature
initial eggs. One possible explanation for this phenom-
enon is that it allows females to be more particular in
their choice of mate, since the small copper is a perch-
ing species, in which males spend part of the day sit-
ting on some object waiting for passing females who
are then pursued and courted. Such behavior may be
particularly advantageous in a monandrous species.
Wiklund (1977) showed that female monogamy in the
pierid butterfly, Leptidea sinapis, is maintained by the
females simply remaining quiescent during courtship,
without any kind of mate-refusal posture. However,
there is no quantitative information on the diurnal ac-
tivity of females of the small copper or on their choos-
ing of mates in the field. A detailed study of preferen-
tial mating will be required for the monandrous small
copper.
ACKNOWLEDGMENTS
We thank Carol Boggs, Deane Bowers and an anonymous referee
for their helpful comments on the manuscript. Thanks are also due
to Roger Kitching for his kind advice. We are grateful to Messrs. T.
Higashi, Y. Nakanishi, T. Ito, T. Imoto, M. Hirota, M. Bon’no, and A.
Hachisuka for their assistance in the field.
LITERATURE CITED
BissOONDATH, C. J. & C. WIKLUND. 1995. Protein content of sper-
matophores in relation to monandry/polyandry in butterflies.
Behay. Ecol. Sociobiol. 37:365-371.
Boccs, C. L. 1981. Nutritional and life-history determinants of re-
source allocation in holometabolous insects. Am. Nat.
117:692-709.
. 1986. Reproductive strategies of female butterflies: varia-
tion in and constraints on fecundity. Ecol. Entomol. 11:7-15.
Boccs, C. L. & L. E. GILBert. 1979. Male contribution to egg pro-
duction in butterflies: evidence for transfer of nutrients at mat-
ing. Science 206:83-84.,
Boccs, C. L. & W. B. Warr. 1981. Population structure of pierid
butterflies. IV. Genetic and physiological investment in off-
spring by male Colias. Oecologia 50:320-324.
Burns, J. M. 1968. Mating frequency in natural populations of skip-
pers and butterflies as determined by spermatophore counts.
Proc. Natl. Acad. Sci. 61:852-859.
DUNLAP-PIANKA, H., C. L. Boccs & L. E. GILBERT. 1977. Ovarian
dynamics in heliconiine butterflies: Programmed senescence
versus eternal youth. Science 197:487-490.
LEDERHOUSE, R. C. 1981. The effect of female mating frequency on
egg fertility in the black swallowtail, Papilio polyxenes asterius
(Papilionidae). J. Lepid. Soc. 35:266-277.
LEDERHOUSE , R. C., M. P. AyRES & J. } M. ScriBER. 1989. Evaluation
of spermatophore counts in studying mating systems of Lepi-
doptera. J. Lepid. Soc. 43:93-101.
Sapiro, A. M. 1970. The role of sexual behavior in density-related
dispersal of pierid butterflies. Am. Nat. 104:367—-372.
SucawaRA, T. 1979. Stretch reception in the bursa copulatrix of the
butterfly, Pieris rapae crucivora, and its role in behavior. J.
Comp. Physiol. 130:191-199.
SUZUKI, Y. 1976. So-called territorial behaviour of the small copper,
Lycaena phlaeas daimio Seitz (Lepidoptera, Lycaenidae). Kon-
tyu 44:193-204.
. 1978. Mate-avoiding behaviour in females of the small cop-
per, Lycaena phlaeas daimio Seitz (Lepidoptera: Lycaenidae).
Trans. Lepid. Soc. Jpn 29:129-138.
WaTANABE, M. 1988. Multiple matings increase the fecundity of the
yellow swallowtail butterfly, Papilio xuthus L., in the summer
generations. J. Insect Behav. 1:17-30.
. 1992. Egg maturation in laboratory-reared females of the
srelllonieil butterfly, Papilio xuthus L. (Lepidoptera: Papilion-
idae), feeding on different concentration solutions of sugar.
Zool. Sci. 9:133-141.
WaTANABE , M. & S. ANDO. 1993. Influence of mating frequency on
lifetime fecundity in wild females of small white Pieris rapae
(Lepidoptera: Pieridae). ). Jpn. J. Entomol. 61:691—696.
WATANABE , M., Y. NAKANISHI & M. BON’NO. 1997. Prolonged copu-
lation and spermatophore size ejaculated in the sulfur butterfly,
Colias erate (Lepidoptera: Pieridae) under selective harassments
of mated pairs by conspecific males. J. Ethol. 15:45-54.
WATANABE , M. & K. NozatTo. 1986. Fecundity of the yellow swal-
lowtail butterflies, Papilio xuthus and P. machaon hippocrate S,
in a wild environment. Zool. Sci. 3:509-516.
WATANABE , M., C. WIKLUND & M. Bon’no. 1998. Ejaculation tim-
ing of eupyrene and apyrene sperm in the cabbage white but-
terfly Pieris rapae (Lepidoptera: Pieridae) during copulation.
Entomol. Sci. 1:15-19.
WIKLUND, C. 1977. Courtship behaviour in relation to female monog-
amy in Leptidea sinapis (Lepidoptera). Oikos 29:275-283.
. 1982. Behavioural shift from Ce ee solicitation to mate
avoidance in female ringlet butterflies (Aphantopus hyperan-
thus) after copulation. Anim. Behay. 30:790-793.
Received for publication 29 December 1998: revised and accepted 6
June 2000.
Journal of the Lepidopterists’ Society
54(3), 2001, 88-90
DESCRIPTIONS OF THE IMMATURE STAGES AND OVIPOSTION BEHAVIOR OF
PYRRHOGYRA OTOLAIS (NYMPHALIDAE)
HAROLD F. GREENEY
Yanayacu Biological Station and Center for Creative Studies, Cosanga, Ecuador
c/o Carrion No. 21-01 y Juan Leon Mera, Quito, Ecuador
AND
NICOLE M. GERARDO
Department of Zoology, University of Texas, Austin, Texas 78712, USA
ABSTRACT. The morphology and behavior of the immature stages of Pyrrhogyra otolais (Nymphalidae) are described. The early stages
are compared with those of Pyrrhogyra crameri. An unidentified species of Serjania (Sapindaceae) is the larval host plant in Ecuador. The ovipo-
sition behavior and habits of the adults are discussed.
Additional key words:
The entirely neotropical genus Pyrrhogyra Hiibner
encompasses between nine and twelve species (D’Abr-
era 1987) distributed from Mexico through Central and
South America, with the highest diversity in northern
South America. At least four species occur in our study
area. Our current understanding of species relationships
within this genus is poor (D’Abrera 1987) and larval de-
scriptions and hostplant relationships will provide useful
information for correctly separating species. Adults are
fast flyers and are frequently encountered in forest light
gaps where they feed on dung, fruit (DeVries 1987) and
carrion (pers. obs.). Known host plants are all within the
family Sapindaceae (DeVries 1987) and the larvae show
close behavioral and morphological affinities to such
genera as Catonephele Hiibner, Nessaea Hiibner, Cal-
licore Hiibner, Diaethria Billberg, Temenis Hiibner, Eu-
nica Hiibner, and Epiphile Doubleday (DeVries 1987).
Pyrrhogyra otolais Bates is distributed from Mexico
to Peru, and seems confined to wet forests of lower ele-
vations. Despite this widespread butterfly’ formal de-
scription nearly 150 years ago, its early stages and host-
plant relationships have remained undescribed. This
study presents observations on hostplant use, larval mor-
phology and behavior, as well as oviposition behavior.
MATERIALS AND METHODS
All observations and rearings were conducted at the
La Selva Biological Station located in the Sucumbios
providence of eastern Ecuador. The station is located ap-
proximately 75 km east-south-east of Coca at an eleva-
tion of approximately 250 m. All studies were conducted
adjacent to the oxbow lake Garza Cocha. The forest in
this area is predominantly intact, though some areas
have been clear-cut for coffee and manioc plantations.
On 14 November 1997, at 1330, two female P. oto-
lais were observed to oviposit on a small seedling of an
unidentified species of Serjania Mill. (Sapindaceae).
Ecuador, larva, pupa, Serjania, Sapindaceae.
Six eggs and a second instar were collected by cutting
off the leaves to which they were attached. Larvae
were reared in individual plastic cups. Fresh leaves
were added and frass was removed semi-daily. Subse-
quently, larvae in various stadia were found and reared
in a similar fashion. Vouchers of each stadium were
preserved. Seven larvae were reared to adults. All head
capsules, pupal exuviae, and vouchers are in the per-
sonal collection of the senior author.
RESULTS
Larval host plant. Serjania sp. (Sapindaceae) is the
larval host plant of P. otolais at our study site in eastern
Ecuador. The strikingly colored, new, red leaves of this
plant are easily seen in light-gap areas of the forest and
in the forest understory. New leaves are present from
November to January. The fact that adult P. otolais are
present for most of the year in this area and can be quite
common, suggests that adults are fairly long-lived.
Oviposition behavior. Upon locating a potentially
suitable oviposition plant, the two observed females be-
gan flying rapidly in tight circles around the plant. Dur-
ing this period they landed on leaves of any plant in the
vicinity. After several minutes they began landing more
and more frequently on the host plant. While on the
plant they walked about for several seconds rapidly tap-
ping their forelegs on the leaf. After a single egg was laid,
they repeated the rapid flight and leaf-tapping sequence
for about 30 seconds before laying another egg. Eggs
were always laid singly on the fresh red leaves of new
shoots, but multiple eggs were eventually placed on the
same leaf. Most commonly, eggs were placed dorsally
along the mid-rib of the leaf, but were occasionally
placed at the base of the petiole or on the stem nearby.
Egg. (n = 20+, Fig. la). Pale-yellow to white, darken-
ing slowly after being laid, shape a truncated cone with
strong longitudinal ridges and many closely spaced cross
VOLUME 54, NUMBER 3
Fic. 1. Egg and larval head capsules of P. otolais. (a) egg: not
drawn to scale (b) Second instar (c) Fourth instar (d) Pupa: not
drawn to scale.
ribs, a slight constriction near the top gives them almost
a bell shape. First instars hatch by flipping open the en-
tire top of the egg like a lid. Larvae were never observed
to eat the egg shell upon emergence.
Larva. First instar (n = 6). Head dark brown and de-
void of scoli; body light yellow with dark setae. Second in-
star (n = 9, Fig. 1b). Head black with dorso-lateral scoli;
body same color as first instar, all segments with pairs of
short, rounded scoli dorsally, all with bulbous tips, brown
at base and white at apex; A7 and A8 with middorsal scoli
formed by rosettes of 3 and 5 spines respectively. Early
third instar (n = 7). Head black with dorso-lateral scoli
curving backwards, each scolus with several irregularly
placed chalazae along the shaft and ending in a rosette of
five spines, shaft of each scolus white immediately below
apical rosette; body dull yellow, T3—A6 dorsally dark
greenish-gray color with fine white patteming; Al-A6
with a pair of small white spots dorsally on either side of
midline, all segments with dorsal, bifid scoli on either
side of midline forming a row along the body, similar
rows of single scoli laterally and ventro-laterally; T1 with
single middorsal scolus; T2 with three-pronged middor-
sal scolus; T3 with five-pronged middorsal scolus; A2-A8
each with single, bifid, middorsal scolus; A9 with mid-
dorsal rosette of four scoli; A7 with middorsal, three-
branched scolus in addition to the bifid scolus; A8 with
additional five branched scolus; all scoli black and
sparsely covered with small chalazae. Late third instar (n
89
= 7+). Similar to early third instar but with orange bod-
ies, dorsal portions of T3—A7 pale yellow with dark brown
markings. Fourth instar. (n = 10+, Fig. 1c) Head as de-
scribed for third instars; body as above with the addition
of small, single scoli near the base of each proleg. Fifth
instar (n = 17). Head bright orange except for the scoli
and all spines, no white patch below the rosette at the tip
of the scoli; body differs from fourth instar in being more
brightly orange and the dorsal areas of segments T3-A7
bright yellow with dark brown or black markings.
Pupa. (n = 11, Fig. ld) Mostly lime-green with
brown markings along the ridges, cremaster dark
brown, strong thoracic keel coming to an abrupt peak
around the mesothorax, strong ridge along the dorsal
edge of the wing pads extending onto the head and
eyes, another strong ridge across the dorsum of the
second abdominal segment. All individuals that pu-
pated in the laboratory did so on the side of the con-
tainer and were attached at nearly right angles to the
sides of the container but curved downward. Only one
pupa was found in the field, and this was attached to
the near-horizontal surface of the hostplant leaf.
Larval behavior. Typical of other nymphalid genera,
first and second instar larvae rest on “frass chains” built
from the edge of the leaf (DeVries 1987, Otero & Aiello
1996). Later instars rest along the midrib of the leaf
when not feeding. They rest with the head oriented to-
wards the apex of the leaf, and angled so that the scoli
project forwards. When disturbed, larvae rear back onto
their prolegs and thrash wildly about, occasionally rais-
ing their terminal abdominal segments simultaneously.
DISCUSSON
As noted for other species of Pyrrhogyra (DeVries
1987), the early stages of P. otolais show affinities to the
early stages of other sapindaceous feeders such as Teme-
nis Hiibner and Epiphile Doubleday (Nymphalidae).
First, second, and early third instars are almost identical
to those of P. crameri Aurivillius, which can be found
during the same time of year in this area. Pyrrhogyra
crameri larvae, however, feed on Paullinia Linn. (Sapin-
daceae). The second and third instars can be separated
from those of P. otolais by the presence of only three
spines in the middorsal rosette of segment AS. The be-
havior of these two species while resting as larvae on
their host plant and as adults is very similar. Adults of
both species are seen frequently patrolling light gaps,
usually perching several meters from the ground.
ACKNOWLEDGMENTS
We thank R. Hill and S. Wheeler for their help in collecting larval
host plants, and R. Raguso and M. S. Singer for critical comments on
the manuscript. H. Greeney gives a special thanks to Bob and Katt
Azar for their encouragement and financial support. N. Gerardo
90
thanks the Thomas Watson Foundation for supporting her studies
abroad. We thank the PBNHS for their continued support.
LITERATURE CITED
D’AsrERA, B. 1987. Butterflies of the Neotropical Region. Part III.
Brassolidae, Acraeidae, & Nymphalidae (partim.). Hill House,
Victoria, Australia.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
DEVRIES, P. J. 1987. The butterflies of Costa Rica and their natural
history, Vol. 1, Papilionidae, Pieridae, Nymphalidae. Princeton
Univ. Press, Princeton, New Jersey.
OTERO, L. D. & A. AIELLO. 1996. Descriptions of the immature stages
of Adelpha alala (Nymphalidae). J. Lepid. Soc. 50:329-336.
Received for publication 6 April 1999: revised and accepted 25 Oc-
tober 1999.
Journal of the Lepidopterists’ Society
54(3), 2001, 91-95
EFFECTS OF ADULT FEEDING AND TEMPERATURE REGIME ON
FECUNDITY AND LONGEVITY IN THE BUTTERFLY
LYCAENA HIPPOTHOE (LYCAENIDAE)
KLAUS FISCHER AND KONRAD FIEDLER
Department of Animal Ecology I, University of Bayreuth, D-95440 Bayreuth, Germany
ABSTRACT. When fed highly concentrated sucrose solution, adult females of the Purple-Edged Copper butterfly Lycaena hippothoe L.
laid significantly more eggs (mean = 464) than those individuals given water only (mean = 65). Longevity was also three to five times greater,
whereas hatching rate of the eggs was not affected by the mother’s nutrient intake. Stored resources acquired during the larval stage supported
realization of only 14% of the fecundity of the fed females. Hence, L. hippothoe butterflies depend far more on adult-derived resources than
other nectar-feeding butterflies for which comparable data exist. These findings may be important for the population dynamics of the species,
as reduced availability of nectar sources presumably constrains realized fecundity.
Additional key words: Adult diet, reproduction, nutritional ecology, population dynamics.
Oogenesis in insects is typically a nutrient-limited
process, triggered only if sufficient nourishment is avail-
able (Wheeler 1996). The required resources can be ac-
quired during the larval or adult stage (or both), de-
pending on the insects’ life cycle (Boggs 1997a). In the
holometabolous Lepidoptera, resources for reproduc-
tive investment may stem from the herbivorous larval
stage or from the adult stage, during which they collect
liquid food such as nectar (Wheeler 1996). Although it
is generally thought that butterflies and moths usually
rely more on reserves accumulated during the larval
stage to supply egg production (Wheeler 1996), there is
broad variation with regard to the importance of adult
feeding (Hill 1989, Murphy et al. 1983). Many adult
moths do not feed at all and rely completely on reserves
accumulated during the larval stages (“capital breeders”
sensu Tammaru & Haukioja 1996). At the other ex-
treme, female Heliconius butterflies collect protein-rich
pollen which supports the laying of eggs through the
adult life span of several months (Gilbert 1972, Dunlap-
Pianka et al. 1977). Such species can be referred to as
“income breeders” (Tammaru & Haukioja 1996), for
which successful reproduction is essentially mediated
by resources available to the adults.
In many temperate-zone nectar-feeding butterflies
(cf. Boggs & Ross 1993, David & Gardiner 1962, Karls-
son & Wickman 1990, Labine 1968, Murphy et al. 1983,
Stern & Smith 1960) and at least some moths (Adler
1989, Cheng 1970, Leahy & Andow 1994, Leather
1984, Miller 1989), even though substantial protein is
not acquired after adult eclosion, carbohydrate inges-
tion can profoundly affect longevity and fecundity. Cer-
tain tropical butterflies even appear to supplement their
nitrogen budget by visiting protein-rich mud-puddles
(Beck et al. 1999). The degree of dependence on nectar,
however, varies between species (e.g., Boggs 1997b).
The majority of previous studies are concerned with an-
alyzing correlations between butterfly abundance and
availability or diversity of potential nectar sources (e.g.,
Douwes 1975, Grossmueller & Lederhouse 1987, Hill
1992, Loertscher at al. 1996, Schultz & Dlugosch 1999).
Controlled experimental studies, in contrast, are avail-
able only for a small range of taxa.
With far more than 5000 extant species, the Ly-
caenidae are the second-largest family of true butter-
flies in the world (Heppner 1991). However, hitherto
almost no experimental data on the role of adult feed-
ing in lycaenid butterflies are available (with exception
of the Australian species Jalmenus evagoras; Hill &
Pierce 1989). This paper reports the effects of differ-
ent feeding and temperature regimes on fecundity,
survival and hatching rate of the Purple-Edged Cop-
per butterfly Lycaena hippothoe, in order to assess the
relative importance of income vs. stored resources.
Furthermore, we discuss the relevance of our results
for the population dynamics of this species.
STUDY ORGANISM, MATERIAL AND METHODS
Lycaena hippothoe is a widespread butterfly ranging
from northern Spain in the west throughout much of
the northern Palaearctic region eastwards to the east-
ernmost parts of Siberia and China (Ebert & Rennwald
1991, Lukhtanov & Lukhtanov 1994). In Central Eu-
rope, adults fly in one generation from about mid-June
through late July (Fischer 1998). The species inhabits
different kinds of wetland as well as unimproved grass-
land. The principal larval hostplant is Rumex acetosa L.
(Polygonaceae), a common and widespread perennial
herb occurring in various types of grassland. Recent de-
clines of L. hippothoe populations stimulated concern
among nature conservation authorities and resulted in
the inclusion of this species into “Red Data Lists” in
various European countries (cf. Marttila et al. 1999,
Pretscher 1998, SBN 1987, Tax 1989).
In June 1997 and 1998, respectively, freshly
emerged females (not older than one or two days,
judged by wing wear) of L. hippothoe were caught in
the Westerwald area (western Germany; see Fischer
92
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
a
fo)
oO
a
So
Oo
Accumulated fecundity (no. of eggs)
: ta +
0 4 8 12 16 20 24 28 32 36 40
Days
Fic. 1. Mean accumulated fecundity (solid lines without sym-
bols: +SE of respective means) of Lycaena hippothoe over time un-
der different feeding and temperature regimes (rhombuses: “fed
25°C”, squares: “fed”, triangles “water’).
1998) for rearing experiments and to provide the foun-
dation of a captive breeding stock. Females were
placed individually in glass jars (1 L) containing moist-
ened filter paper and the jars covered with gauze.
Each jar contained a bunch of the larval foodplant R.
acetosa (in H,O) as oviposition substrate.
For measuring the effects of feeding and tempera-
ture regime three experimental treatments were used.
(1) Ambient daylight and temperature conditions (from
mid-June to mid-July 1997, wild-caught females), but-
terflies fed with a highly concentrated sucrose solution.
(2) Conditions as before (from mid-June to mid-July
1998, wild-caught females), but butterflies were pro-
vided with water only. (3) In an environmental cham-
ber at a constant temperature (25°C) and a photope-
riod of L18:D6 (in 1998), fed with a highly
concentrated sucrose solution (these females origi-
nated from captive breeding stock, F generation). The
three treatments are hereafter referred to as “fed” (1),
“water” (2), and “fed 25°C” (3). For egg-laying under
ambient conditions glass jars were put on a table out-
side in a sheltered area. The day-night regime during
the periods was about L16.30:D7.30 (time between
sunrise and sundown excluding civil twilight).
In the morning (i.e., before onset of oviposition activ-
ities) of each day eggs were removed and counted. Eggs
were then exposed to a constant temperature of 20°C
(photoperiod L18:D6) to assess hatching rate. Females
were dissected after death occurred, and the number of
mature oocytes remaining in their ovaries was deter-
mined under a stereomicroscope. Furthermore, 10
freshly emerged and unfed females from captive breed-
ing stock were dissected to analyze the status of oogene-
sis at emergence. Only already yolked oocytes with a di-
ameter of at least 0.5 mm were counted.
Number of eggs
3
.
1 23 4 ©- 6 YY & GY 10 11 12 13
Fic. 2. Fecundity in a single female Lycaena hippothoe pro-
vided with water only. On two occasions (indicated by arrows) the
female was allowed to feed on highly concentrated sucrose solution
for about half an hour each time.
Differences in mean values were tested using the
Mann-Whitney U-test and the Tukey-Kramer post-hoc
comparison after the non-parametric Kruskal-Wallis
H-test (Sachs 1997), since data distributions deviated
strongly from normality. Throughout the text all means
are given +1 SD.
RESULTS
Egg production and hatching rate. Total life-
time egg production was strongly affected by access to
nutrients, but not by temperature regime (see below).
Females fed highly concentrated sucrose solution in
both treatments laid significantly more eggs than those
individuals given water only (H,,, = 14.3, p = 0.0008;
Table 1). The latter achieved, on average, only 14.0 %
of the fecundity of the fed individuals (64.9 + 30.1 vs.
464.2 + 207.1 eggs, data for “fed” and “fed 25°C”
pooled). The number of eggs laid in the “water”-group
coincides with the well developed oocytes at emer-
gence (mean 60.9 + 18.8, n = 10; U-test: z = 0.44, p =
0.66, n.s.). Likewise, total potential fecundity (mea-
sured as the total number of eggs laid plus mature
oocytes remaining after death) was much reduced in
the water-only treatment, but did not differ between
the two sugar-feeding treatments.
In females with no access to carbohydrate sources,
substantial oviposition occurred only during the first
three days of the experiment (Fig. 1). One of those but-
terflies was provided with sucrose solution on two days,
after which it resumed laying eggs again (Fig. 2). These
eggs laid after sucrose feeding were excluded from fur-
ther data analysis.
No significant difference in egg production was
found between females fed at a constant temperature
of 25°C compared to those fed under ambient tem-
VOLUME 54, NUMBER 3
93
Table 1. Fecundity, hatching rate and longevity (means + SD) of female Lycaena hippothoe under different feeding and temperature
regimes (“fed 25°C”: in environmental chamber at a constant temperature of 25°C, fed with sucrose solution; “fed”: ambient temperature con-
ditions, fed with sucrose solution; “water”: ambient temperature conditions, access to water only). Figures within one row followed by the same
letter do not differ significantly (Tukey-Kramer post-hoc comparison after Kruskal-Wallis H-test, threshold for significance p < 0.05).
Fed 25°C
Number of females 6
Number of eggs laid 422.3 + 187.9"
Range (eggs laid) 165-690
Highest egg number/day 98
Number of oocytes 43.0 + 41.0*
Number of eggs and oocytes 465.3 + 163.7?
Relative realized fecundity [%] 88.1 + 11.98
Hatching rate of eggs [%] 83.0 + 11.98
Female longevity [days] 16.8 + 7.8*
Oviposition period [days] 10.8 + 8.28
Fed Water
7 8
500.1 + 245.7" 64.9 + 32.1
126-808 9-102
110 65
26.9 + 23.18 28.5 + 22.88
527.1 + 240.4" 93.4 + 14.9»
93.0 + 6.4" 66.6 + 29.8%
813 = 178? 84.1 + 11.8?
31.7 + 9.7? 6.1 + 3.6°
28.0 + 10.1? 4.0 + 1.55
perature conditions (Table 1). However, under the am-
bient temperature regime it took the females far
longer to reach saturation of the fecundity curve
(about 30 compared to 10 days; Fig. 1). The “steps” in
the fecundity curve of the individuals fed under an
ambient temperature regime are caused by days with
adverse weather conditions and hence no egg-laying.
Realized relative fecundity, expressed as the ratio of
eggs laid relative to the total number of eggs laid plus
remaining mature oocytes after death, was high in fed
individuals and lower, though not significantly so, in
the water-fed butterflies (H,,, = 4.5, p = 0.1; Table 1).
In contrast, hatching rate of eggs was invariably high
(80-85 %) and not affected by adult feeding (H,,, =
0.1, p = 0.95). As all females produced viable eggs,
they obviously were mated.
Longevity. Feeding as well as temperature regime
strongly influenced longevity (Table 1). Sucrose-fed fe-
males (both treatments) lived three to five times longer
than water-fed individuals (H,,, = 14.5, p = 0.0007). Fur-
thermore, sucrose-fed individuals under ambient cli-
matic regime survived twice as long as the group fed at
25°C. Extended survival also distinctly increased the du-
ration of an individual’s oviposition period (Bh pss L511, jo
= 0.0005). Compared to females provided with water
only, among sucrose-fed butterflies the period over
which eggs were laid increased by a factor of 2 (at 25°C)
to 7 (at ambient temperature conditions).
DISCUSSION
Our data demonstrate the exceptional importance
of adult feeding for reproduction in female L. hip-
pothoe. Overall, fecundity in our experiments was
much higher than suggested for this species in the lit-
erature (Bink 1992), and sucrose-feeding increased to-
tal egg production, oviposition period as well as
longevity. Therefore, nourishment from nectar sources
is essential for maintenance of basic metabolic func-
tions as well as for egg production.
Adult-derived resources seem to be even more im-
portant for egg production and longevity than in many
other butterfly species. For example, in Pararge aegeria
(Nymphalidae) egg number is approximately four times
higher after sugar-feeding than after water-feeding
(Karlsson & Wickman 1990). In the checkerspot but-
terfly, Euphydryas editha (Nymphalidae), as well as in
Jalmenus evagoras (Lycaenidae), sugar-feeding roughly
doubles egg production and longevity relative to indi-
viduals kept with no access to food or supplied with wa-
ter only (Murphy et al. 1983, Hill & Pierce 1989). In L.
hippothoe, in contrast, the increase in fecundity was
seven-fold, and longevity was three to five times
greater. Obviously, substantial egg production in L. hip-
pothoe relies strongly on adult-derived rather than lar-
val-derived carbohydrates. Thus, in the continuum be-
tween capital and income breeders the species is
positioned far to the latter side. Until recently, carbo-
hydrate intake was largely regarded as a means of ac-
quiring flight fuel, and its importance for oogenesis was
not clear (Wheeler 1996). But now the incorporation of
carbohydrates into developing oocytes has been con-
firmed directly by radiotracer studies (Boggs 1997a),
and our results further corroborate the relevance of
sugar-feeding for egg production in butterflies.
Surprisingly, hatching rate was not affected by
adult feeding. We conclude that unfed females are
able to lay those eggs only, in which development is
already under way at adult emergence (cf. Boggs &
Ross 1993). This is supported by the coincidence be-
tween the number of well developed oocytes found
in females at emergence with the total egg produc-
tion in the water-fed group (cf. Hill & Pierce 1989,
Watanabe 1992). Oogenesis of those eggs must rely
exclusively on nourishment gathered during larval
stage (probably supplemented by male-derived nutri-
ents received by the female at mating; cf. Boggs
1990, 1997a). The production or maturation of addi-
tional eggs is apparently severely constrained in L.
SIEEEESSRSEEsnnEnEeetereniememeainmmneen
Sep TET
Se ee
— —=
ee ee
94
hippothoe butterflies without adult carbohydrate in-
take. In contrast to the sucrose-fed females, most
oocytes remaining after death were poorly developed
in the water-fed individuals. Obviously, females are
supplied with a rather small amount of stored carbo-
hydrates, which are abundantly available from adult
nectar resources. On the other hand nitrogenous
compounds, which are scarce in the diet of most
(temperate-zone) adult butterflies, are commonly
capital reserves acquired more or less entirely during
the larval stages (but see Beck et al. 1999, Erhardt &
Rusterholz 1998). These stored reserves are used
throughout adult life (Boggs 1997a), whereas larval-
derived carbohydrates decline rapidly and need to be
complemented by income.
Our findings could be of importance for the popula-
tion dynamics of L. hippothoe in at least two ways. As
the number of eggs laid obviously depends on adult re-
sources in this species, adverse weather conditions can
influence population dynamics not only through a
higher adult or larval mortality (e.g., Singer & Ehrlich
1979, Pollard et al. 1997), but also through a decrease
in fecundity. A high incidence of rainy or overcast days
would limit not only the time available for egg-laying,
but also for feeding. Unlike factors affecting oviposi-
tion rate only, there is no compensation for a reduced
egg production caused by lack of nourishment (Gos-
sard & Jones 1977), as nutrient limitations should im-
pede compensation of time limitations through an in-
crease in oviposition rate. In contrast to other species
able to produce substantial egg numbers even without
any adult feeding (e.g., Euphydryas editha; Murphy et
al. 1983), population dynamics of L. hippothoe should
be far more affected if access to adult nutrient re-
sources is limited.
Second, modern agricultural techniques lead to a
reduction of nectar sources (i.e., flowers) through a
high mowing frequency and recurrent application of
fertilizers (Barabasz 1994, Ellenberg 1996, Erhardt
1995, Nigmann 1997). Taking into account the dra-
matic loss of traditionally managed hay meadows in
central European landscapes (e.g., Erhardt 1995, Er-
hardt & Thomas 1991), the concomitant reduction of
flower availability could be an important factor for re-
gional declines in this sedentary species.
Our study revealed that L. hippothoe butterflies must
be regarded, to an unexpected degree, as income breed-
ers. It remains to be tested whether other temperate-
zone lycaenids rely in a similar manner on nectar re-
sources for reproduction. If this were the case, this could
explain the decline of other lycaenid species with com-
mon and widespread hostplants (e.g., L. tityrus Poda, L.
virgaureae L.., sharing the same hostplant, Rumex ace-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
tosa, with L. hippothoe) in modem European land-
scapes. Hence, the importance of the availability of adult
resources for the reproductive biology and persistence of
butterfly populations, which is most commonly inferred
from field data on spatial distributions (e.g., Hill 1992,
Douwes 1975, Loertscher et al. 1996), deserves more ex-
perimental studies on a broader range of taxa.
ACKNOWLEDGMENTS
We wish to thank the Koblenz district government for granting per-
mission to pursue this study, and W. E. Miller and an unknown re-
viewer for valuable comments on the manuscript. This study was sup-
ported by grants from the Friedrich-Ebert-Foundation to K. Fischer.
LITERATURE CITED
ADLER, P. H. 1989. Sugar feeding of the adult corn earworm (Lepi-
doptera: Noctuidae) in the laboratory. J. Econ. Entomol.
§2:1344-1349.
BarRaBASZ, B. 1994. The effect of traditional management methods
modifications on changes in meadows of Molinio-Arrhen-
atheretea class. Wiad. Bot. 39:85-94.
BECK, J., E. MUHLENBERG & K. FIEDLER. 1999. Mud-puddling be-
havior in tropical butterflies: in search for proteins or minerals?
Oecologia 119:140-148.
BINK, F. A. 1992. Ecologische Atlas van de Dagvlinders van Noord-
west-Europa. Schuyt & Co., Haarlem, 512 pp:
Boccs, C. L. 1990. A general model of the role of male-donated nu-
trients in female insects’ reproduction. Am. Nat. 136:598-617.
1997a. Dynamics of reproductive allocation from juvenile
and adult feeding: radiotracer studies. Ecology 78:192—202.
1997b. Reproductive allocation from reserves and income in
butterfly species with differing adult diets. Ecology 78:181-191.
& C. L. Ross. 1993. The effect of adult food limitation on
life history traits in Speyeria mormonia (Nymphalidae). Ecology
74:433-441.
CHENG, H. H. 1970. Oviposition and longevity of the dark-sided cut-
worm, Euxoa messoria (Lepidoptera: Noctuidae), in the labora-
tory. Can. Entomol. 104:919-925.
Davin, W. A. L. & B. O. C. GARDINER. 1962. Oviposition and the
hatching of the eggs of Pieris brassicae in a laboratory culture.
Bull. Entomol. Res. 53:91—109.
Douwes, P. 1975. Distribution of a population of the butterfly
Heodes virgaureae. Oikos 26:332-340.
DUNLAP-PIANKA, H. L., C. L. Boccs & L. E. GILBert. 1977. Ovar-
ian dynamics in Heliconiine butterflies: Programmed senes-
cence versus eternal youth. Science 197:487—-490.
Ebert, G. & E, RENNWALD (EDS.). 1991. Die Schmetterlinge Baden-
Wiirttembergs. Vol. 2. Tagfalter II. Ulmer, Stuttgart, 535 pp.
ELLENBERG, H. 1996. Vegetation Mitteleuropas mit den Alpen. 5.
edition. Ulmer, Stuttgart, 1095 pp.
ERHARDT, A. 1995. Ecology and conservation of alpine Lepidoptera.
pp. 258-276. In A. S. Pullin (ed.), Ecology and conservation of
butterflies. Chapman & Hall, London.
& H.-P. RusterHorz. 1998. Do peacock butterflies (Inachis
io L.) detect and prefer nectar amino acids and other nitroge-
nous compounds. Oecologia 117:536—-542.
& J. A. THomas. 1991. Lepidoptera as indicators of change in
the semi-natural grasslands of lowland and upland Europe. pp.
213-236. In N. M. Collins & J. A. Thomas (eds.), The conserva-
tion of insects and their habitats. Academic Press, London.
FiscuHEr, K. 1998. Population structure, mobility and habitat selec-
tion of the butterfly Lycaena hippothoe (Lycaenidae: Lycaenini)
in western Germany. Nota Lepid. 21:14-30.
GILBERT, L. E. 1972. Pollen feeding and reproductive biology of He-
liconius butterflies. Proc. Natl. Acad. Sci. USA 69:1403-1407.
Gossarp, T. W. & R. E. Jones. 1977. The effects of age and weather
on egg-laying in Pieris rapae L. J. Appl. Ecol. 14:65-71.
VOLUME 54, NUMBER 3
GROSSEMUELLER, D. W. & R. C. LEDERHOUSE. 1987. The role of
nectar source distribution in habitat use and oviposition by the
tiger swallowtail butterfly. J. Lepid. Soc. 41:159-165.
HEPPNER, J. B. 1991. Faunal regions and the diversity of Lepi-
doptera. Tropical Lepidoptera 2, Suppl. 1: 1-85.
HI, C. J. 1989. The effect of adult diet on the biology of butter-
flies. 2. The common crow butterfly, Euploea core corinna. Oe-
cologia 81:258-266.
1992. Temporal changes in abundance of two lycaenid but-
terflies (Lycaenidae) in relation to adult food sources. J. Lepid.
Soc. 46:174-181.
& N. E. Pierce. 1989. The effect of adult diet on the biol-
ogy of butterflies. 1. The common imperial blue, Jalmenus
evagoras. Oecologia 81: 249-257.
KARLSSON, B. & P. O. WickMaN. 1990. Increase in reproductive effort
as explained by body size and resource allocation in the speckled
wood butterfly, Pararge aegeria. Funct. Ecol. 4:6009-6017.
LABINE, P. A. 1968. The population biology of the butterfly Eu-
phydryas editha. VUI. Oviposition and its relation to patterns of
oviposition in other butterflies. Evolution 22:799-805.
Leany, T. C. & D. A. ANDow. 1994. Egg weight, fecundity, and
longevity are increased by adult feeding in Ostrinia nubilalis
(Lepidoptera: Pyralidae). Ann. Ent. Soc. Am. 87:342-349.
LEATHER, S. R. 1984. The effect of adult feeding on the fecundity,
weight loss and survival of the pine beauty moth, Panolis flam-
mea (D&S). Oecologia 65:70-74.
LOERTSCHER, M., A. ERHARDT & J. ZETTEL. 1996. Microdistribution
of butterflies in a mosaic like habitat: the role of nectar sources.
Ecography 18:15-26.
LuKHTANOv, V. & A. LUKHTANOV. 1994. Die Tagfalter Nordwest-
asiens. Herbipoliana vol. 3. Eitschberger, Marktleuthen, 440 pp.
Martriia, O., K. SAARINEN & J. JANTUNAN. 1999. The national but-
terfly recording scheme in Finland: first seven-year period
1991-1997. Nota Lepid. 2217-34.
MILLER, W. E. 1989. Reproductive enhancement by adult feeding:
effects of honeydew in imbibed water on spruce budworm. J.
Lepid. Soc. 43:167-177.
Mureny, D. D., A. E. LAUNER & P. R. EHRLICH. 1983. The role of
adult feeding in egg production and population dynamics of the
checkerspot butterfly Euphydryas editha. Oecologia 56:257-263.
NIGMANN, U. 1997. Tagfaltergemeinschaften in nordbayerischen
Feuchtwiesen-Talriumen—Diversitiit, Struktur und Dyni amik.
PhD Dissertation, Univ ersity of Bayreuth, 154 pp.
POLLARD, E., J. N. GREATOREX-Davigs & J. A. THomas. 1997.
Drought reduces breeding success of the butterfly Aglais ur-
ticae. Teall Ent. 22:315- 318,
PRETSCHER, P. 1998. Rote Liste der Grofischmetterlinge (Macrolepi-
doptera). pp. 87-111. In Bundesamt fiir Naturschutz (ed.), Rote
Liste gefiihrdeter Tiere Deutschlands. Schriftenreihe fiir Land-
schaftspflege und Naturschutz 55. Bonn-Bad Godesberg.
Sacus, L. 1997. Angewandte Statistik. 8. edition. Springer, Berlin,
877
SBN (oaipinmnect ire BUND FUR NATURSCHUTZ, ED.). 1987. Tag-
falter und ihre Lebensriume. Basel, 516 pp. ‘
SCHULTZ, C. B. & K. M. Diucoscu. 1999. Nectar and hostplant
scarcity limit populations of an endangered Oregon butterfly.
Oecologia 119:231-238.
SINGER, M. C. & P. R. Euruicu. 1979. Population dynamics of the
checkerspot butterfly Euphydryas editha. Fortschr. Zool.
25:53-60.
STERN, V. M. & R. F. SMITH. 1960. Factors affecting egg production
and oviposition in populations of Colias philodice eurytheme
Boisduval. Hilgardia 29:411—-454.
TAMMARU, T. & E. Hauktoya. 1996. Capital breeders and income
breeders among Lepidoptera—consequences to population dy-
namics. Oikos 77:561—564.
Tax, M. H. 1989. Atlas van de Nederlandse dagvlinders. Vlinder-
stichting, Wageningen, 248 pp.
WATANABE, M. 1992. Egg maturation in laboratory-reared females of
the swallowtail butterfly, Papilio xuthus L. (Lepidoptera: Papil-
ionidae), feeding on different concentration solutions of sugar.
Zool. Sci. 9:133-141.
WHEELER, D. 1996. The role of nourishment in oogenesis. Ann.
Rey. Entomol. 41:407-431.
Received for publication 26 July 1999: revised and accepted 9 Feb-
ruary 2000.
GENERAL NOTES
Journal of the Lepidopterists’ Society
54(3), 2001, 96-97
NOTES ON THE BEHAVIOR OF SPEYERIA IDALIA (DRURY) (NYMPHALIDAE) LARVAE WITH IMPLICATIONS THAT
THEY ARE DIURNAL FORAGERS.
Additional key words: Speyeria idalia, diumal feeding behavior, tallgrass prairie, Kansas.
Populations of the regal fritillary, Speyeria idalia (Drury), are in de-
cline throughout its range (Hammond & McCorkle 1983, Debinski &
Kelly 1998). This decline has heightened interest in the study of the life
history of this butterfly species, especially the juvenile stages. Under-
standing the larval behavior of S. idalia is an important step in under-
standing the butterfly’s overall life history and ultimately, implement-
ing management plans to protect it along with the remaining tracts of
native tallgrass prairie. Scudder (1889) was the first to note our lack of
knowledge of the natural history of S. idalia larvae. Much of what is
known, such as the daily feeding pattems, is anecdotal and has not yet
been corroborated by laboratory experiments. Larvae of the genus
Speyeria have been reported to feed on violet plants (Viola spp.) at
night and remain hidden during the day, away from their host plants
(Holland 1931, Ehrlich & Ehrlich 1961, Royer 1988). In Kansas, S.
idalia feeds primarily on two species of violet: V. pedatifida G. Don
(the prairie violet) and, to a lesser extent, V. pratincola Greene (the
blue prairie violet). If the larval strategy is to leave their host plant dur-
ing the day, the larvae must either relocate the host plant upon which
they last fed or find a new one. We studied larval S. idalia both in the
field and in the laboratory to determine their daily activity patterns.
We first attempted to rear the larvae according to the procedure
of Mattoon et al. (1971), but survivorship was very low. Therefore, it
was necessary to collect larvae in the field. Field studies were con-
ducted in a native tallgrass prairie (dominated by little bluestem,
Schizachyrium scoparius Michx.; Indian grass, Sorghastrum nutans
(L.) Nash; and big bluestem, Andropogon gerardii (Vitman) 8.0 km
west of the town of Wamego, in Pottawatomie Co., Kansas (TIOS,
R9E, Sec. 1)). For a more through description of the study site, see
Kopper (1997). Searching violet plants at night proved useless. We
spent most of our time searching for larvae on and around violet
plants during the day in mid-April and early May. Even in the day lar-
vae were exceedingly difficult to find, resulting in the small number
(n = 12) used in this study. Where the larvae were found and the be-
havior that they exhibited prior to collection was recorded (Table 1).
Of the 12 field-collected larvae available, three were parasitized by
Hymenoptera and three died from unknown causes, leaving us with six
larvae to use for bioassays. We released one 4th instar larva back to the
prairie and observed it continuously for 24 hours beginning at 0900
CST. Additional studies were conducted in the greenhouse. Larvae (n
= 5) were released into a prairie plot located in the Kansas State Uni-
versity greenhouses. This plot consisted of a portion of native tallgrass
prairie (slated for road development) that was dug out to a depth of 40
cm and transplanted into an open-top plywood box (1.83 x 1.22 m).
The temperature within the greenhouse was maintained as close to
field conditions as possible (28°C during the day, 18°C during the
night; 60% RH). The plot was watered once a week. Natural light was
used for illumination, so the photoperiod also mirrored field condi-
tions. The larvae were observed every other hour over staggered eight-
hour blocks of time to continuously monitor the larvae for 1 week.
We also obtained three larvae from eggs that we reared in the lab
and used to film larval behavior. A sand-filled arena (24 em diameter
desiccation chamber) was used for the assay; violet leaves were placed
into water vessels, which were buried in the sand. The density of violet
leaves in the arena was similar to densities observed in the field. The
bioassay was kept at ~20°C and 60% RH. A continuously operating,
red low intensity (25 w.) incandescent light source permitted us to film
and observe larval behaviors during the scotophase (dark period). The
ratio of photoperiod:scotophase that the larvae would experience in the
field was maintained in the laboratory, but the phases were reversed.
Larvae were held for 1 week under these conditions to allow them to
entrain to the new light regimen prior to the bioassay. The larvae were
filmed and observed bihourly over various times of the day for 1 week.
We found that, in the field, the larvae were considerably more ac-
tive during the day and inactive throughout most of the night. When
inactive, the early instar (1st—early 3rd) larvae could be found in the
curls of the young violet leaves, whereas the later instar (late
3rd-6th) larvae were found at the base of the violet plant. These re-
sults may be biased, because we spent a great deal of time searching
for larvae in violet clumps. During the day (1000-1600 hours), lar-
vae were feeding, walking, or inactive. When not at the base of vio-
let plants, larvae were found feeding on violet plants (both leaves
and flowers) and walking in curved directional paths.
We calculated the probability that larvae would move from one
behavior to another by observing larvae (laboratory larvae were ob-
served bi-hourly for one week and the field larva was observed over
a 24-hour period) and assessing the change in larval behavior. Larval
behavior was characterized into three categories: moving, inactive,
and feeding. The change in larval behavior was pooled separately for
larvae in both laboratory and field observations. Because of the low
sample size (n = 3 for the laboratory assay and n = | for the field as-
say) used for these observations, these results should be interpreted
with caution. Also, any differences between laboratory and field data
may be due to the nature of the laboratory bioassay, such as a greater
likelihood of running into leaves in the arena than finding a plant in
the field. Furthermore, temperature or time of day was regrettably
not recorded and which may have also influenced larval behavior.
Following a period of inactivity, the larvae often started to move as
opposed to feeding (52.4% and 75.0% laboratory and field, respec-
TaBLE 1. Behavior of larvae found in the wild. All larvae were collected during daylight hours. Time, when recorded, is expressed in mili-
tary time. nr = not recorded.
Individual Date Time
1 4/16/97 nr
2 4/22/97 nr
3 4/23/97 10:00-10:30 h
4 4/23/97 10:00-10:30 h
5 4/24/97 10:00-11:00 h
6 4/28/97 11:00-12:00 h
i 4/28/97 11:00-12:00 h
8 5/1/97 nr
9 5/8/97 10:30-11:30 h
10 5/9/97 nr
11 5/9/97 nr
12 5/9/97 nr
Instar Behavior when found
4th Inactive at the base of V. pratincola.
3rd Inactive at the base of V. pedatifida.
3rd Inactive at the base of V. pedatifida.
4th Inactive at the base of V. pedatifida.
3rd Inactive at the base of V. pedatifida.
3rd Feeding on V. pedatifida leaves.
3rd Inactive at the base of V. pedatifida.
3rd Feeding on V. pedatifida flower.
4th Inactive at the base of V. pedatifida.
5th Walking towards V. pedatifida.
5th Inactive at the base of V. pedatifida.
5th Feeding on V. pedatifida leaves.
VOLUME 54, NUMBER 3
tively). After moving, the larvae typically became inactive again, only
feeding 27.8% in the laboratory and 25.5% in the field. However, in
the field, encountering food is not guaranteed, and these percent-
ages do not take into account distance traveled. Once a larva in the
field had fed, it would either remain inactive or move with equal
probability. However, larvae in the laboratory would remain inactive
after feeding 81.3% of the time.
Larvae would eat the violet leaves rapidly during feeding bouts
and, when not feeding, larvae would hide in clumps of grass or
walk. McCorkle and Hammond (1988) observed similar feeding
behavior for S. zerene hippolyta. The fifth instar larva that was re-
leased back to the prairie walked in a curved path (25.40 m over a
24 hour period) and fed only on violet plants that were in its path
(although it walked past violet plants as close as a centimeter away
and apparently did not perceive them). Wind direction did not
seem to matter as the larvae walked equally close to violets up and
down wind without noticing them. As the sun set, the larva walked
and fed progressively less, until it became inactive at the base of a
grass clump and remained there until the following morning. The
larvae studied in the laboratory and greenhouse displayed a similar
diurnal feeding pattern. The inactivity associated with nightfall
may be due to a reduction in temperature, however temperature
does not drop substantially as soon as the sun sets in eastern
Kansas, leading us to believe that light is a more important cue for
activity than temperature.
This study of a small sample of S. idalia larvae in Kansas tall-
grass prairie indicates that they do not forage entirely at night.
More study is needed covering a larger geographic region in order
to determine how widespread this behavior is within S. idalia and
the genus Speyeria.
ACKNOWLEDGMENTS
We thank Alberto B. Broce and Matt R. Whiles, Department of
Entomology, Kansas State University and William F. J. Parsons,
Department of Entomology, University of Wisconsin for helpful
comments on earlier versions of this manuscript. Voucher speci-
mens (Lot # 076) are deposited in the KSU museum of Entomo-
logical & Prairie Arthropod Research. This is contribution No. 99-
515-J of the Kansas Agricultural Experiment Station, Kansas State
University, Manhattan, Kansas 66506.
Journal of the Lepidopterists’ Society
54(3), 2001, 97-100
Si
LITERATURE CITED
DeEBINSKI, D. M. & L. KELLy. 1998. Decline of Iowa populations of
the regal fritillary (Speyeria idalia) Drury. J. lowa Acad. Sci.
105:16—22.
EHRLICH, P. A. & A. H. EHRLICH. 1961. How to know the butter-
flies. Wm. C. Brown Co., Dubuque, Iowa. 262 pp:
HAMMOND, P.C. & D. V. McCorkK LE. 1983. The decline and extinc-
tion of Speyeria populations resulting from human environ-
mental disturbances (Nymphalidae: Argynninae). J. Res. Lepid.
22:21. 7-224.
HOLLAND, W. J. 1931. The butterfly book. Second ed. Doubleday.
New York, New York. 424 pp. i
KopPeER, B. J. 1997. Behavioral ecology of the regal fritillary, Speye-
ria idalia (Drury) (Lepidoptera: Nymphalidae), in Kansas tall-
grass prairie: reproductive diapause, factors influencing ovipo-
sition site selection, and larval foraging behavior. M.S. Thesis,
Kansas State University, 105 pp.
MaTToon, S. O., R. D. Davis, & O. D. SPENCER. 1971. Rearing
techniques for species of Speyeria (Nymphalidae). J. Lepid.
Soc. 25:247-256.
McCorkLg, D. V. & P. C. HAMMonpD. 1988. Biology of Speyeria
zerene hippolyta (Nymphalidae) in a marine-modified environ-
ment. J. Lepid. Soc. 42:184-195.
Royer, A. R. 1988. Butterflies of North Dakota. Minot State Uni-
versity, Minot, North Dakota. 192 pp.
SCUDDER, S. H. 1889. Butterflies of the eastern United States and
Canada with special reference to New England. Published by
the author. Cambridge, Massachusetts. Vol. 1, pp. 1-776; vol. 2,
pp. 767-1774; vol. 3, pp. 1775-1958, pls. 1-89, 3 maps.
BRIAN J. Kopper,’? Davip C. MARGOLIES, AND RALPH E. CHARL-
TON, Department of Entomology, Kansas State University, Manhat-
tan, Kansas 66506-4004, USA.
‘Present address: Department of Entomology, University of Wis-
consin, 237 Russell Labs, Madison, Wisconsin 53706, U.S.A.
? Corresponding author.
Submitted for publication 14 August 1999 , revised and accepted 25
July 2000.
BIOLOGY OF ADELPHA MYTHRA FEEDING ON ASTERACEAE, A NOVEL PLANT FAMILY FOR THE NEOTROPICAL
LIMENITIDINAE (NYMPHALIDAE), AND NEW DATA ON ADELPHA “SPECIES-GROUP VII”
Additional key words: Adelpha syma, Adelpha cocala, life history, Rosaceae, Rubiaceae.
The Neotropical genus Adelpha Hiibner (Nymphalidae) includes
about 85 species (Keith Willmott pers. comm.) spread from western
USA to Uruguay, and occurring in a wide variety of habitats and veg-
etation types (Aiello 1984). Species determination is very difficult in
some Adelpha groups, and the natural divisions of the genus are not
yet fully resolved, although a number of species relationships have
been proposed on the basis of the immatures (Aiello 1984). Unfortu-
nately, immatures are known for only 32 species of Adelpha, solely 21
of which have some portion of the early stages illustrated. Thus, al-
though a cladistic analysis of the genus is needed, it would be impos-
sible at this time. Because information on additional species is essen-
tial to a better understanding of the genus (DeVries 1987; Aiello
1991), it is important that any new data about Adelpha immatures be
reported (Otero & Aiello 1996).
This paper describes the immature stages of Adelpha mythra (Go-
dart 1824) and A. syma (Godart 1824), reports their larval host plants,
and discusses the position of both species within Adelpha, based on
their immatures.
Study sites and methods. Adelpha mythra, a montane species in
southeast Brazil, is one of 16 species of Adelpha known in the Santa
Genebra Forest Reserve (22°44’S, 47°06’W, altitude 600-630 m), a
250 ha fragment of semideciduous forest in Campinas, Sao Paulo
State, SE Brazil (see additional information on the area in Morellato
and Leitao-Filho 1995). In January 1999, a female A. mythra was ob-
served there ovipositing on the scandent vine, Mutisia coccinea St.
Hil. (Asteraceae). The egg did not hatch, so this very unusual “record”
was thought to be an oviposition mistake of this female. However,
from February to April 1999, A. mythra was reared from first to
fourth instars collected on the same plant species and also on Bathysa
meridionalis (Rubiaceae) in several parts of the Serra do Japi
(23°11’S, 46°52’W), a mountain range (700-1300 m altitude) covered
by semideciduous forest, in Jundiai, Sao Paulo State, SE Brazil
(Brown 1992). Immatures of Adelpha syma were also found on Rubus
(Rosaceae) in the Serra do Japi, and immatures of A. cocala were dis-
covered feeding on a Rubiaceae in the Parque Ecolégico do Voturua
(46°22’W, 93°57S, altitude 20-100 m), a 200 ha fragment of lowland
subtropical rainforest in the city of Sao Vicente, coastal Sao Paulo
State, SE Brazil.
98
Fic. 1. Fifth instar larvae of Adelpha syma (above) and A.
mythra (below)
Larvae were reared in plastic cages cleaned daily, following Fre-
itas (1991). Adults, head capsules and pupal skins are in the collec-
tion of the first author. Larval food plant vouchers, identified by Dr.
Jorge Tamashiro, have been deposited in the herbarium of the Uni-
versidade Estadual de Campinas.
Descriptions of immatures and host plants of Adelpha
mythra. The only egg observed was greenish brown, sculptured
with hexagonal pits, with spines arising from the pit junctions, con-
sistent with eggs described for other species of Adelpha. The egg
was placed on the upper surface of the leaf, near the apex. It was laid
rapidly by a startled female, and did not hatch.
First and second instars were not described in detail. The third
instar had conspicuous head scoli and a faintly visible variegated pat-
tern that, with few changes, was maintained and intensified through
the remaining larval stadia. The fifth instar (Fig. 1) was variegated
green, with a pattern of oblique lateral stripes. The green areas
changed to light orange as the larvae neared pupation. In the final
larval stadium, the body scoli were short and thick, with a dense cov-
ering of spines; the scoli on A2 were arched posteriorly. The distri-
bution of the scoli (Fig. 2) was the same as for most Adelpha species.
The mature larva was about 25 mm long.
The first through fourth instars initiated feeding at the apex of a
leaflet, leaving the midvein intact and extending it with fecula and
silk to form “frass chains” (Aiello 1984). They rested upon these
structures when not feeding. In addition, they attached dead leaf
fragments and clumps of fecula to the base of the chains. When
feeding on Bathysa meridionalis, larvae built “frass chains” on other
parts of the same large leaf.
The pupa (Fig. 3A) showed an elongated general profile (about
17 mm long), with segment A2 produced and curved anteriorly, and
segment T2 pointed and directed posteriorly. The head horns were
pointed and shaped like tiny asymmetrical leaves, curving out from
the sides of the head. The general color was brown, with dark lines
on the wing pads and no reflective areas.
Mutisia coccinea (Asteraceae), a scandent shrub common in wet
second growth habitats, was observed as the larval food plant of
Adelpha mythra both in Santa Genebra and in the Serra do Japi. In
the latter site, A. mythra was also reared on Bathysa meridionalis L.
B. Smith & Downs (Rubiaceae), a plant with enormous leaves (up to
1m in length and width), of montane habitats, where it occurs most
often near watercourses. Bathysa meridionalis is used by many other
Adelpha species in that site (Brown 1992 and AVLF pers. obs.).
Immatures and host plants of Adelpha syma. The egg was
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
SPIRACLE
A. mythra LONG SHORT
SCOLUS
SIMPL
SCOLUS
LEG PROLEG
Fic. 2. Above, Chaetotaxy of first instar larva of Adelpha syma.
Below, Distribution of scoli in a fifth instar larva of A. mythra.
greenish brown, with sculpturing and ornamentation as described
for other species of Adelpha, placed on the upper surface of the leaf,
near the apex. The height and diameter were about 0.8 mm; the du-
ration was not determined.
The first instar was entirely brown, with pale body setae about
0.06 mm long, arranged as in Fig. 2. The head capsule was pale
brown without ornamentation. The final body length was 4 mm. The
second instar was entirely brown, with a spiny brown head. The body
bore short stubby scoli. The final body length was 6 mm. The third
and fourth instars showed a clearly visible variegated pattern, with
head spines longer. The final lengths were 9 mm in the third and 15
mm in the fourth larval stadia. The fourth and fifth instars had the
same general pattern. The fifth instar (Fig. 1) was variegated with
green, cream and brown, showing a general pattern of oblique lat-
eral stripes. The green areas change to light orange before pupation.
The body scoli were short and thick, with dense spines, and the scoli
of A2 arching posteriorly. The distribution of the scoli was the same
as in A. mythra. The fully grown larva was 25 mm long. Data on
head capsule widths for all instars are in Table 1. The first through
fourth instars constructed frass chains and had a behavior similar to
that described for A. mythra.
The pupa (Fig. 3B) showed an elongated general profile (about
18 mm long), with segment A2 projecting and curved anteriorly, and
segment T2 pointed and directed posteriorly. The head homs were
very small and pointed, curved out from the sides of the head. The
general color was brown, with dark lines on the wing pads and no re-
flective areas.
The host plant in the Serra do Japi was Rubus brasiliensis Mart.
(Rosaceae), a common blackberry of sunny second growth habitats, es-
pecially in montane sites. Rubus rosifolius Sm., an introduced species,
is also used as larval food in other montane sites in SE Brazil.
Positions within Adelpha. The scolus shape and the general
pattern of the larvae, and the general form of the pupae, suggest
TABLE 1. Head capsule widths of Adelpha syma
Larval instar Range (mm) Mean sD n
ie 0.56-0.58 0.57 0.011 @
a 0.78—-0.82 0.80 0.014 5
3° 1.18-1.22 1.18 0.042 6
4° 1.64-2.03 1.81 0.151 13
5° 2.73-3.12 2.89 0.133 11
VOLUME 54, NUMBER 3
that both Adelpha mythra and A. syma belong to the Species-
Group VII of Aiello (1984). The immatures of both species are
very similar to those of A. cocala, the main difference being that
the A2 process of the pupa is much longer in A. mythra and more
arched in A. syma than in A. cocala (Fig. 3C). In A. mythra, the
length of the A2 projection approaches the condition observed in
A. phylaca (as described by Miiller 1886) (Fig. 3D), a species be-
longing to Group II of Aiello (1984). However, the T2 projections
of A. mythra and A. syma are sloped posteriorly, and not curved
upward as in Group II pupae, giving pupae in the two groups dis-
tinctive general appearances. The pupal head horns of A. mythra
(Fig. 3A) are similar in shape to those of A. cocala (Fig. 3C), but
are farther apart at their bases and are more tapered. The head
horns of A. syma (Fig. 3B) are similar to those of A. phylaca (Fig.
3D), but are more curved.
Additional species of Adelpha need to be reared in order to
clarify the scenario based on the morphology of the immatures and
to make a cladistic analysis possible. Some species groups are
based on only one, two or three species, and the immatures of
many common species remain to be discovered, or their descrip-
tions are not sufficiently detailed to permit assignment to a species-
group. Additional descriptions of Adelpha immatures with figures
are important, especially when they show apparent deviations from
the eight known species groups (Aiello 1984, Otero & Aiello 1996).
Host plant use. Among the Nymphalidae, the association with
Asteraceae as larval food plants is found in only a few groups (es-
pecially Melitaeinae and Acraeinae) (Ackery 1988, Freitas 1991).
——————_
5mm
99
The record of Asteraceae as a larval host of the Limenitidinae rep-
resents a new plant family for neotropical Adelpha. Larvae of Adel-
pha species have been recorded as feeding on Aquifoliaceae, Aster-
aceae, Bombacaceae, Caprifoliaceae, Combretaceae, Ericaceae,
Fagaceae, Flacourtiaceae, Icacinaceae, Malpighiaceae, Melastom-
ataceae, Moraceae, Ochnaceae, Piperaceae, Rosaceae, Rubiaceae,
Tiliaceae, Ulmaceae, Urticaceae, Verbenaceae and Vochysiaceae
(Jones & Moore 1883, Miiller 1886, Moss 1933, Biezanko et al.
1966, Aiello 1984, 1991, DeVries 1987, Ackery 1988, Brown 1992,
Otero & Aiello 1996, Diniz & Moraes 1997, Constantino 1998, and
this work). Some themes may be recognized in the different
species-groups (Aiello 1984). Basically the species of Adelpha can
be sorted into rubiaceous feeders and non-rubiaceous feeders. Four
examples of species feeding on both Rubiaceae and other families
have been reported (A. serpa, A. boreas tizona, A. syma and A. co-
cala), and A. mythra is the fifth recorded case. Although interest-
ing, this pattern must be considered with caution, because some
plant identifications need to be confirmed by additional field obser-
vations.
ACKNOWLEDGMENTS
This study was conducted as part of a Ph.D. Thesis on Nymphali-
dae systematics and evolution. K. Willmott, Deane Bowers and an
anonymous referee reviewed and made helpful critics and comments
on the manuscript. This research was supported by fellowships from
the Brazilian CNPq to AVLF and KSB.
Fic. 3. Pupae of Adelpha mythra (A), A. syma (B), A. cocala (C) and A. phylaca (D) (A. phylaca redrawn from Aiello, 1984, in a different
scale; the bar means 7.3 mm).
——————ESESsS
VOLUME 54, NUMBER 3
LITERATURE CITED
AcCKERY, P. R. 1988. Hostplants and classification: a review of
nymphalid butterflies. Biol. J. Linn. Soc, 33:95-203.
AIELLO, A. 1984. Adelpha (Nymphalidae): deception on the wing.
Psyche 91:1-45.
AIELLO, A. 1991. Adelpha ixia leucas: Immature stages and position
within Adelpha (Nymphalidae). J. Lepid. Soc. 45(3):181-187.
BIEZANKO, C. M., A. RUFFINELLI & C. S. CARBONELL. 1966. Lepi-
doptera del Uruguay. Notas complementarias. III. Boletin
Facul. Agron. Univ. Repub. (Montevidéo) 91:1-53.
Brown, K. S. JR. 1992. Borboletas da Serra do Japi: Diversidade,
habitats, recursos alimentares e variagao temporal, pp. 142-187,
18 figs. In L. P. C. Morellato (ed.), Historia natural da Serra do
Japi. Ecologia e preservagao de uma area florestal no sudeste do
Brasil. Campinas, Editora da Unicamp/Fapesp.
CONSTANTINO, L. M. 1998. Butterfly life history studies, diversity,
ranching and conservation in the Chocé rain forests of western
Colombia (Insecta: Lepidoptera). Shilap Revta. Lepid.
(Madrid) 26 (101):19-39.
DEVRIES, P. J. 1987. The butterflies of Costa Rica and their natural
history. Princeton University Press, Princeton, New Jersey.
Diniz, I. R. & H. C. Moraes. 1997. Lepidopteran caterpillar fauna
of cerrado host plants. Biodiversity and Conservation
6:817-836.
Freitas, A. V. L. 1991. Variagao morfoldégica, ciclo de vida e sis-
tematica de Tegosa claudina (Eschscholtz) (Lepidoptera,
100
Nymphalidae, Melitaeinae) no estado de Sao Paulo, Brasil.
Revta. bras. Ent. 35:301-306.
Jones, E. D. & F. Moors. 1883. Metamorphoses of Brazilian Lepi-
doptera from San Paulo, Brazil, with nomenclature and de-
scriptions of new forms. Second series. Proceedings of the Lit-
erary and Philosophical Society of Liverpool 37:229-259.
MoreLiato, L. P. C. & H. LEITAO-FILHO. 1995. Introdugao, pp.
15-18. In L. P. C. Morellato & H. Leitao-Filho (eds.), Ecologia
e preservacao de uma floresta tropical urbana. Reserva de Santa
Genebra. Campinas, Editora da Unicamp.
Moss, M. 1933. Some generalizations on Adelpha, a neotropical
genus of nymphalid butterflies of the group Limenitidi. Novit.
Zool. (Tring) 39(1):12-20.
MULLER, W. 1886. Sudamerikanische Nymphalidenraupen: Versuch
eines naturlichen Systems der Nymphaliden. Zoologische
Jahrbucher (Jena) 1:417-678.
OTERO, L. D. & A. AIELLO. 1996. Descriptions of the immature stages
of Adelpha alala (Nymphalidae). J. Lepid. Soc. 50(4):329-336.
ANDRE VICTOR LUCCI FREITAS, KEITH S. BROWN, JR., Museu de
Historia Natural, Instituto de Biologia, Universidade Estadual de
Campinas, CP 6109, 13083-970, Campinas, Sao Paulo, Brazil, AND
ANNETTE AIELLO, Smithsonian Tropical Research Institute, Box
2072, Balboa, Ancon, Panama.
Received for publication 9 August 1999; revised and accepted 2
October 2000.
BOOK REVIEWS
Journal of the Lepidopterists’ Society
54(3), 2001, 101
HAWKMOTHS OF THE WORLD: AN ANNOTATED AND ILLUSTRATED RE-
VISIONARY CHECKLIST (LEPIDOPTERA: SPHINGIDAE), by Ian J. Kitching
and Jean-Marie Cadiou. Published by The Natural History Mu-
seum, London and Cornell University Press, Ithaca, New York. 226
pp. 8 color plates. Hardcover, 8.8 x 11.3 inches, ISBN 0-801437-34-
2. Available from the publisher, Price $95.
Description: This large-format book documents the current
nomenclatorial status of more than 3800 species and genus-group
names for the Sphingidae. Kitching and Cadiou recognize 203 gen-
era and 1272 species as valid, and include all known available and
unavailable synonyms in the checklist. The authors are to be con-
gratulated for their encyclopedic scholarship in updating and cor-
recting sphingid nomenclature and their comprehensive and critical
review of the status of species and generic names. Nevertheless, as
the authors acknowledge, there is still much work to be done on this
charismatic moth family, including a full revision and a thorough
phylogenetic analysis.
The book begins with a 34-page introduction containing short
sections on morphology of adults and early stages, natural history,
economic impact, rarity, and conservation (the authors argue that
sphingid species are not necessarily as rare as they may seem).
There is a brief sketch of the history of higher-level classification,
and a table presenting a partially ordered “current best estimate”
classification based on unpublished cladistic analyses by Kitching.
This is followed by notes on biogeography, and a description of prior
faunal lists and catalogs (some of the most important or recent are
Rothschild and Jordan (1903, A revision of the lepidopterous family
Sphingidae, Novitates Zoologicae, 9 (suppl.):1-972.), Hodges (1971,
Sphingoidea, Moths of America North of Mexico including Green-
land, 21:1-158), d’Abrera (1987, Sphingidae Mundi, Hawk moths of
the world, E. W. Classey, Farringdon, U.K., 226 pp.), Bridges (1993,
Catalog of the family-group, genus-group and species-group names
of the Sphingidae of the world, C. A. Bridges, Urbana, Ilinois, 282
pp-), Pittaway (1993, The Hawkmoths of the Western Palaearctic,
Harley Books, Colchester, U.K., 240 pp.), Carcasson and Heppner
(1996, Sphingoidea, 118, Sphingidae. In J. B. Heppner, ed., Atlas of
Neotropical Lepidoptera, checklist. 4B. Drepanoidea, Bomby-
coidea, Sphingoidea. Association for Tropical Lepidoptera,
Gainesville Florida, pp. 50-60, 62.), Zhu and Wang (1997, Lepi-
doptera Sphingidae. Fauna Sinica (Insecta) 11:1-410) and Danner,
Hitschberger and Surholt (1998. Die Schwdrmer der westlichen
Palaearktis, Bausteine zu einer Revision (Lepidoptera: Sphingidae).
Textband. Herbipolania 4(1):1-368; Tafelband. Herbipolania
4(2):1-720). A section on methodology describes the authors’ con-
cepts of genus, species and subspecies, and explains their differ-
ences of opinion on circumscription of taxa with respect to Danner
et al.’s (1998, op.cit.) more finely split treatment of the Palearctic
fauna. The introduction ends with instructions on how to interpret
abbreviations and annotations to the checklist itself.
The checklist is organized in nested alphabetical order, first by
genus, then by species within each genus, then by subspecies within
species. Synonyms, infrasubspecific names, and nomina nuda are
listed beneath the valid species or subspecies name to which they
correspond. Author and year of publication are given for each name,
and all original descriptions are included in the References Cited
section. There are eight good-quality color plates: six illustrate
pinned specimens of 48 species (including six holotypes, two
paratypes, and two lectotypes), and two show photos of living speci-
mens at rest in the wild. The latter are slightly grainy and the color
reproduction seems a bit saturated. The dust jacket is illustrated
with rather dark images of five species named after either Cadiou or
Kitching which are aia reproduced elsewhere in the book.
The bulk of the text comprises 627 nomenclatorial notes. These
range from short reports on lectotype designations, transcriptions of
data labels and other rather dry taxonomic minutia, to long and
sometimes fascinating detective stories surrounding cases of mis-
taken identity, nomenclatorial squabbles that are reminiscent of
Francis Hemming’s pronouncements (Hemming 1967, The generic
names of the Rane rflies and their type-species (Le pidopte ra:
Rhopalocera). Bull. BM(NH) Ent. Suppl. 9:1-509), and philosophi-
cal explorations of the ICZN Code (1985, International Commission
on Zoological Nomenclature, International Code of Zoological
Nomenclature, International Trust for Zoological Nomenclature and
the British Museum (Natural History), London.)) and its relation to
species concepts.
The book concludes with a comprehensive Literature Cited sec-
tion, descriptions of two newly-recognized species and two new sub-
species, a one-page Addenda section, and good subject and taxo-
nomic indices. Two of the four descriptions of new taxa are
nomenclatorial corrections of discrepancies between current con-
cepts and old nominal types, the third gives a differentiated geo-
graphical form subspecific status and the last is a previously unrec-
ognized species. Only two of the new taxa are illustrated.
In general, the book seems well-organized and error-free. I no-
ticed only one error (a misnumbered footnote reference), which is
quite remarkable given the complex cross-referencing between the
checklist and the notes.
Two Criticisms: As the authors state (p. 24), “Sphingidae Mundi
[d’Abrera 1985, op.cit.| remains the only readily available color
guide to adult hawkmoths on a global basis . . . and will continue to
be used to identify these moths,” They justify their checklist as a
means to correct the numerous nomenclatorial and identification er-
rors in that work, (as well as a means to forestall entry into the liter-
ature of the “almost 1000 errors” in the Bridges’ [1993, op.cit.] cata-
log). Although it exhibits impressive scholarly rigor, in my view
Kitching andl Cadiou’s book will not be of much use or interest to the
casual reader, or even to the serious amateur, because the practical
information it contains is almost completely inaccessible, buried in
the midst of a jumble of nomenclatorial notes that are arranged in
the alphabetical order of the checklist. For example, suppose you
identify a specimen as Oxyambulyx placida Moore by comparing it
to the pictures in d’Abrera’s book. To discover that d’Abrera’s speci-
men was misidentified, you would need to stumble upon Kitching
and Cadiou’s footnote 56 (to Ambulyx semiplacida Inoue), which is
not cross-referenced to either the synonym Oxambulyx Rothschild
& Jordan, nor the species A. placida (Moore). Given the authors’ ac-
knowledgment of the continuing importance of d’Abrera’s book to
sphingid workers, two extremely practical sections to have included
in Kitching and Cadiou would have been, 1) a concise list of corri-
genda to d’Abrera’s nomenclature and identifications, and 2) sup-
plemental illustrations of the 200+ taxa d’Abrera excluded from
Sphingidae Mundi.
A sign of our mercenary times is the Natural History Museum's
promotion of what is essentially a rather narrowly-focussed mono-
graph as a trade book. The inclusion of color plates (and of the term,
“Tllustrated” on the glossy dust jacket) is a lure that will entice many
hawkmoth fanciers to buy this fall but I suspect that most of them
will find its contents lar gely impenetrable. Once upon a time, the
British Museum (Natural History) published scholarly nomenclato-
rial works in its Bulletin (e.g., Hemming 1967), and distributed
them free of cost. Nowaday: 's, the more revenue-oriented Natural
History Museum encourages its systematists to produce marketable
products: another recent “example is Malcolm Scoble’s geometrid
catalog (1999. Geometrid Moths of the World: a catalogue. CSIRO
publications: Melbourne, 1400 pp., $395). I can think of no better
way to prolong the lepidopterological imperialism of wealthy west-
er nations than by publishing catalogs and checklists of global fau-
nas in prohibitively expensive formats that the researchers from
countries where most of the taxa occur generally cannot afford. It is
onerous enough for Ecuadorean le -pidopterists to have to go to Lon-
don to see the holotypes of Ecuadorean moths. The least the Nat-
ural History Museum could do is to give something back to the “type
localities” by making the knowledge ear ed from its holdings acces-
sible at moderate cost, or better yet, free on the WWW.
ANpDREW V. Z. BROWER, Department of Entomology, Oregon
State University, Corvallis, Oregon 97331, USA.
VOLUME 54, NUMBER 3
Journal of the Lepidopterists’ Society
54(3), 2001, 102
CHECKLIST OF THE GEOMETRIDAE (LEPIDOPTERA) OF THE FORMER
U.S.S.R., by Jaan Viidalepp. 1996. Published by Apollo Books Aps.,
Kirkeby S Sand 19, DK-5771 Stenstrup, Denmark. 111 pages. Sole
cover, 25 x 17 cm., ISBN 87-88757-0: 6, Price DKK 200.
This small book is a sequentially numbered annotated distribu-
tional checklist of the 1486 species of Geometridae recorded from
the former U.S.S.R. and updates Jaan Viidalepp’s previous list of
1247 species (1976-1979, A list of Geometridae (Lepidoptera) of the
USSR. I-IV. Entomol. Obozrenie, 55 (4): 842-852, 56 (3): 564-576,
57 (4): 752-761, 58 (4): 782-798. (in Russian)). This is one of the au-
thor’s first publications in English.
In a brief forward Viidalepp explains the checklist represents a
personal effort to summarize his own fieldwork (1961-1992) and
compile information previously published in regional faunas. A list
of included geopolitical regions and their corresponding faunistic
literature follows. The short reference list includes 39 systematic
and faunistic papers published between 1976 and 1996 and 35 of
these are in Russian or German. For literature prior to 1976 the
reader is referred to the author's previous published list (Viidalepp,
op. cit.). Following the 83 page checklist, two indices are provided,
the first to higher categories, and the second to species group
names
The region covered by Checklist of the Geometidae of the For-
mer U.S.S. R includes the lar ge Russian Federation extending from
eastern Europe to the Baer and north to the Arctic Owesm. the
Baltics, Belarus, Ukraine, Moldova, Georgia, Armenia, Azerbaijan,
Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan.
To present distributional information for this vast and politically
challenged segment of the world Viidalepp utilizes an unconven-
tional combination of political names and physical geographic fea-
tures. For Asia in particular emphasis is placed on stable geomor-
phic features such as mountain ranges. Unfortunately the author
does not provide a gazetteer or map to familiarize the reader with
this geographically complex region. For many species entries ex-
tralimital distributional Peeuion is also provided with neighbor-
ing states listed when distributions extend outside the territory of
the former U.S.S.R. For holarctic species, North America is some-
times included in the extralimital distribution but inconsistently so.
Many well known holarctic geometrids such as Epirrhoe alternata
(Miiller), Rhewmaptera (Hydria) undulata (Linnaeus), and Ectropis
crepuscularia (Denis and Schiffermiiller) as well as palearctic in-
troductions to North America including Thera juniperata (Lin-
naeus), Operophtera brumata (Linnaeus), Pasiphila [Rhinoprora in
this list; Chloroclystis in North American lists] rectangulata (Lin-
naeus), Aplocera plagiata (Linnaeus) and erties aestivaria
(Hiibner) are not acknowledged to occur on the North American
continent.
The geometrid fauna Viidalepp treats includes the following sub-
families followed by the number of species listed (the corresponding
number of North American species recorded north of Mexico is
given second for comparison based on the list compiled by D. C.
Ferguson (pp. 88-107 In R. W. Hodges et al., 1983, Check List of
the Lepidoptera of America North of Mexico, E. W. Classey, Ltd.,
London, England, 284 pp.): Archiearine (4/2), Alsophilinae
(Oenochrominae sensu lato, part) (14/3), Larentiinae (706/467),
Sterrhinae (204/96), Geometrinae (66 / 76), Ennominae (488/760),
and Orthostixinae (4/0). At least some of the genera included in the
last subfamily by Viidalepp and others might be more appropriately
placed in the Ennominae but their placement remains unresolved
and they cannot be included in a strict concept of Oenochrominae
(see Miiller, B., 1996, Geometridae pp. 218-249 In O. Karscholt &
J. Razowski (eds.), The Lepidoptera of Europe, Apollo Books, Sten-
strup, Denmark, 380 pp.; and Holloway, J. D., 1996, The Moths of
Borneo Part 9: Family Geometridae, Subfamilies Oenochrominae,
Desmobathrinae and Geometrinae, Malayan Nature Journal 49:
147-326, published by Southdene Sdn. Bhd., Kuala Lumpur,
Malaysia). The total geometrid fauna catalogued by Viidalepp (1486
sequentially numbered entries; doubtful records also included
within the list but not numbered) is comparable to the total number
of geometrid species in North America north of Mexico (1404).
However in the Eurasian fauna the subfamilies Larentiinae and
Sterrhinae are disproportionately represented as exemplified by 253
species of Eupithecia Curtis and 69 species of Idaea Treitschke in
the Viidalepp list (corresponding figures for North America north of
Mexico are 157 and 26 respectively). Although Viidalepp’s checklist
provides distributional information for each species, an overview of
faunistic affinities and biogeographic distributional pattems within
Eurasia is not provided.
It is unfortunate Viidalepp does not discuss or more completely
reference recent systematic literature. Approximately 200 species
group and over 20 generic names listed in the Checklist of the
Geometidae of the Former U.S.S.R were published since 1970 re-
flecting considerable recent systematic effort especially in Asian
countries. Species group and generic synonymies are incomplete
and only selectively provided. Many recent new combinations, new
synonymies, status changes, and revivals cannot be traced using this
checklist alone. In an attempt to be inclusive, a number of unpub-
lished taxa are listed with notations such as “sp. n. (in print)” [=in
press?] and “sp. n. (in prep.)”. Even more tentative is larentiine
species number 185: “G. nov. sp. n.” listed in the tribe Cidariini. The
tribe “Hierochthoniini (trib. n.)” appears on page 61 of the list and
includes the geometrine genera Hierochthonia Prout and Hissarica
Viidalepp but without description of the new higher category.
In summary the Checklist of the Geometidae of the Former
U.S.S.R is a distributional list that also serves as an introduction to
the geometrid fauna of a large and diverse region. Lepidopterists
unfamiliar with the Eurasian fauna and its biogeography can delve
further with the aid of the Zoological Record, a good atlas, and the
references provided. English speaking geometrid specialists eagerly
await additional faunistic and systematic publications treating the
Geometridae found within the former U.S.S.R.
GEORGE J. BALOGH, 6275 Liteolier, Portage, Michigan 49024, USA.
Journal of the Lepidopterists’ Society
54(3), 2001, 102-104
THE Motus oF BORNEO, Part 9, Family Geometridae (incl.
Orthostixini), subfamilies Oenochrominae, Desmobathrinae, Geo-
metrinae, and Ennominae addenda, by. Jeremy Daniel Holloway.
1996. Malayan Nature Journal 49: 147-326. Published in the
Malayan Nature Journal and also produced in paper covers by
Southdene Sdn. Bhd., Kuala Lumpur, Malaysia. 427 figures, 12 color
plates. Soft cover, sewn binding, 17.9 x 25.3 cm., ISBN: 983-99915-
3-1. Available from Southdene Sdn. Bhd., P.O. Box 10139, 50704
Kuala Lumpur, Malaysia; Phone: 603-4022-2643; FAX: 603-4022-
2267; e-mail: hsbar@pc.jaring.my; website: www.edi.co.uk/barlow;
Price $26.00, £18.00 (including surface mail overseas).
THE Motus oF BorNEO, Part 10, Family Geometridae, subfamilies
Sterrhinae, Larentiinae, and addenda to other subfamilies, by Je-
remy Daniel Holloway. 1997. Malayan Nature Journal 51: 1- 242.
Published in the Malayan Nature Journal and also produced in pa-
per covers by Southdene Sdn. Bhd., Kuala Lumpur, Malaysia. 608
figures, 12 color plates. Soft cover, sewn binding, 17.9 x 25.3 cm.,
ISBN: 983-99915-4-1. Available from Soutitdlone Sdn. Bhd., P.O.
Box 10139, 50704 Kuala Lumpur, Malaysia; Phone: 603-4022- 2643.
FAX: 603-4022-2267; e-mail: hsbar@pc.jaring.my; website:
www.edi.co.uk/barlow. Price $30.00, £20.00 (including surface mail
overseas).
These volumes are the eighth and ninth of an estimated eighteen
part series documenting the macrolepidoptera of the second largest
non-continental island, Borneo. For the family Geometidae alone
this is an ambitious effort. Together with part 11 of the series (En-
nominae, published in 1994), a fauna of 1079 species is treated,
equivalent to approximately one-quarter of the estimated geometrid
VOLUME 54, NUMBER 3
fauna in the Oriental region. The author, Jeremy Holloway, has been
working on the faunistics and biogeography of Indo- Australian Lep-
idoptera for decades and in 1995 was awarded the Karl Jordan
Medal of the Lepidopterists’ Society for his many contributions.
Borneo is an important piece of the lar ge Indo-Australian bio-
geographic puzzle that has fascinated biogeographers since Alfred
Wallace. In a broad sense understanding the fauna of Bomeo re-
quires consideration of taxa that range orn Africa to Oceanea, and
from Australia to the Himalayan region. Holloway’s contribution to
the higher classification of the Caometnies as outlined in this series
extends well beyond the region to the global level as evident in more
recent systematic treatments of the family (see Minet, J. & M. J.
Scoble, 1999, The Drepanoid / Geometroid Assemblage, pp.
301-320, In N. P. Kristensen (ed.), Handbook of Zoology, Volume IV
Arthropoda: Insecta, Part 35 Lepidoptera, Moths and Butterflies,
Volume 1: Evolution, Systematics, and Biogeography, Walter de
Gruyter, Berlin and New York, and Scoble, M. J. (ed.), M. S. Par-
sons, M. R. Honey, L. M. Pitkin, & B. R. Pitkin, 1999, Geometrid
Moths of the World: a Catalogue (Lepidoptera, Geometridae),
CSIRO Publishing and Apollo Boo in association with The Nat-
ural History Minera, London, England; Collingwood, Australia
and Stenstrup, Denmark, 2 volumes). Of the 1079 species of
Geometridae treated in The Moths of Borneo series, 17% are newly
described, 11% contribute new synonymy, and 4% are revived from
synonymy. The lowland forests ‘of Borneo harbor many endemic
species and the upper montane forests are rich in endemic Larenti-
inae, especially the tribe Trichopterygini.
Parts 9 and 10 of The Moths of Borneo are similar in format. The
introduction of part 9 is a world review of the family group names
previously referred to subfamilies Oenochrominae sensu lato and
Geometrinae as well as a discussion of the classification of these sub-
families. The introduction of part 10 includes a synopsis of the di-
versity of Bommean Geometridae with analysis of ecological and bio-
geographic affinities. A brief section offering a tentative
classification of the world Geometridae follows. Family group names
and subfamily classification are reviewed in the systematic accounts
of the Sterrhinae and Larentiinae.
Within the systematic accounts each genus is cited with its au-
thor, type species (including type locality), and synonyms. A generic
description follows based primarily on ‘adult external features and
genitalia. Brief descriptions of the early stages are provided where
known (not illustrated except for a few colon photos in the plates),
larval hosts enumerated, and the geographic distribution of the
genus summarized. Particularly for new or revived genera, extralim-
ital taxa are listed as new combinations. Species accounts include
synonymy, citation of original descriptions, diagnosis, taxonomic
notes, geographical distribution, habitat notes, eal where known, bi-
ology and larval hosts. Recent systematic and biological literature is
often cited and considerable biological information is published
here for the first time. The total number of new taxa published in
parts 9 and 10 attest to Holloway’s effort to document the fauna (22
genera, 108 species, and 3 subspecies). At the conclusion of each
part useful appendices include lists of new taxa, new combinations,
new synonyms, and status changes and revivals. Finally, a checklist
of Bornean taxa is provided and annotated with general distribution
and habitat information. The checklists serve as incliogs to the pages
on which species accounts appear, a general index is not provided.
The extensive literature cited should appeal to the geometrid biblio-
phile.
With the exception of the first four figures of part 9 (line draw-
ings of wing venation) and the first two text figures of part 10, all fig-
ures are photographs of male and female genitalia. Legends are
listed separately in the pages that precede the genitalia figures. The
genitalia photographs vary in size and are arranged in a loose “cut
and paste” format. Figures are not always sequentially numbered.
Quality of the preparations and clarity of the cy oe varies.
Central structures of the male genital capsule and the female
sterigma are not always optimally displayed and aedeagi are often
photographed without vesica inflation. All to frequently the aedea-
gus is not illustrated or only the male genitalia are figured even
103
when female material was available for study. I confess a preference
for line drawings of genitalia but this would be prohibitive in a series
of this magnitude. Nevertheless, selective use of line drawings can
serve to differentiate close taxa as well as illustrate important generic
and family group characters. The color plates of both parts are of
reasonable quality. For smaller moths photographic enlargement
would have enhanced details of maculation. It is difficult to judge
color fidelity without specimens for comparison. Some plates of the
green geometrids in part 9 appear color shifted toward blue or show
reflections (perhaps attributable to scale wear) and many plates of
part 10 are dominated by yellow tints. Specimens depicted on the
color cover of part 10 appear crisper and show better color balance
than photographs of the same specimens on the corresponding
plates.
For the remainder of this review I will overview Holloway’s con-
tribution to geometrid systematics in The Moths of Borneo at the
subfamily, generic, and specific levels.
Present higher classification of the Geometridae is far from sat-
isfactory and requires revision on a world basis. While it is accepted
that traditional groups such as the Oenochrominae sensu lato are
unnatural assemblages (see Minet & Scoble 1999, op. cit.), satisfac-
tory phylogenetic alssilRontiom f is elusive. Holloway’s proposed clas-
sification might best be viewed as provisional, and is based largely on
similarities of adult structures, especially those of the alyalortnen
Holloway appreciates that careful comparative study of the early
stages is needed to shed further light on geometrid systematics.
“Ta the Bornean fauna, only the alu bodied members of Sar-
cinodes Guenée are included in the Oenochrominae sensw sticto.
Holloway revives the subfamily name Desmobathrinae to encom-
pass “delicately built ‘oenochromine’ genera with elongate, slender
appendages.” This subfamil ly aeltnales a number of cosmopolitan
mostly tropical genera placed in the tribe Desmobathrini and the
monobasic Tingle Australian tribe Eumeleini. Two genera, Heteralex
Warren, and Naxa Walker, traditionally referred to Oenochrominae
are tentatively transferred to Ennominae with the latter genus
(along with Orthostixis Hiibner) included in the tribe Orthostixini.
The larvae of Naxa are colonial and known to feed on Oleaceae with
the larvae and pupae suspended in extensive silken webs. In part 10
Holloway publishes new biological information regarding Naxa gut-
tulata Warren observed feeding on a fern (Diplazium sp.); walor
photographs of the larva and pupa suspended in their silken web are
recorded on plate 4. In his review of ‘oenochromine’ family group
names Holloway concludes the Holarctic genus Alsophila Hiibner
should be plac edina separate subfamily, Alsophilinae, a decision ac-
cepted by Minet and Scoble (1999, op. cit.).
The Geometrinae of Indo-Australia include groups quite unfa-
miliar to Lepidopterists of the Northern Hemisphere. Holloway re-
tains the robust bodied aposematic moths of the genus Dysphania
Hiibner within the subfamily but emphasizes the distance of this
unique group from the remainder of the Geometrinae by dividing
the subfamily into two tribes, Dysphaniini and Geometrini. In tum
it is proposed, though with some reservation, that all geometrine
family groups other ian Dysphaniini be assigned as subtribes of
Geometrini and given names ending in “-iti’. Many genera of
Bomean Geometrini are robust Inala moths with falcate forewings
or gray and brown moths with boarmiine facies.
A number of Bornean Sterrhinae should be more familiar to
northern Lepidopterists with the genera Scopula Schrank, Idaea
Treitschke, and Cyclophora rubaer well represented. Holloway re-
views evidence for viewing the sterrhines as two broad lineages
Timandrini / Cosymbinii andl Scopulini / Sterrhini / R hodostrophi-
ini) based on characters of both the adult and early st: ages. He fur-
ther argues for inclusion of the Neotropical tribe Cyll opodini in the
latter lineage, and places the Old World tribe Rhodometrini in the
‘ormer, at least in his phylogenetic diagram (Part 10, Fig. 2)
Given the absence of many recognized tribes of ‘Larontnne in
the tropics, Holloway’s discussion of hi igher classification of this sub-
family is less inclusive. It is suggested that the Trichopterygini may
represent a sister group for ‘inte ‘remainder of the Larentiinae. Four
Australian and Oriental larentiine genera (two newly described by
104
Holloway) cannot be placed in presently recognized tribes indicative
of the unsettled state of higher classification for the subfamily.
On the generic level, Holloway’s revisionary work establishes a
large area Nek of new combinations and many new genera. As
pointed out by Scoble et al. (1999, op.cit.), restricting the definition
of previously lar ge and overly inclusive genera such as the larentiine
genus Chloroc lystis Hiibner based on study of a regional fauna may
Sane unstudied extralimital taxa not corresponding to new generic
concepts. Holloway both revived and erected new genera in his
treatment of Chloroclystis sensu lato and reassigned many Indo-
Australian species. Some reassignments are hasty and seem prema-
ture. For example, eight species are formally transferred to the lar-
entiine genus Bosara Walker without study of the genitalia even
though Holloway admits final placement awaits eontiamelion by dis-
section. A number of unplaced species now reside in the provisional
genus “Chloroclystis” of the Scoble, et al. (1999, op.cit.) catalog.
At the species level, documenting the macro-fauna of a Agere
tropical landmass such as Borneo is daunting but Holloway pulls off
a commando performance. Many factors eeatabate to the difficult
task of determining specific limits including allopatry in the insular
realm of Indo- Australia lack of sufficient study material, ambiguous
association of sexes, and inadequate old descriptions. Holloway ac-
knowledges that some treatments are provisional. To identify species
considerable reliance is placed on brief diagnoses and discussion of
similar species. Keys to genera and species are not provided even
though this would certainly aid in the sorting and recognition of taxa
especially for speciose groups. For many species only the male gen-
italia or the male genital capsule are illustrated. A number of new
species and genera are described in the absence of description and
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
illustration or even mention of the male aedeagus. This frequently
occurs in the green geometrids. Even if the aedeagus lacks diagnos-
tic characters, that fact should be demonstrated.
Access to the enviable store of type material in The Natural His-
tory Museum, London, and other European museums has enabled
Holloway to place many Indo-Australian taxa and recognize unde-
scribed species. Although synonymies are listed for each species,
useful type information such as type locality, repository, condition,
sex, and lectotype designation is frequently lacking. Unless other-
wise noted, holotypes of species described in The Moths of Borneo
are deposited in The Natural History Museum.
Holloway rejects the stipulation of the Code of Zoological
Nomenclature that requires species group names to agree in gender
with genus group names and gives all specific names the orthogra-
phy at the original description. This convention is also adopted in
the Scoble catalog for the geometrid fauna of the world, a conve-
nience that I sincerely hope will become standard practice.
I cannot overstress the ambitious nature of the project Jeremy
Holloway has undertaken. Borneo harbors a geometrid fauna equal
to 77% of the number of species known from America north of Mex-
ico, a fauna better understood as a consequence of Holloway’s stud-
ies. Lepidopterists interested in the faunistics, biogeography, and bi-
ology of Indo-Australian Lepidoptera will want to add The Moths of
Borneo to their library. The parts treating the Geometridae are of
particular value to geometrid specialists worldwide and will inform
and inspire future systematic efforts.
GEORGE J. BALOGH, 6275 Liteolier, Portage, Michigan 49024,
USA.
Date of Issue (Vol. 54, No. 3): 25 September 2001
eho EDITORIAL STAFF OF THE JOURNAL
i ey asses ~M, Deane Bowers, Editor ~
. Entomology Section
University of Colorado Museum
Campus Box 218
University of Colorado
Boulder, Colorado 80309, U.S.A.
ema: bowers@spot. colorado.edu
; Associate Editors:
ee “ Gekarpvo Lamas (Peru), Kener W. Putuir (USA), Roperr K; Rossins (USA),
ade ee Feux A. H. eames Won Davin L. Wacner (USA), Curisvern Wik. OND (Sweden)
- NOTICE TO CONTRIBUTORS
‘to a ae may aca math any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Technical Comments,
bituaries, Feature Photographs, and Cover Illustrations: Obituaries must be authorized by the President of the Society. Requirements
tographis and Cover Illustrations are stated on page 111 in Volume 44(2). Journal submissions should be sent to the editor at the above
m muscripts concerning new state records, current events, and notices should be sent to the News, Phil Schappert, Dept. Zoology, Univ.
nm, TX 78712. Requests to review books for the Journal and book review manuscripts should be sent to M. Alma Solis, Syst. Entomol. Lab,
itional Museum of Natural History, MRC 127, Washington, DC 20560. Journal contributors should submit manuscripts in triplicate,
te ae tirely double-spaced, with wide margins, on one side only of white, letter-sized paper. Prepare manuscripts according to the following
‘submit them flat, not folded. Authors are expected to provide a Ses cay of the final accepted manuscript in a standard PC or
based word proce ing format.
[ informati Pees should precede the text of Aaiales and Profiles.
Ww Up to five key weeds or terms notin ve title should seca Articles, ‘Profiles, General Notes, aad Technical Com-
ntributors Sila write ‘vith precision, ae sak economy, and should use e the active voice and first person siheneyer eters Titles
cit, descriptive, and as short as possible. The first mention of a plant or animal in the text should include the full scientific name with
easurements should be given in metric units; times in terms of the 24-hour clock (0930 h, not 9:30 AM). Underline only where
x yBelerettoes in the text of Articles, Profiles, ‘General Notes, and Technical Comments should be given as Sheppard (1959) or
[a, 1961b) and listed alphabetically under the heading Lirenarure Ciren, in the following format without underlining:
, M. 19 Natural selection and heredity. 2nd ed. Hutchinson, London. 209 pp. '
ee Some contributions to population genetics resulting from the study of the Lepidoptera. Adv. Genet. 10: +165- 216.
ily half of symmetrical objects such as adults with wings spread should be illustrated, unless whole illustration is crucial, Pho-
nd drawin: s should | be mounted on stiff, white backing, arranged in the desired format, allowing (with particular regard to lettering) for re-
ournal page. Illustrations larger than letter-size are not acceptable and should be reduced photographically to that size or smaller. The
and eee numbers as cited i in the text spould be se on the back of each illustration. Figures, both line Sheree and Pees
aged: contact the editor for submission fears and.cost..
ould be numbered consecutively i in Arabic numerals. Headings for tables should not be capitalized. Tabular material must be
ro ets, and placed oe the main text, with the Baapienale desired position indicated in the text. Vertical lines as well as ver-
iould be. avoided. ;
: When appropriate, manuscripts must ‘ame a rpeblic Rs where specimens documenting identity of organisms can be
ports that require Lie include life cneweaeal host associations, immature Bee SS and i uae enquiries.
Fe or authors affliated ovith iaeGentiotis page one are $50 per Aatale page. For srluiels without institutional support, page
21 + Journal page. For authors who are not members of the Society, page charges are $75 per Journal page. Authors unable to pay page
reason should: apply to the editor at Lae time ais subpiission for a reduced rate or free publication. Authors of Book Reviews and
PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A.
-CENpRAL Notes,
Notes on the behavior of Seacae
Kopper, 1 David C. Margoli
log of ‘Adelphia mythra feedi
_ data on cae ore ven
*
i Boox Rens
Number 4
“ISSN 0024-0966
EPIDOPTERISTS”
pee
re)
, Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
“en
Publié par LA SOCIETE DES LEPIDOPTERISTES.
egeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
icado por LA SOCIEDAD DE LOS LEPIDOPTEROLOGOS
ag
Heraus;
THE LEPIDOPTERISTS’ SOCIETY
EXECUTIVE CoUNCIL
Joun W. Brown, President ‘ ... Susan S. Borxin, Vice President
Micuaen J. Smiru, Immediate Past President Mirna M. Casacranng, Vice President
Manuew A, BaucazAr-Lara, Vice President Dawip CG. Irrner, Treasurer
Ernest H. Wittias, Secretary
Members at large:
Ronald L. Rutowski M. Deane Bowers : AS George L. Balogh
Felix A. H. Sperling Ron Leuschner Andrew V. Z. Brower
Andrew D. Warren Michael Toliver Brian Scholtens
EpIroriAL BoaRD
Rosert K. Rossiys (Chairman), Joun W, Brown (Member at large)
M. Deane Bowens (Journal)
WituraM EB. Mituer (Memoirs)
Puiu J. Scuarrert (News)
. Honorary Lire MEMBERS OF THE SocIETY
Cnarzes L. Remincton (1966), E.G. Munroe (1973), IAN EF. B. Common (1987),
Joun G. FrancLemonr (1988), Linconn P. Brower (1990), Doucias C. Fercuson (1990),
Hon. Miriam Rotuscuitp (1991), Craupe Lemame (1992), Frepericxk H. Rinpce (1997)
The object of The Lepidopterists’ Society, which was formed in May 1947 and formally constituted in December 1950; is “to promote the scier
of lepidopterology in all its branches, . . . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas
by both the professional worker and the amateur in the field; to secure cooperation in all measures” directed towards these aims.
Membership in the Society is open to all persons interested in the study of Lepidoptera. All members receive the Jowrnal and the News of The L
idopterists’ Society. Prospective members should send to the Assistant Treasurer full dues for the current year, together with their full name, address,
and special lepidopterological interests, In alternate years a list of members of the Society is issued, with addresses and special interests.
Active members—annual dues $45.00 within the U.S., $50.00 outside the U.S. —
Affiliated members—annual dues $10.00 within the U.S., $15.00 outside the U.S:
Student members—annual dues $20.00 within the U.S., $25.00 outside the U.S.
Sustaining members—annual dues $60.00 within the U.S., $65.00 outside the U.S.
Life members—single sum $1,800.00 : idee
Institutional subscriptions—annual $60,00 haat opera
Airmail postage for the News $15.00 :
Send remittances, payable to The Lepidopterists’ Society, to: Kelly M. Richers, Asst. Treasurer, 9417 Carvalho. Court, Bakersfield, CA 93311; and a
dress changes to: Julian P. Donahue, Natural History Museum, 900 Exposition Blyd., Los Angeles, CA 90007-4057. For information about the Soci-
ety, contact: Emest H. Williams; Department of Biology, Hamilton College, Clinton, NY 13323. To order back issues of the Memoirs, write for avail- ti eh
ability and paces to Kelly M. Richers, Asst. Treasurer, 9417 Carvalho Court, Bakersfield, CA 93311. Ge ete
The additional cost for members outside the U.S, is to cover mailing costs.
Journal of The Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by The Lepidopterists’ Society, 4 Los Angeles Come Musennt of
Natural: History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA and at additional mailing offices.
POSTMASTER: Send address changes to The Lepidopterists’ Society, “/o Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007- 405
Cover illustration: Alesa amesis (Riodinidae), fifth instar, detail of the thorax with an insert showing entire larva. Scale line = 1 mm. This species i is from
Eeuador, Sucumbios, Garza Cocha. Drawing by C, M. Penz. ;
JOURNAL OF
Toe LeEpPIpoPrTERISTs’ SOCIETY
Volume 54
Journal of the Lepidopterists’ Society
54(4), 2000, 107-110
2000 Number 4
IMMATURE STAGES OF THE MARBLED UNDERWING, CATOCALA MARMORATA (NOCTUIDAE)
JOHN W. PEACOCK
185 Benzler Lust Road, Marion, Ohio 43302, USA
LAWRENCE F. GALL
Entomology Division, Peabody Museum of Natural History, Yale University, New Haven, Connecticut 06520, USA
ABSTRACT. The immature stages of C. marmorata are described and illustrated for the first time, along with biological and foodplant notes.
Additional key words: underwing moths, Indiana, life history, Populus heterophylla.
The Marbled Underwing, Catocala marmorata Ed-
wards 1864, is generally an uncommon species whose
present center of distribution is the central and south
central United States east of the Mississippi River
(Fig. 1d). Historically, the range of C. marmorata ex-
tended somewhat farther to the north, as far as south-
ern New England (open circles in Fig. 1d; see Holland
1903, Barnes & McDunnough 1918, Sargent 1976),
but the species has not been recorded from these lo-
calities in the past 50 years, and the reasons for its ap-
parent range contraction remain unknown.
We are not aware of any previously published infor-
mation on the early stages or larval foodplant(s) for C.
marmorata. The foodplant has long been suspected to
be willow (Salix) or poplar (Populus) (Salicaceae),
based on the external morphology and wing pattern of
the adults. In his recent studies on this species, Willis
(1991) was unable to obtain viable ova from numerous
captive females, even from one female that was kept
alive for 81 days. He dissected 40 females that died in
captivity, and concluded that females may not regu-
larly oviposit until September or October, which is six
to eight weeks after their emergence in the field.
Here we report on the successful rearing of C. mar-
morata, from ova deposited by a wild-caught female
from southern Indiana, and offer suggestions on a po-
tential wild larval foodplant for this species.
REARING NOTES
Ova were secured from a worn female C. marmorata
collected at a baited tree at 2300 CST on 11 September
1994, in Point Twp., Posey Co., Indiana. The habitat is
mesic lowland flatwoods, with internal swamps of two
types: (1) buttonbush (Cephalanthus occidentalis L.)
(Rubiaceae), cypress (Taxodium distichum L.
(Richaud)) (Taxodiaceae), and swamp cottonwood
(Populus heterophylla L.); and (2) overcup oak (Quer-
cus lyrata Walt.) (Fagaceae) and swamp cottonwood.
The female was confined in a large grocery bag (17.8 x
30.5 x 43.2 cm) on a shaded porch at outside ambient
temperature (15—30°C). She was offered a 20% sucrose
solution daily on a small piece of sponge; a new sponge
piece with solution was provided every other day. The
female moth lived until 4 October 1994 (24 days), dur-
ing which time she deposited 404 ova on the sides and
bottom of the bag, and on the sponges. Ova were re-
moved daily and placed onto filter paper in empty plas-
tic film containers, and slightly moistened every 2-3
days. Ova were overwintered in the film containers at
ambient outside temperatures (—2 to 10°C) until 23
January 1995, when they were transferred to a refrig-
erator and kept at 5°C until April 1995. Refrigerated
ova were removed and misted lightly once every two
weeks before being returned to cold storage.
108 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fic. 1. Immature stages and distribution of Catocala marmorata. a, lateral view of 5th (last) instar larva. b, closeup of head capsule of 5th in-
star larva. e, dorsal view of egg as revealed by scanning electron microscopy (60x). d, distribution of C. marmorata in North America; open
circles, pre-1950 records; filled circles, post-1950 records; each circle represents a county in which the species has been recorded; shadin
represents distribution of Populus heterophylla (after Little 1977 and McCormac 1993).
o
(=)
On 13 and 27 April 1995, batches of ova were re- of 404 ova survived winter storage to produce larvae.
moved from the refrigerator and held at 22°C. First in- The duration between 5% and 95% total first instar
star larvae began emerging 13-19 days after the ova emergence from a single clutch of eggs, a measure to
were removed from the refrigerator. A total of only 23 assess hatch synchronization as it pertains to life his-
VOLUME 54, NUMBER 4
tory strategies of Catocala (D. F. Schweitzer & L. F.
Gall, unpubl. data), was 4 days for this clutch. Newly
emerged larvae were placed individually in petri
dishes on moistened filter paper, and presented with
fresh leaves of sandbar willow (Salix exigua Nutt.) and
cottonwood (Populus deltoides Bartr.). Although no
precise data were kept, the larvae appeared to eat both
foods equally well, and their development was compa-
rable on both plants, with little apparent difference in
the duration of larval instars and pupae, or adult size.
The larvae were reared indoors at 20-24°C through
the fourth instar using fresh cut foliage that was col-
lected daily. Most of the ultimate (Sth) instar larvae
were placed in fine-mesh nylon sleeves on field grown
willow or cottonwood saplings to complete develop-
ment. Thirteen of the original 23 larvae survived to the
adult stage. The average duration for each stage was as
follows: ova, 15 days; Ist instar, 4 days; 2nd instar, 4.5
days; 3rd instar, 4.7 days; 4th instar, 5 days; 5th instar,
7.6 days; pupa, 26.1 days. A description of the mature
larva is as follows:
oth (final) instar larva. Fig. la, b. Length 9 cm;
head capsule width 4 mm; body color light mousy
brown tinged with pink; dorsal tubercles pink; spira-
cles pinkish brown; finely dispersed black to brown
specks forming interrupted but nearly complete dorsal
and lateral lines along length of body, most apparent in
the vicinity of the dorsal tubercles; 5th abdominal seg-
ment with slightly elevated, 2 mm wide transverse pro-
tuberance, slightly lighter than body color, with a lat-
eral “saddle patch” slightly darker brown than body
color, all lines formed by specks being darker and
more prominent on this segment; Sth abdominal seg-
ment with a pair of ventrally projecting, 2 mm long tu-
bercles; lateral filaments present along entire length of
body, pink, dense, 1-2 mm in length, simple (not bi-
furcate or multifurcate); ventral surface of body seg-
ments dirty pink, with black spots on each abdominal
segment, edged and overlayed with orange on 4th
through 7th abdominal segments; capitad surface of
head capsule flattened but not strongly produced ad-
dorsally, with two 1-2 mm wide, dark, nearly continu-
ous lateral stripes from antennae to dorsal margins; an-
tennae and true legs pinkish; setae on head capsule
and body pinkish, sparse.
On 16 September 1996, two additional female C.
marmorata were captured at the Posey Co. site, and
confined for ova. The protocols for handling these fe-
males and their ova and larvae were largely as de-
scribed above. These two females together laid several
hundred ova, with subsequent first instar hatches again
of only several dozen larvae. Measures of hatch syn-
chronization for first instar larvae from these broods
109
were 6 and 12 days, respectively. Larvae of each brood
were reared indoors successfully on Populus deltoides.
DISCUSSION
It is unlikely that either of the laboratory foodplants,
Populus deltoides and Salix exigua, is the wild larval
foodplant of C. marmorata at the Indiana site. S. ex-
igua was not located anywhere in the area where the
adult females were taken, and P. deltoides occurs only
rarely in the general vicinity, and then not in close
proximity to the female collection site. However, both
black willow, S. nigra Marsh, and swamp cottonwood,
P. heterophylla, occur within 200 m of the female col-
lection site. S. nigra is common and widely distributed
in the eastern United States, as is P. deltoides, and both
of their geographic ranges greatly exceed that of C.
marmorata. The indeterminate foliating schedule of S.
nigra leaves is considerably longer than that of most
Populus species, and the egg hatch synchronization
measures of 4, 6, and 12 for C. marmorata are compa-
rable to Catocala species that are known to use the
more determinate-foliating Populus as opposed to
Salix as their principal wild larval foodplants (D. F.
Schweitzer & L. F. Gall, unpubl. data). During 1995
and 1996, many adult C. marmorata, particularly fe-
males in September, rested in close proximity to P. het-
erophylla trees at the Indiana site, and the overall geo-
graphic range of P. heterophylla better approximates
that of C. marmorata (Fig. 1d; but note this tree is not
present in the central Appalachian Mountains). These
facts suggest that P heterophylla is more plausible
than either S. nigra or P. deltoides as a possible wild
foodplant.
Adult C. marmorata were fairly common at the In-
diana site in 1995, with a total of 18 moths observed in
48 hours, both on baited trees and in bait traps at
night, and on tree trunks during daylight hours. On
warm days, adults were seen resting head up from 0.5
to 2 m above the ground on the trunks of large, gray-
barked trees, especially overcup oak (Quercus lyrata).
These observations accord well with those of Willis
(1991), who collected most of his C. marmorata by
tapping large trees with light-colored bark, including
white oak (Q. alba L.), red oak (Q. rubrum L.), maple
(Acer spp.) (Aceraceae), hickory (Carya spp.) (Juglan-
daceae), white pine (Pinus strobus L.) (Pinaceae) and
tulip poplar (Liriodendron tulipifera L.) (Magnoli-
aceae). Although both Holland (1903) and Sargent
(1976) considered C. marmorata “rare,” and Covell
(1984) considered it “uncommon to rare,” Willis (1991)
is more probably correct in describing the species as
“not as rare as sometimes implied ... but rather...
[with] a somewhat localized distribution.”
110
We suspect our success in obtaining viable ova was
in part the result of utilizing female C. marmorata col-
lected late in the season. It is possible that female C.
marmorata mate many weeks after emergence, and/or
undergo a period of reproductive diapause from the
time of initial emergence in July and August, as is true
for some western willow-poplar feeding Catocala
species (D. C. Hawks, pers. comm.). Irrespective, our
poor success at obtaining hatchable C. marmorata
eggs contrasts sharply with the more typical 90-100%
hatching success with most other Catocala species,
and suggests that at least some aspects of the handling
and/or overwintering biology of C. marmorata remain
incompletely understood.
ACKNOWLEDGMENTS
This study was facilitated by the generous assistance of John Shuey
who introduced JWP to the Indiana site. Both he and Tom Carr
helped with the collection of female moths. Valerie Giles and David
Wagner provided the larval photographs, and John Furlough (Ohio
State Museum of Biological Diversity) identified the Salix exigua.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
LITERATURE CITED
BARNES, W. & J. McCDuNNouGH. 1918. Illustrations of the North
American species of the genus Catocala. Mem. Am. Mus. Nat.
Hist., n.s. 111:1—47.
CovELL, C. V., JR. 1984. A field guide to the moths of eastern
North America. Houghton Mifflin, Boston. 496 pp.
HOLLAND, W. J. 1903. The moth book. A popular guide to a knowl-
edge of the moths of North America. Doubleday, Page & Co.,
New York. 479 pp.
LITTLE, E. L. 1977. Atlas of United States trees. Vol. 4. Minor east-
ern hardwoods. U.S. Dept. Agric. Misc. Publ. No. 1342. Gov-
emment Printing Office, Washington, D.C.
McCormac, J. S. 1993. Swamp cottonwood (Populus heterophylla)
in Ohio. Michigan Botanist 32:35-39.
SARGENT, T. D. 1976. Legion of night. The underwing moths. Univ.
Massachusetts Press, Amherst. 222 pp.
WILLIs, G. D. 1991. Observations of C. marmorata (Noctuidae). J.
Lepid. Soc. 45:373-374.
Received for publication 15 May 2000; revised and accepted 15 De-
cember 2000.
Journal of the Lepidopterists’ Society
54(4), 2000, 111-118
TAXONOMIC CLARIFICATION OF NOTOCELIA ROSAECOLANA (DOUBLEDAY)
AND N. TRIMACULANA (HAWORTH) (TORTRICIDAE)
WILLIAM E. MILLER
Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108 USA
RICHARD L. BROWN
Mississippi Entomological Museum, P. O. Box 9775, Mississippi State, Mississippi 39762 USA
AND
KEVIN R. TUCK
Department of Entomology, Natural History Museum, Cromwell Road, London SW7 5BD, UK
ABSTRACT. The strikingly similar Old World olethreutines Notocelia rosaecolana (Doubleday) and N. trimaculana (Haworth) have long
been differentiated with seven structural and wing-pattern characters. Because these characters are quantitative and difficult to apply, it is unclear
whether one or both taxa were accidentally introduced into North America. We evaluate the seven characters by examining each one (y) relative
to forewing length (x), a surrogate for body size, in a basic sample of 60 specimens of both taxa from two continents, using the allometric equation
y = a(x). All seven characters proved to be body-size dependent and to lack discontinuities necessary for discrete states, thus rendering them di-
agnostically inadequate. Fortunately, a new qualitative diagnostic character emerged from this study: presence or absence of melanic sex scales
near the base of the male hindwing hair pencil. These scales were absent in the putative type of N. trimaculana but present in those of N.
rosaecolana, thus clarifying species identities and showing that N. rosaecolana is the only member of the pair thus far present in North America.
Additional key words: Olethreutinae, allometry, body size, Rosaceae, nitrogen.
Although strikingly similar, the olethreutines Noto-
celia rosaecolana (Doubleday 1850) and N. trimacu-
lana (Haworth 1811) are considered separate entities
in the Old World (Benander 1950, Bentinck & Di-
akonoff 1968, Bradley et al. 1979, Hannemann 1961,
Kuznetsov 1987, Nasu 1980, Obraztsov 1965, Ra-
zowski 1987, Van Deurs 1956). The similarity, which
encompasses genitalia as well as wing pattern, is ac-
knowledged by most authors, with Hannemann (1961)
adding that the taxa cannot be separated with cer-
tainty. Wing patterns (Figs. 1, 2) demonstrate the di-
agnostic difficulty, the illustrated specimens having
been reliably identified by a newly discovered qualita-
tive structural character introduced later in this paper.
The close similarity of these taxa has posed a prob-
lem in North America because it is unclear whether
one or both are invading immigrants. The first Ameri-
can record is Kearfott’s (1910) report from New Jersey
of what he called Eucosma suffusana ({Lienig &]
Zeller), a junior synonym of Notocelia trimaculana.
This and subsequent specimen records were verified
or reported by Brown (1973), Heinrich (1923), and
Procter (1946). Later, Bradley et al. (1979) asserted
that N. rosaecolana is the correct name for the taxon in
North America. Their claim presumably rests on the
fact that the reported larval foodplant in North
America is Rosa (Rosaceae) (Ferguson 1975, Heinrich
1923, Keartott 1910), which is the foodplant of puta-
tive N. rosaecolana in Eurasia. Doubleday (1850) de-
scribed this species from specimens that developed on
Rosa, probably the same specimens Douglas (1849)
reared shortly before. In announcing the discovery of
putative N. rosaecolana in Quebec, Landry (1995)
used the nomenclature of Bradley et al. (1979), but
suggested that both taxa might be present in North
America.
The adult characters purported in the past to differ-
entiate the two taxa are quantitative, and hence poten-
tially ambiguous. These traditional characters consist
of four structural and three wing-pattern traits. The
states of these characters diagnosing N. rosaecolana
and N. trimaculana, respectively, are: (1) strong vs.
weak forewing costal curvature; (2) more vs. less
obliqueness of forewing costal strigulae; (3) more
(5-7) vs. fewer (4-5) pairs of forewing costal strigulae;
(4) male forewing costal fold two-fifths of forewing
length vs. one-half; (5) wide vs. narrow forewing width;
(6) lighter vs. darker forewing coloration; (7) long and
narrow vs. short and broad male socii (Benander 1950,
Bentinck & Diakonoff 1968, Bradley et al. 1979, Han-
nemann 1961, Kuznetsov 1987, Nasu 1980, Razowski
1987, Van Deurs 1956). The N. rosaecolana states of
characters (1), (2), and (6) were noted in the original
description (Doubleday 1850). States of an eighth
quantitative structural character mentioned by
Bentinck and Diakonoff (1968)—thickness of male
valval neck—contradict their illustrations, so this char-
acter is not considered further here.
The above sources give range of wing span or
forewing length for both taxa as body size indicators.
112 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics. 1-7. Notocelia wings and wing details. 1, N. rosaecolana male from St. Johns, Newfoundland (CNC) showing where forewing width
(W) and curvature (C) were measured. Arrow points to barely visible hindwing anal fold containing hair pencil. Melanic sex scales, not visible,
are present near base of the hair pencil. Forewing length 7.7 mm. 2, N. trimaculana male from “England” (Hodgkinson Coll., No. 54698)
(BMNH). Arrow points to barely visible hindwing anal fold containing hair pencil. Melanic sex scales are absent near base of the hair pencil.
Forewing length 6.9 mm. 3, Hair pencil in hindwing anal fold of male N. trimaculana from “Germany” (MEM), with melanic sex scales absent
near the base of the hair pencil. 4, Hindwing anal fold of male N. trimaculana from “Germany” (MEM) with hair pencil removed to show ab-
sence of melanic sex scales. 5, Hindwing anal fold of male N. rosaecolana from Ithaca, NY (MEM) with hair pencil removed to show presence
of melanic sex scales (arrow). Blackness of melanic sex scales is not evident in scanning electron micrographs. 6, Surface of normal scale adja-
cent to hindwing anal fold and hair pencil of male N. rosaecolana from Ithaca, NY (MEM). Bar = 2 microns. 7, Surface of melanic sex scale in
anal fold beneath hair pencil of male N. rosaecolana from Ithaca, NY (MEM). Bar = 2 microns.
There is fairly close agreement in forewing length
among the sources after spans are converted to lengths
by an empirically derived equation (Miller 1977). The
resulting ranges for putative N. trimaculana and puta-
tive N. rosaecolana are, respectively, 6.8—8.3 mm, and
7.3-9.1 mm. The larger body size of N. rosaecolana
was noted in its original description (Doubleday 1850).
None of the seven traditional characters has been
shown to be independent of body size. If characters
are body-size dependent without discontinuities, they
may erroneously appear to assume dichotomous states
at different ends of the body-size spectrum, more so if
they are positively allometric relative to body size. We
examine the seven characters with respect to body size
to decide whether they are sufficient for differentiat-
ing N. rosaecolana and N. trimaculana. Also, we intro-
duce and elucidate a new qualitative structural charac-
ter that for the first time clearly differentiates the taxa.
MATERIALS AND METHODS
We gathered data on the seven traditional charac-
ters from 60 pinned specimens of combined N.
rosaecolana and N. trimaculana, half male and half fe-
male, which are referred to as the basic sample. To
VOLUME 54, NUMBER 4
Wing width
(W) (mm)
3.6
W = 0.336 (F'"°)
r? = 0.79,P <0.01), 60n
3.2
2.8
2.4
Forewing
curvature a
(C) (mm) ¢=0.015 (Fes) ee °
oe °
08 r° =0.60,P <0.01,60n f
000 @ °
0.6
0.4
0.2
Socius length
(So) (mm)
0.4
0.3
0.2 So)-10,0059)(Fo)
r=0.51,P <0.01, 20n
0.1
Costal fold
length (Ff)
(mm) e eo e
0.0819
Ff = 2.368 (F )
7 =0.02,P >0.60,30n
6.0 7.0 8.0 9.0
Forewing length (F) (mm)
IMs}
Fics. 8-11. Relations of structural character measurements to forewing length in 30 females and 19-30 males of combined N. trimaculana-
N. rosaecolana. 8, Wing width. 9, Forewing curvature. 10, Socius length. 11, Costal fold length. Solid circles are males, hollow circles females.
Some points represent more than one observation.
114
avoid accidentally including closely related taxa such as
N. roborana (Denis & Schiffermiiller), we used only
pristine specimens with well preserved wing patterns.
Using standard procedures, we made genitalia prepa-
rations of nearly half of the basic sample. To count
presence and absence of melanic scaling associated
with the male hindwing hair pencil—the new charac-
ter—and to measure respective forewing lengths, we
more than quadrupled the number of males in the ba-
sic sample, creating what is referred to as the aug-
mented sample. Specimens of the basic sample origi-
nated in the U.K., France, Germany, the U.S.
(Connecticut, Maryland, Massachusetts, Michigan,
New Jersey, New York, Pennsylvania, Vermont), and
Canada (Newfoundland, Ontario, Quebec); specimens
of the augmented sample had the same origins plus
Japan. We labeled each specimen of the basic sample
“Voucher, Miller-Brown-Tuck 2001.”
To gather character data from specimens, we mea-
sured dimensions, and categorized wing patterns, as
follows: (1) Forewing width of both sexes in mm at W
in Fig. 1; (2) Forewing curvature of both sexes mea-
sured in mm from costal edge at C in Fig. 1 to the per-
pendicular reference line; (3) Male socius length in
mm from tip to cleft at the uncus; (4) Male forewing
costal fold length in mm; (5) Number of pairs of
strigulae in one forewing of females only, as the costal
fold interfered with counting such strigulae in males;
(6) Maximum slant of female forewing costal strigulae
scored subjectively from 1 for slight to 4 for extreme;
(7) Proportion of white or near white in the forewings
of both sexes estimated subjectively to the nearest 10%.
We measured maximum length of one forewing as a
surrogate for body size, including fringe and excluding
tegula. Forewing length is a sensitive and reliable in-
dex of body size, with body mass in olethreutine adults
increasing approximately as the cube of forewing
length (Miller 1977). In the basic sample, using an oc-
ular micrometer, we measured wing variables to the
nearest 0.08 mm at nominal 10.5x magnification, and
socius length to the nearest 0.02 mm at nominal 45x
magnification. In the augmented sample, we measured
wing length to the nearest 0.5 mm with a ruler.
To examine character measurements (y) relative to
forewing length (x), we used the power or nonlinear
form of the allometric equation, y = a(x’), where a and
b are parameters (Smith 1980). We obtained parame-
ter values using the Quasi-Newton method of estima-
tion for nonlinear models (SYSTAT 1992). Because r?
values in SYSTAT nonlinear output are rounded to one
decimal place (nearest 10%), we obtained more pre-
cise r? values with the exponential regression option of
StatWorks (Rafferty et al. 1985), which produced
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
trend lines virtually identical to those of the allometric
equation. We used absolute male forewing costal-fold
length in the analysis rather than fold/wing length ratio
to avoid the statistical complication of fold length ap-
pearing on both sides of the equation.
Male hindwing anal folds and associated structures
were photographed with a LEO S 360 scanning elec-
tron microscope at an acceleration voltage of 15.0 kV.
Specimen preparation for scanning electron mi-
croscopy followed Adamski and Brown (1987).
Museum and collection abbreviations are as follows:
AMNH, American Museum of Natural History, New
York, NY; BL, collection of B. Landry, Ottawa, ON;
BMNH, Natural History Museum, London, UK;
CAES, Connecticut Agricultural Experiment Station,
New Haven, CT; CMP, Carnegie Museum, Pittsburgh,
PA; CNC, Canadian National Collection, Ottawa, ON;
JDG collection of J. D. Glaser, Baltimore, MD; LDG,
collection of L. D. Gibson, Florence, KY; MEM, Mis-
sissippi Entomological Museum, Mississippi State,
MS; MNHP, Muséum National d’Histoire Naturelle,
Paris, France; UMC, University of Missouri, Colum-
bia, MO; UMSP, University of Minnesota Entomology
Museum, St. Paul, MN; USNM, National Museum of
Natural History, Washington, DC; WDA, Washington
State Dept. of Agriculture, Olympia, WA.
RESULTS
All seven traditional characters proved to be body-
size dependent in their expressions, and to lack dis-
continuities necessary for discrete character states
(Figs. 8-14). The size dependency of three structural
characters—forewing width, forewing curvature, and
male socius length—is direct, with forewing length ex-
plaining from 51 to 79% of character variability (Figs.
8-10). The size dependency of the remaining struc-
tural character, length of male forewing costal fold, is
indirect; fold length is stable at a mean 2.8 mm re-
gardless of forewing length (Fig. 11). Thus relative
fold length is greater at shorter forewing lengths, and
less at longer forewing lengths. Toward the shorter
end at 6.5 mm of forewing length, the costal fold/wing
length ratio is 0.43 (2.8/6.5 = 0.43)—near that for pu-
tative N. trimaculana—whereas toward the longer end
at 8.0 mm of forewing length, the ratio is 0.35 (2.8/8.0
= 0.35)—near that for putative N. rosaecolana. In all
the wing pattern characters, body-size dependency is
direct, with forewing length explaining from 48 to 75%
of character variability (Figs. 12-14). There were no
pronounced sexual differences in characters measured
on both sexes (Figs. 8, 9, 12).
Allometry is evident in several characters. Allometry
refers to the numerical change in one body part or
VOLUME 54, NUMBER 4
Percentage
forewing
white (Wf)
1.726
Wf = 1.199 (F’*°)
50 :
rr =0.50,P <0.01,60n
40
30
20
10
No. strigulae
pairs per
forewing (Sn)
U
Maximum slant of
Strigulae (Ss) ees
(categories) Ss = 0.00102 (F )
3 PF = 0.48,P <0.01,30n
2
1
0
6.0 7.0
1.576
Sn = 0.261 (F)
fP=0.75,P <0.01,30n
8.0 9.0
Forewing length (F) mm
Fics. 12-14. Relations of scale pattern character measurements to forewing length in 30 females and 30 males of combined N. trimaculana-
N. rosaecolana. 12, Percentage forewing white. 13, No. strigulae pairs per forewing. 14, Maximum slant of strigulae. Solid circles are males, hol-
low circles females. Some points represent more than one observation.
character relative to that in overall body size or in an-
other body part or character (Smith 1980). If the expo-
nent b in the allometric equation is >1, positive allom-
etry is indicated; if it is <1, negative allometry is
indicated; and if it is ~1, isometry is indicated. Thus
wing width, with a rounded exponent of 1.1, is essen-
tially isometric (Fig. 8). Forewing curvature, male so-
cius length, and number of pairs of forewing strigulae,
with rounded exponents of 1.9, 2.0, and 1.6, respec-
tively, are positively allometric (Figs. 9, 10, 13). Length
of male costal fold, with a rounded exponent of 0.08, is
negatively allometric (Fig. 11). Percentage forewing
white (Fig. 12) and maximum slant of forewing strigu-
lae (Fig. 14) may also exhibit positive allometry, but
less definitely so because these variables were arbitrar-
ily scaled. Allometry increases the apparent contrast in
character states at opposite ends of the body-size spec-
trum. Incidentally, the visual impact of some of these
relations is reduced because their vertical axes were
compressed for economy of reproduction.
In contrast to the traditional characters, all seven of
which are quantitative, the new diagnostic character
discovered during this study is qualitative. It consists
of melanic scales which are possessed by one taxon but
not the other. When present, these scales occur in a
fold between hindwing vein 3A and the anal wing mar-
gin near the base of a hair pencil lying in the fold (Figs.
3-5). The scales are assumed to have a sexual function.
116
The hair pencils of both species are similar except for
their bases, the base appearing thicker and darker
when the melanic scales are present. Hair pencil col-
oration ranges from brown to black in each species,
perhaps depending on degree of exposure to light and
fading. In the basic sample, the melanic sex scales were
present in 10 males, and absent in the remaining 20.
In both species, ordinary scales on either side of the
fold in which the hair pencil lies exhibit definable fen-
estrae, cross ribs, and scutes on longitudinal ridges
(Fig. 6) and are similar to ordinary wing scales in the
anal and cubital regions of other olethreutines (Brown
& Miller 1983). In contrast, the melanic sex scales are
coated with a substance that can be seen extruding
from some fenestrae (Fig. 7). Similar coatings on
melanic sex scales in males of the olethreutine Cydia
caryana (Fitch) are removable by solvents, and have
been postulated to be accreted glandular scent compo-
nents (Brown & Miller 1983). The melanic sex scales
reported here could be important in isolating males of
one Notocelia species from females of the other.
The ability to separate males based on the new char-
acter enabled us to verify more subtle differences be-
tween the two taxa. We believe the following character
states apply more often than not, and that they may be
useful for separating females until a qualitative diag-
nostic character emerges for them: The forewing pat-
tern is more precisely defined in N. trimaculana, with
the dark markings having more definite boundaries
than in N. rosaecolana; the forewing apex is more
acute in N. trimaculana, and the apical scales more
reddish brown than in N. rosaecolana; and the trans-
verse silvery bars of the forewing ocellus are closer to-
gether in N. trimaculana than in N. rosaecolana.
Based on the augmented sample, the mean, com-
puted standard deviation, and range of forewing
lengths of males with and without melanic sex scales
are, respectively, 7.9 + 0.5 mm (6.5-9 mm) (n = 76)
and 7.3 + 0.5 mm (6-8.5 mm) (n = 59). Despite a
broad overlap, the mean difference, 0.6 mm, is highly
significant (P < 0.001, Student t-test).
In the basic sample, range in forewing length (6.1-8.6
mm) (n = 60) is similar to that published for the com-
bined taxa (6.8-9.1 mm). Range interval of forewing
length (higher range limit minus lower range limit) in
the basic sample (8.6 — 6.1 = 2.5 mm) is likewise similar
to that published for the combined taxa (9.1 — 6.8 = 2.3
mm). In the augmented sample, forewing length range
(6-9 mm) and range interval (9 — 6 = 3 mm) (n = 135)
are similar to the corresponding published values for
the combined taxa (6.8—9.1 mm and 2.3 mm).
The new diagnostic character begs a review of
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
names currently used for the two species. Presence or
absence of melanic sex scales should enable species
identities to be clearly established from the respective
male primary type specimens. Besides trimaculana
Haworth, 1811, and rosaecolana Doubleday, 1850,
the name suffusana Duponchel (1843) is used instead
of trimaculana by some moder authors apparently
because Obraztsov (1965) considered trimaculana of
Haworth to be a misidentification. Further complicat-
ing matters, the name suffusana was known for many
years in collections as “suffusana Kuhlwein.”
Kuhlwein never published the name even though
credited as the author by [Lienig &] Zeller (1846)
who also reported Crataegus as its foodplant. The
name suffusana |Lienig &] Zeller was originally used
for the taxon introduced into North America, but suf-
fusana |Lienig &] Zeller is a junior homonym of suf-
fusana Duponchel. In addition, swffusana [Lienig &]
Zeller is inferred to be a junior synonym of trimacu-
lana Haworth because of the Crataegus rather than
Rosa foodplant, and the liklihood that Duponchel also
used Kuhlwein’s name for his suffusana.
According to Horn et al. (1935-37), the collections
of Haworth and Doubleday are in BMNH, and that of
Duponchel in MNHP. Searches of those collections
did indeed reveal male examples presumably used in
describing the three taxa. Specimens of such vintage
are usually considered types, in the present case a
holotype each for trimaculana Haworth and suffusana
Duponchel, and several syntypes of rosaecolana Dou-
bleday. These examples of trimaculana and suffusana
were found to lack melanic sex scales, while those of
rosaecolana had them.
The present known North American distribution by
states and provinces of the taxon possessing the
melanic sex scales—N. rosaecolana—is Alabama, Con-
necticut, Delaware, Kentucky, Maine, Maryland, Mas-
sachusetts, Michigan, Missouri, New Jersey, New
York, Newfoundland, Nova Scotia, Ohio, Ontario,
Pennsylvania, Quebec, Tennessee, Vermont, Virginia,
Washington state, and West Virginia (AMNH, BL,
CAES, CMP, CNC, JDG, LDG, MEM, UMC, UMSBP,
USNM, WDA). None of our North American male
specimens lacked the melanic sex scales (n = 49).
DISCUSSION
The demonstration that all seven traditional charac-
ters used to distinguish N. rosaecolana and N. trimac-
ulana are governed by body size, and that they lack
discontinuities with change in body size, renders them
inadequate for diagnostic use. Analogous situations
have been demonstrated in other olethreutine genera,
VOLUME 54, NUMBER 4
such as Endothenia (Miller 1983) and Epiblema
(Miller & Pogue 1984). As for the larvae, Swatschek
(1958) does not explicitly differentiate N. rosaecolana
and N. trimaculana, but his key separates them by
head color. MacKay (1959), however, considered head
color a dubious diagnostic character for olethreutine
larvae. Comparisons of forewing length range and
range interval of study specimens with published val-
ues for the two taxa show that body size in the study
specimens is representative.
Were it not for the new qualitative diagnostic char-
acter discovered during this study, results of the analy-
sis of the seven traditional characters would cast doubt
on whether N. rosaecolana and N. trimaculana are in
fact different species. Although the new character dis-
tinguishes males only, it is sufficient to confirm that the
two taxa are indeed separate entities. Females thus far
cannot be reliably diagnosed, but certain wing charac-
ters verified in males enable tentative separation of
some females.
The difference in mean body size between the taxa
may be explainable by foodplant differences. Larvae of
both are said to feed on terminal foliage of their re-
spective foodplants: N. rosaecolana on the shrub genus
Rosa, and N. trimaculana on the arboreal genera
Crataegus, Prunus, and Pyrus (Rosaceae) (Benander
1950, Bradley et al. 1979, Hannemann 1961, Kuz-
netsov 1987, Nasu 1980, Razowski 1987, Swatschek
1958, Van Deurs 1956). These four plant genera differ
in foliar nutrient quality. Nitrogen concentration in
foodplant tissues is positively linked to growth in phy-
tophagous insects (Mattson & Scriber 1987). Concen-
trations of foliar nitrogen in the three arboreal genera
range from 1.7 to 2.9%, with one high outlier at 3.4%
(Blinn & Buckner 1989, Cannon et al. 1960, Chase &
Young 1978, Gerloff et al. 1964, Henry 1972, Vang-Pe-
tersen 1973). In contrast, concentrations of foliar ni-
trogen in the shrub genus Rosa under regimes of culti-
vation and fertilization range from about 3.0 to 5.6%,
with one low outlier at 2.2% (Armitage & Tsujita 1979,
Di Benedetto et al. 1995, Johansson 1979a, 1979b).
Thus the foliage of cultivated Rosa can often be a third
or more richer in nitrogen than the foliage of the arbo-
real foodplant genera. Significantly, Bradley et al.
(1979) state that in the U.K., the larger bodied N.
rosaecolana occurs especially on cultivated roses. In-
terestingly, elevated foliar nitrogen in Rosa may be an
artifact of cultivation because foliar nitrogen reported
for wild Rosa ranges from 1.5 to 2.2% (Henry 1972)—
essentially the same as for the arboreal genera. Thus
the body size difference between the two Notocelia
taxa could also be an artifact.
ILAL7/
Five reared adults available to us approximated body
size expectations. All North American in origin, two
males and two females developed on Rosa (CAES,
USNM). These range in forewing length from 7.0 to 8.2
mm, which places them among the largest two-thirds of
the basic sample. Both males possess the melanic sex
scales of N. rosaecolana. The fifth specimen developed
on Robinia pseudoacacia 1. (Leguminosae) (CAES),
which constitutes a new foodplant record for the N.
rosaecolana-trimaculana group. This adult, which can-
not be identified with certainty because it is a female,
measures 7.4 mm in forewing length, and thus also falls
among the largest two-thirds of the basic sample. Foliar
nitrogen values for Robinia range from 3.1 to 4.0%
(Blinn & Buckner 1989, Day & Monk 1977), which is in
the foliar nitrogen range of cultivated Rosa.
Results of examining the putative types for melanic
sex scales confirm the currently prevailing nomencla-
ture of the two species, while clearly making suffusana
Duponchel a junior synonym of trimaculana Haworth.
The results also show that males of the invading
species in North America are N. rosaecolana rather
than N. trimaculana. Of course, the absence thus far of
N. trimaculana males in North America makes it likely
that North American females of the complex are also
N. rosaecolana.
In conclusion, we would underscore that invalidat-
ing traditional but unreliable diagnostic characters can
have the positive effect of fostering fruitful searches
for new and strong diagnostic characters.
ACKNOWLEDGMENTS
For lending specimens in their care or possession and for other
help, we thank J. S. Miller and E. L. Quinter (AMNH), C. T. Maier
(CAES), J. Minet (MNHP), J. E. Rawlins (CMP), P. T. Dang (CNC),
J. W. Brown (USNM), K. B. Simpson and R. W. Sites (UMC), P. J.
Clausen (UMSP), E. H. LaGasa (WDA), M. Sabourin, B. Landry, J.
D. Glaser, and L. D. Gibson. We also thank J. C. Luhman for help in
translating Polish literature, and S. J. Weller, M. Sabourin, and J. W.
Brown for useful manuscript reviews. Figure 2 is reproduced by
permission of the Trustees of the British Museum, copyright owners
of BMNH specimen images. Scanning electron micrographs were
made by W. Monroe, Mississippi State University Electron Micro-
scope Center.
LITERATURE CITED
ADAMSKI, D. & R. L. BRowN. 1987. A new Nearctic Glyphidocera
with descriptions of all stages (Lepidoptera: Blastobasidae:
Symmocinae). Proc. Entomol. Soc. Washington 89:329-343.
ARMITAGE, A. M. & M. J. Tsujita. 1979. The effect of nitrogen con-
centration and supplemental light on the growth and quality of
‘Caliente’ roses. HortSci. 14:614—-615.
BENANDER, P. 1950. Andra Familjegruppen Vecklarfjirilar, Tortric-
ina. Fjarilar, Lepidoptera. Smafjarilar, Microlepidoptera 10.
Svensk Insektfauna 39. 173 pp.
BENTINCK, G. G. & A. DiAKONOFF. 1968. Nederlandse Bladrollers
(Tortricidae). Monogr. Nederl. Entomol. Vereen. 3, 200 pp. +
99 pls.
118
BLINN, C. R. & E. R. BUCKNER. 1989. Normal foliar nutrient levels
in North American forest trees: a summary. Minn. Agr. Exp.
Sta. Bull. 590. 28 pp.
BRADLEY, J. D., W. G. TREMEWAN & A. SMITH. 1979. British tortri-
coid moths. Tortricidae: Olethreutinae. Ray Society, British
Museum (Natural History), London. 336 pp.
Brown, R. L. 1973. Phylogenetic systematics: its application to the
genus Epiblema (Lepidoptera). M.S. thesis, Univ. of Arkansas,
Fayetteville. 179 pp.
Brown, R. L. & P. R. MILLER. 1983. Studies of Lepidoptera hind-
wings with emphasis on ultrastructure of scales in Cydia
caryana (Fitch) (Tortricidae). Entomography 2:161—295.
CANNON, T. F., L. C. CHapwick & K. W. ReEIscH. 1960. The nutri-
ent element status of some ornamental trees. Proc. Am. Soc.
Hort. Sci. 76:661—666.
Cuase, A. Il. & H. E. YouNc. 1978. Pulping, biomass, and nutrient
studies of woody shrub and shrub sizes of tree species. Maine
Agr. Exp. Sta. Bull. 749. 36 pp.
Day, F. P. & C. D. Monk. 1977. Seasonal nutrient dynamics in the
vegetation on a southem Appalachian watershed. Amer. J. Bot.
64:1126-1139.
D1 BENEDETTO, A., M. BEFUMO, G. Rossi & C. BoscHi. 1993/94
[1995]. Diagnostico de la fertilidad en rosas para corte medi-
ante analisis foliares. Revista Fac. Agron. Univ. Buenos Aires
14:229-234.
DousLebay, H. 1850. Descriptions of lepidopterous insects of the
genera Hypenodes, Eupithecia and Spilonota, recently discov-
ered in Britain. Zoologist 8(Append.):cv—cvi.
Douctas, J. W. 1849. Larvae on the leaves and catkins of sallows,
etc. Zoologist 7:2346.
DuPONCHEL, P. A. J. 1843. Supplément a histoire naturelle des
Lépidoptéres d'Europe. Vol. 4. Méquignon-Marvis, Paris.
Fercuson, D. C. 1975. Host records for Lepidoptera reared in
eastern North America. U.S. Dept. Agr. Tech. Bull. 1521, 49 pp.
GERLOFF, G. C., D. G. Moore & J. T. Curtis. 1964. Mineral con-
tent of native plants of Wisconsin. Wisc. Agr. Exp. Sta. Res.
Rept. 14. 27 pp
HANNEMANN, H. J. 1961. Kleinschmetterlinge oder Microlepi-
doptera. I. Die Wickler (s. str.) (Tortricidae). Die Tierwelt
Deutschlands 48. 233 pp. + 22 pls.
Haworrn, A. H. 1811. Lepidoptera Britannica. . . . Part 3, pp.
377-512.
HEINRICH, C. 1923. Revision of the North American moths of the
subfamily Eucosminae of the family Olethreutidae. U.S. Natl.
Mus. Bull. 123. 298 pp.
Henry, D. G. 1972. Foliar nutrient patterns in the subordinate
vegetation of six Minnesota forest stands. M.S. thesis, Univ. of
Minnesota, Twin Cities. 222 pp. "
Horn, W., I. KAHLE & R. KoRSCHEFSKY. 1935-37. Uber entomolo-
gische Sammlungen, Entomologen & Entomo-Museologie. En-
tomol. Beihefte Berlin-Dahlem. 536 pp.
JOHANSSON, J. 1979a. Leaf composition of flowering shoots from
different greenhouse rose cultivars as influenced by rootstock
and season. Acta Agr. Scand. 29:85-92.
. 1979b. Main effects and interactions of N, P, and K ap-
plied to greenhouse roses. Acta Agr. Scand. 29:191-208.
Jones, F. M. & C. P. Kimpatu. 1943. The Lepidoptera of Nan-
tucket and Martha’s Vineyard Islands, Massachusetts. Nan-
tucket Maria Mitchell Assoc. Publ. 4. 217 pp.
Kearrort, W. D. 1910. Family Tortricidae, pp. 537-552. In
Smith, J. B. (ed.), Annual report of the New Jersey State Mu-
seum including a report of the insects of New Jersey, 1909.
Trenton, New Jersey.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Kuznetsov, V. I. 1987. Family Tortricidae (Olethreutidae, Cochyl-
idae)—tortricid moths, pp. 279-956. In Medvedev, G. S. (ed.),
Keys to the insects of the European part of the USSR. U.S.
Dept. Agr. & [U.S.] National Science Foundation, Washington,
D.C. [Translation. |
LANpDRY, B. 1995. Premiéres mentions de quatre espéces de Lépi-
doptéres au Québec. Fabreries 20:6—-14.
[Lienic, F.] & P.C. ZELLER. 1846. Anmerkungen zu Lienigs Lepi-
dopterologischer Fauna von Livland und Curland. Isis von
Oken 1846:175-302.
MacKay, M. R. 1959. Larvae of the North American Olethreutidae
(Lepidoptera). Canad. Entomol. Suppl. 10. 338 pp.
Mattson, W. J. & J. M. Scriper. 1987. Nutritional ecology of in-
sect folivores of woody plants: nitrogen, water, fiber, and min-
eral considerations, pp. 105-146. In Slansky, F. & J. G. Rod-
rigues (eds.), Nutritional ecology of insects, mites, spiders, and
related invertebrates. Wiley-Interscience, New York.
MILLER, W. E. 1977. Wing measure as a size index in Lepidoptera:
the family Olethreutidae. Ann. Entomol. Soc. Am. 70:253-256.
. 1983. Nearctic Endothenia species: a new synonymy, a
misidentification, and a revised status (Lepidoptera: Tortrici-
dae). Great Lakes Entomol. 16:5-12.
MILLER, W. E. & M. G. PocuE. 1984. Ragweed borer (Lepi-
doptera: Tortricidae: Eucosmini): taxonomic implications of an
allometric analysis of adult characters. Ann. Entomol. Soc. Am.
1-227 — 231"
Nasu, Y. 1980. The Japanese species of the genus Notocelia Hiib-
ner (Lepidoptera: Tortricidae). Tinea 11(4):33-43.
Osraztsov, N.S. 1965. Die Gattungen der palaearktischen Tortri-
cidae. Il. Die Unterfamilie Olethreutinae. 6. Tidj. Entomol.
108:365-387.
POWELL, J. A. 1983. Tortricidae, pp. 31-41. In Hodges, R. W.,, T.
Dominick, D. R. Davis, D. C. Ferguson, J. G. Franclemont, E.
G. Munroe & J. A. Powell (eds.), Check list of the Lepidoptera
of America north of Mexico. E. W. Classey & The Wedge Ento-
mological Research Foundation, London, England.
PROCTER, W. 1946. Biological survey of the Mount Desert region,
7. The insect fauna. Wistar Institute of Anatomy and Biology,
Philadelphia. 566 pp.
RAFFERTY, J., R. NORLING, R. TAMARU, C. MCMaAtH & D. Mor-
GANSTEIN. 1985. StatWorks. Cricket Software, Philadelphia.
98 pp.
eee J. 1987. Motyle (Lepidoptera) Polski VII—Uzupel-
nienia I Eucosmini. Monogr. Fauny Polski 15. 253 pp.
SmitH, R. J. 1980. Rethinking allometry. J. Theor. Biol. 87:97-111.
SWATSCHEK, B. 1958. Die Larvalsystematik der Wickler (Tortrici-
dae und Carposinidae). Abh. Larvalsyst. Insekt. 3, 269 pp.
Akademie Verlag, Berlin.
SYSTAT. 1992. Statistics, version 5.2 ed. SYSTAT Inc. 724 pp.
Van Deurs, W. 1956. Sommerfugle VIII. Viklere. Danmarks
Fauna 61. 292 pp.
VANG-PETERSEN, O. 1973. Leaf analysis I. Content of nutrients in
leaf dry matter in apple, pear, plum, cherry, black currant and
red currant in relation to level of nitrogen, potassium and mag-
nesium. Tids. Plant. 77:393-398. [Danish, English summary. ]
WinTER, W. D., ed. 1993. The Northeast, pp. 46-48. In Season
summary 1992. News Lepid. Soc. 1993(2).
YOUNG, T. W., G. H. SNYDER, F. G. MARTIN & N. C. Haysip. 1973.
Effects of nitrogen, phosphorus, and potassium fertilization on
roses on Oldsmar fine sand. J. Am. Soc. Hort. Sci. 98:109-112.
Received for publication 5 January 1998; revised and accepted 15
February 2001.
Journal of the Lepidopterists’ Society
54(4), 2000, 119-130
THE GENUS HYPOPHYLLA BOISDUVAL, 1836 (RIODINIDAE)
WITH DESCRIPTIONS OF NEW TAXA
CuRTIS JOHN CALLAGHAN
Ay. Suba 130-25 Casa 6, Bogota, Colombia
ABSTRACT. The genus Hypophylla Boisduval, 1836 is reinstated as the genus of a monophyletic group of riodinid butterflies previ-
ously included in the genus Calospila Geyer, 1832. Two new species, Hypophylla caldensis, n. sp. and Hypophylla idae, n. sp. are de-
scribed from Colombia. Additional taxa included in the genus Hypophylla are H. zeurippa Boisduval, 1836; H. lasthenes (Hewitson, 1870),
reinst. stat., n. comb.; H. flora (Staudinger, 1887), n. comb.; H. martia (Godman, 1903), n. comb.; H. sudias (Hewitson, [1858]), n.
comb.; H. sudias callaghani (Constantino and Salazar), 1998, n. comb., n. stat.; and H. argenissa (Stoll, [1790]) n. comb. Notes are pro-
vided on the habitats, distribution, behavior, systematics of individual species, as well as keys for the determination of both males and fe-
males.
Additional key words: Trans-Andean region, México, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, perching be-
havior, morphology.
Over the many years in which I have been studying
the riodinids of the inter-Andean valleys and western
coastal regions of Colombia, I collected a number of
specimens of the genus Calospila, two of which I rec-
ognized as being undescribed taxa. My interest thus
having been aroused, I examined other similar species
of the genus Calospila. My conclusion was that these
taxa form a distinct monophyletic group that belongs
to the genus Hypophylla Boisduval. The purpose of
this review is to 1) re-describe the genus Hypophylla
Boisduval, 2) present morphological and biological in-
formation on each species comprising the genus and
near relatives, 3) describe two new species in the
genus, and 4) provide a key to the identification of
both males and females.
MATERIALS AND METHODS
During course of the study I examined material at
the Museo de la Universidad de Caldas, Manizales,
Colombia, the Musée National d’Histoire Natural,
Paris (MNHN), the Universidad Nacional Autonoma
de México (MZFC), the collections of Dr. Schmidt-
Mumm, Bogota, and Dr. Francisco Delgado of Santi-
ago, Veraguas, Panama (FD) and of myself (CJC). I
studied the relevant type material at the British Mu-
seum (Natural History), and the Humboldt Universi-
tat, Berlin. I examined 167 specimens and made 46
genitalia preparations. Measurements were made with
an ocular micrometer and calipers. Reference to wing
cells and veins follows the Comstock-Needham system
in Miller (1969) and the genitalia terminology is from
Klots (1970). My field trips in Colombia, Costa Rica
and Panama over a 25 year period provided data on
the habitats and adult habits of Hypophylla.
HYPOPHYLLA Boisduval, 1836, reinstated genus
Type species. Hypophylla zeurippa Boisduval, 1836
Description. Male (Figs 26, 27)—Average Fw length = 16.5 mm.
Forewing costa slightly curved to apex, apex slightly falcate, distal
margin slightly convex; hindwing with costa straight, distal margin
rounded. Venation (Fig. 6), gens with 4 radial veins, Rl and R2
branching before, and R3 after the discal cell, and Cul branching
before. On the hindwing Rs branches before and Cul at the end oe
the discal cell, Vein 3A extends to the tornus. Dorsal surface wing
ground color reddish-brown. Forewing costal margin dark brown to
apex, wing dull purple from middle of cell to apex and to tornus, rest
of wing reddish-brown; a reddish-brown line at end of discal cell and
two more inside with two additional black lines in cell Cu2-1A+2A
below; an irregular median band of connected reddish-brown spots
reaches from the costa to Cul, then is displaced basad such that it is
closer to the cell, reaching 1A+2A; a submarginal black band ex-
tends from the apex to the inner margin, widest from costa to M3;
fringe black. Hindwing reddish-brown above 1A+2A and anal mar-
gin light grey, and distal margin from tornus to Cul dark yellow-
orange, fringe black. Some specimens have variable blue scaling at
apex. Merlin includes a black spot on the costa, with a black line
at end and two within the discal cell, and two more below in cell
Cu2-1A+2A, median band of unconnected spots, one on costa
slightly offset basad from next 4 in a straight row to Cul, then two
more offset basad to 1A+2A. Long hair-like scales are scattered in
cell Cu2-2A. Ventral surface ground color dark grey-brown with
black markings. Forewing with black markings as on dorsal surface.
Hindwing pattern same as dorsal surface, but with a faint submar-
ginal band reaching from apex to torus, distad of which is a row of
faint spots between the veins, those at the apex and tornus with
more extensive variable black scaling.
Head, thorax, abdomen dark reddish-brown dorsad, uniform
white scaling ventrad, antennae brown, ventrad with white scales
between sections, club weak; orbit white, frontoclypeus white ven-
trad, with light brown scaling above; labial palpi (Fig. 1) dark brown,
with short, dense scaling, last segment 0.32 of second, protruding
beyond face when oneal dorsally. Forelegs (Fig. 3 ) white, with coxa
thin and wedge-shaped, trochanter branching off slightly more than
half way to the tip, tarsus unimerous; middle (Fig. 4) and hind legs with
a tibial spur and a group of 3 spines on distal tip of tarsal segments.
Genitalia (Fig. 8)—Uncus deeply bifurcated and lobes rounded;
vinculum narrow, widening medially, broad at base, connected to
wide, spade-like saccus; melee as two caudad projecting rounded
processes, connected with a broad transtilla w hich has two long, thin
caudad projecting processes, the right side (looking cephalad) longer;
pedicel squared caudad; bifurcated process on transtilla Fiore
reaching slightly beyond the end of the aedeagus; aedeagus pointed
with one cornuti; posterior edge of 8th sternite slightly bifurcated.
Female (Figs. 28, 29) —Av erage forewing length 15.5 mm.
Forewing height to length 1:1.44. Dewal surface wing ground color
light brown with ventral maculation appearing through. Forewing
with 3.0 mm wide uneven pale yellow band reaching from costa al-
most to tornus, enclosing a variable spot of brown scaling at anal an-
gle, band slightly jagged. at end of cell. Ventral surface forewing api-
al area dark brown, bordered basad by pale yellow area
}
—
=e
Ze
S
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fig. 1. Male palpi. Fig. 2. Female palpi. Fig. 3. Male foreleg. Fig. 4. Male middle leg. Fig.5. Female foreleg. Fig.6. Venation
of Hypophylla.
corresponding to dorsal band, which extends along costa to base and
stops slightly short of tomus. Submedian to base lighter grey-brown
with brown maculation corresponding to that of male. Hindwing grey-
brown, darker on submarginal area with same maculation as male.
Head, thorax and abdomen dark brown dorsad, white scaling
ventrad; frons white basad, labial palpi (Fig. 2) dimorphic, longer
than male, third segment 0.62 of second. Forelegs (Fig. 2) normal.
Genitalia (Fig. 18)—Papillae anales blade-like, rounded, setose,
with a small point between lobes; ostium bursae opening as a wide
funnel with the top folded dorsad and a small sclerotized ridge, duc-
tus seminalis joins at the left side of the base, looking ventrad; cor-
pus bursae with the signa as two slightly invaginated sclerotized
patches.
Systematics: Most of the species proposed for the genus Hy-
pophylla were originally described in the genus Lemonias. In his re-
vision of that genus, Stichel (1911) resurrected the genus Poly-
stichtis (Hubner 1816) for these and other butterflies, with the
genus type being Lemonias parthaon (Dalman 1823). As the name
Polystichtis was invalid as a subjective junior synonym of Emesis
Fabricius, 1807, Hemming (1967) placed these butterflies under the
next available name, Calospila Geyer, 1832.
As defined by Stichel, the genus Calospila includes at least two
morphologically distinct groups; a section of species related to C.
parthaon, the type of Calospila (including C. gygas (Stichel 1911)
and C. zeanger (Stoll 1790)), and part of another section called the
Argenissiforms by Stichel (1911), containing the species in this
study. Hypophylla is related to Calospila, with which it shares vena-
tion (which probably led Stichel to include both under Polystichtis),
sexually dimorphic labial palpi, and the slightly indented eighth ab-
dominal sternite in the male.
Morphological synapomorphies which support the monophyly of
Hypophylla are 1) the long, thin asymmetric posterior projecting
processes of the transtilla of the male genitalia, 2) the branching of
the ductus seminalis from the left side of the base of the ductus bur-
sae of the female genitalia, 3) the formation of the opening of the os-
tium bursae which is folded over as a thin short membrane and with
a variable internal flange, and 4) the signa in the corpus bursae as
two slightly invaginated round sclerotized patches in all but one
species,
In addition to the foregoing, the members of Hypophylla can be
separated from those of Calospila by the general structure of the os-
tium bursae (compare Figs. 17 and 18), the separation of the male
valvae and the male pedicel which is uniform and not inflated at the
base (compare Figs. 7 and 8). Superficially, members of Hypophylla
may be recognized by their generally larger size, dull blue coloration
on the male dorsal forewing and orange-yellow at the tornus and
submarginal areas of the hindwing.
Among the Stichel’s Argenissiforms, there are two that do not be-
long. Calospila candace H. Druce (1904), from southeast Brazil, is a
typical Calospila, as I verified through examination of the male and
female genitalia. The other is Calospila fannia (Godman 1903),
which is likewise a Calospila. (J. Hall, pers. comm.).
The male, and, especially female, genitalia show consistent phe-
notypic variation and are a good basis for the separation of species.
VOLUME 54, NUMBER 4
Important characters in the male include the shape of the uncus,
shape of the valvae and caudad projecting process on the transtilla,
shape of the pedicel, rami and the adeagus. In females, important
characters include the shape of the ostium bursae, position of the
ductus seminalis and the signa in the corpus bursae.
With the changes and new species proposed in this review, the fol-
lowing synonymic list summarizes the classification of Hypophylla:
HYPOPHYLLA Boisduval, 1836
zeurippa zeurippa Boisduval, 1836,
eumedes Doubleday, 1847, nom. nud.
lasthenes (Hewitson, 1870), reinst. stat., n. comb.
martia (Godman, 1903), n. comb.
caldensis Callaghan, n. species
flora (Staudinger, 1887), n. comb.
idea Callaghan, n. species
sudias sudias (Hewitson, [1858]), n. comb.
sudias callaghani (Constantino and Salazar), 1998, n. comb.,
n. stat.
argenissa (Stoll, [1790]), n. comb.
petronius (Fabricius, 1793)
staudingeri (Godman, 1903)
ECOLOGY AND BEHAVIOR
Distribution and habitat. Members of Hy-
pophylla exclusively inhabit the transandean region,
from México (Chihuahua, Guerrero, Chiapas, Oaxaca,
Veracruz) through Central America to Colombia
(Cordillera Oriental, west slope to the Pacific coast)
and western Ecuador. The genus is not recorded from
the eastern (Amazonian) slope of the Cordillera Ori-
ental. It shows its greatest radiation in Colombia, with
six species recorded. Four species are found in Costa
Rica (DeVries 1997) and southern Nicaragua, while
two are found in Mexico, and one in Ecuador (J. Hall,
pers. comm.). The assertion in Seitz (1917) that it is
found in Venezuela needs verification. In the MNHN
in Paris there is a typical H. lasthenes male from “Suri-
nam”, which I suspect is a mislabelling.
The majority of the species inhabit tropical lowland
Tropical Humid Forest and Very Wet Humid forests
(Holdridge 1947) below 900 m. Two species in Colom-
bia are found in Premontane Humid Forest zone in
the Coffee belt, between 900 and 1800 m.
Wing pattern and predation. The males of all
species but one have bright yellow patches on the dis-
tal hindwing to attract predators to that part of their
anatomy, as suggested by captured specimens with
beak-bite marks on the hindwing. Females mimic day
flying moths with a yellow band on the forewing, par-
ticularly Geometridae (Laurentiinae, Sterrhinae) and
Arctiidae (Lithosiinae) (DeVries 1997; pers obs.)
Mating behavior. All species of Hypophylla use
perching as mate locating behavior. Perch localities as
well as male behavior differ among species. The males
of six taxa perch inside woods edges, streams or
treefalls, resting with outspread wings on ventral leaf
surfaces, and three, H. caldensis, H. sudias callaghani
12]
and H. idae I found exclusively on hilltops, resting on
the dorsal leaf surfaces (Fig. 52). When Acta. all
species fly with a rapid, loping flight similar to Ewry-
bia, or a satyrid, returning shortly to their original
perches. When not perching, both sexes rest under
leaves with the wings flat.
SPECIES ACCOUNTS
Hypophylla zeurippa Boisduval, 1836
(Figs. 26, 27, 28, 29)
eumedes Doubleday, 1847, nom. nud.
Hypopylla zeurippa was described from a male from Mexico and
as the type of the genus Hypophylla . Boisduval’s type was discov-
ered in the British Museum (Natural History) by Dr. Gerardo
Lamas. The specimen is similar to the H. zeurippa population from
the Pacific coast (Guerrero), which is distinct from the southern and
gulf coast populations.
Diagnosis. The similarity of the male genitalia place H. zeurippa
in a group comprising H. martia, H. lasthenes and H. caldensis. In
fact, I initially considered these four to be conspecific. However, the
distinct female genitalia and consistently different wing patterns and
coloration, ral | lack of intergrades, as well as H. irene. and H.
martia being sympatric in Costa Rica and Panama, convinced me
otherwise. The males of H. zeurippa are easily distinguished by the
dark orange color of the uniform submarginal band on the hind wing
(in some individuals it may be entirely lacking) coupled with the
weak markings on the ventral surface. The females have a narrow
yellow subapical band on the forewing, a characteristic shared with
H. martia; however they may be separated by the post median row
of spots between Rs and Cul ventral hindwing being in a straight
line. H. zeurippa male specimens from Guerrero have more exten-
sive blue on the forewing, and a wider yellow forewing band on the
females.
Range. Hypophylla zeurippa ranges from Mexico (Colima,
Guerrero, Veracruz) south through Guatemala, Honduras, Belize to
central Nicaragua(?).
Material examined. MEXICO: 1 6 Santa Marta, Oaxaca, 500
m, 1 June 1970 (CJC); 8 dd 12 Candelaria, Loxiche, Oaxaca, 500 m,
10 eles 2 63 same Siecaliey Aug. Oct. 1988 (MZFC); 2 dé
Chacalapill al, wears. Loxiche, Oaxaca, 24 July 1990, (MZFC);
104,799 Las Parotas, Atoyac de Alvarez, Guerrero, 25 Oct. 1985; 11
Sept. 1988, 20 Jan. 1987, 25 Nov. 1985, 4 May 1985, 25 Oct. 1985,
11 Sept. 1985, (MZFC); 8d, 82° Rio Santiago, Atoyac de Alvarez,
Guerrero, 20 March 1987, 10 Sept. 1985, 6 July 1985, 23 Oct. 1985,
9 July 1985, 26 Nov. 1985, 22 July 1984, 23 Oct. 1985, 28 July 1984,
25 July 1984, 5 May 1985, 8 Aug. 1985, (MZFC); 1d Camino a Ton-
ina, Ocosingo, Chiapas, 14 July 1991, (MZFC); 12° Ojo de Agua,
Madrid, Tecoma Colima, 29 Oct. 1982 (MZFC); Baden de Neixpa,
Lazaro Cadenas, Michoacan, 15 Feb. 1996, (MZFC); 1d Metates
San Miguel Aloapam, Oaxaca, Aug. 1991, (MZFC); GUATEMALA:
1d Guatemala, (MNHN).
Hypophylla lasthenes (Hewitson, 1870)
(Figs. SO), Sule Sr, Sis)
Hewitson described Lemonias lasthenes from a male from Chon-
tales, Nicaragua. Godman and Salvin (1886) extended its range to
Costa Rica and Panama, and Stichel (1911) placed this taxon as a
subspecies of Polystichtis zewrippa. Some confusion exists regarding
the type. The specimen in the British Museum (Natural History)
bearing the type label from “Chontales” is typical of southern Mexi-
can specimens of H. zeurippa. Comparison of this specimen with
the original description suggests that it is not the type, especially
with reference to the size ar the orange spot on the dorsal hindwing.
It is possible that both phenotypes occurred at Chontales, but is im-
possible to verify as forest habitat there no longer exists.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fic. 7. Calospila parthaon, male genitalia. Fic. 8. Hypophylla zewrippa, male genitalia. Fic.9. Hypophylla lasthenes, male genitalia.
Fic. 10. Hypophylla martia, male genitalia. Fic. 11. Hypophylla caldensis, male genitalia. Fic. 12. Hypophylla flora, male genitalia.
Diagnosis. A comparison of H. zeurippa and H. lasthenes sug-
gests that they are separate species. Their populations are appar-
ently allopatric with no intergrades, the male genitalia are similar,
but H. lasthenes has more parallel and longer valvae (Fig. 9). The fe-
male genitalia of H. lasthenes (Fig. 19) have the ductus bursae more
swollen and a sclerotized patch opposite the base of the ductus sem-
inalis. Superficially, H. lasthenes males are separated from H.
zeurippa and H. martia by the wider, more rounded and more
brightly colored orange patch on the hindwing that extends from M2
to the inner margin, with a wavy basal edge. The ground color is
lighter, the maculation bolder on the ventral surface, and the fe-
males have a wider, more even forewing band.
Material examined. COSTA RICA: 1d, 299 Chilamate, 100
m, Heredia, 30 June 1992, leg. Callaghan. (CJC); PANAMA: 1,
229 Colén, Puerto Bello, Rio Cuango, 26 April 1991, 1 Aug. 1991
leg. Delgado (FD); 2 4d, 629 Col6n, Coclesito, 50 m, 9 Sept. 1988,
VOLUME 54, NUMBER 4
Fic. 13. Hypophylla idae, male genitalia. Fic. 14. Hypophylla sudias callaghani, male genitalia. Fic. 15. Hypophylla sudias sudias,
male genitalia. Fic. 16. Hypophylla argenissa, male genitalia.
4 Sept.1988, 13 Jan. 1989, 22 Feb. 1989, 18 March 1994, 16 June
1988, leg. Delgado (FD); 1¢ Santa Fé, Belém, 13 March 1987,
leg. Delgado (FD); 12 Alto de Piedra, Santa Fé de Veraguas, 28
Sept. 1989, leg. Delgado, (FD); 1d “Surinam” (MNHN); 1d no
data (MNHN); 12 Panamé, (MNHN); 12 Chiriqui, Panama
(MNHN).
Range and habits. This species is recorded only from northern
Costa Rica to the Caribbean coast in Panama. DeVries (1997) re-
ports this phenotype from Costa Rica on both Atlantic and Pacific
slopes near sea level in mangrove habitats. My experience with this
butterfly in Costa Rica is in Very Humid Tropical Rainforest
(Holdridge 1947) on the edge on a small hilltop near Chilamate,
Heredia (100 m). Two females were captured at 1000 hours resting
with wings outspread on the underside of leaves. At 1230, a male
was taken on the same hilltop likewise resting under a leaf with
wings spread. It is rare in Costa Rica, more common in Panama.
Hypophylla martia (Godman, 1903), n. comb.
(Figs. 34, 35, 36, 37)
This species was described from a male originating from San
Pablo on the Rio San Juan, Chocé, Colombia. The type is in the
British Museum (Natural History).
Diagnosis. H. martia males resemble H. zeurippa in the extent
of blue on the forewing and the uniform dark orange submarginal
band along the distal margin of the hindwing, reaching from the tor-
nus to M2. H. martia may be separated by ane much darker macula-
tion of both wing surfaces, Picci thinner submarginal line on the
rene apex, and more pointed forewing. The male genitalia (Fig.
10) differ in the shorter, thinner valvae and a more truncated
process on the transtilla. The female forewing band, while approxi-
mately the same width, is sharper on H. martia, is not bordered
basad by the end of the discal cell and terminates on the distal mar-
gin. The ductus bursae (Fig. 20) is wider in H. martia and dilated at
the base of the ductus seminalis.
Range and habits. This species ranges from Costa Rica to the
west coast of Colombia. In Costa Rica, there is one record for this
species from Magsaysay in the lowland Atlantic rainforest (DeVries
1997). This species is very rare in Colombia, the only specimen I
have seen from there is the type. It appears to be more common in
Panaméa, where it is sympatric with H. lasthenes. From my data, the
habitat of this species is lowland tropical rain forest.
Material examined. PANAMA: 1¢ es 5 Jan. ne
(CJC); 1d same locality, 16 Dec. 1979, (CJC); 1¢ Los Rios,
Jan.1971, leg. King ( Ie 12 Barro Colorado aay July 13, ica
leg. R. Robbins ( (CJC); 16, 22° Chepo Island, Majé, March 1989 leg.
124
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fic. 17. Calospila parthaon, female genitalia. Fic. 18. _Hypophylla zeurippa, female genitalia. Fic. 19. _Hypophylla lasthenes, female geni-
talia. Fic. 20. Hypophylla martia, female genitalia. Fic. 21. Hypophylla flora, female genitalia. Fic. 22. Hypophylla idae, female genitalia.
Fic. 23. Hypophylla caldensis, female genitalia. Fic. 24. _Hypophylla sudias, female genitalia. Fic. 25. Hypophylla argenissa, female genitalia..
Delgado, (FD); 24, 392 Colén, Coclesito, 23 Jan. 1988, 15 April
1990, 26 Aug. 1988 leg. Delgado, (FD); 1d, 292 Colon, Puerto Bello,
Rio Cuango, 26 April 1991, leg. Delgado (FD).
Hypophylla caldensis Callaghan, new sp.
(Figs. 38, 39, 40, 41)
Description. Male (Figs. 38, 39)—Forewing length of Holotype
16.5 mm and paratypes 16.2 mm. Forewing costa slightly curved to
apex, apex rounded, distal margin slightly convex; hindwing with
costa straight, distal margin rounded. Forewing costa dark brown to
apex; dull, light purple from end of discal cell to apex and to tornus,
rest of wing reddish-brown; a reddish-brown line at end of discal cell
and two more inside, two additional black lines in cell Cu2-1A+2A
below; a median band of connected dark brown spots reaches from
the costa to Cul, then is displaced basad such that it forms a contin-
uous band with the end of the cell to 1A+2A; a marginal black band
extends from the apex to the inner margin, widest from costa to M3;
fringe black. Hindwing costa above M1 to apex dark brown with two
black spots above discal cell and faint irregular blue spot just before
VOLUME 54, NUMBER 4
apex, base to median above 1A+2A reddish brown and anal margin
light grey, postmedial area and distal margins yellow-orange, fringe
black. Discal cell includes a black line at end and two within, and
two below in cell Cu2—1A+2A, median band of unconnected, bold
spots, one on costa slightly offset distad from next 4 in a straight row
to yellow-orange area. Ventral surface ground color dirty white with
black markings. Forewing slightly darker at apex and margin, black
markings as on dorsal surface. Hindwing maculation as on dorsal
surface, and yellow-orange area same, but paler, with a darker mar-
ginal area containing a row of variable marginal black spots, those at
86 - as Or
Fics. 26, 27. Hypophylla zeurippa, male, dorsal/ventral view. Fics. 28, 29. _Hypophylla zeurippa, female, dorsal/ventral view. Fics. 30,
31. Hypophylla lasthenes, male, dorsal/ventral view. Fics. 32, 33. Hypophylla lasthenes, female, dorsal/ventral view. Fics. 34,35. Hy-
pophylla martia, male, dorsal/ventral view. Fics. 36,37. Hypophylla martia, female, dorsal/ventral view.
anal angle and two at apex larger; a post median row of spots contin-
ues below Cul, offset basad and nearly parallel to those above.
Head, thorax, abdomen dark reddish-brown dorsad, uniform
white scaling ventrad, antennae brown dorsad, with white scales
ventrad between sections, club long, weak; orbit white, fronto-
clypeus white ventrad, with light brown scaling above; labial palpi
scaled, protruding beyond face when viewed dorsally.
Genitalia (Fig. 10)—uncus bilobed and rounded, valvae rounded,
narrower and straighter than H. zeurippa, process of transtilla
slightly longer and thinner, posterior edge of Sth sternite slightly bi-
126
furcated, aedeagus pointed with unsclerotized sheath, and a wedge-
shaped comuti.
Female (Figs. 40, 41)—forewing length 15.5 mm. Dorsal wing
ground color “light brown with Rental maculation appearing
through. Forewing with a 3 mm wide uneven pale yellow band from
costa almost to AonntG, enclosing a variable spot of brown scaling at
anal angle, band slightly jagged at end of cell. Ventral forewing api-
cal area dark brown, bordered basad by pale yellow area reflecting
the dorsal band, base light grey with indistinct brown maculation
corresponding to that of male. Hindwing light grey with dark brown
apex and brown maculation as on male. Head, thorax and abdomen
dark brown dorsad, white scaling ventrad; frons white basad, labial
palpi sexually dimorphic, third segment longer than on male.
Genitalia (Fig. 23)—ostium imran as a long, sclerotized tube,
slightly wider at opening where the rim is folded dorsad, and with a
high, sclerotized flange in the middle dorsad; base of tube very
broad where the ductus seminalis separates; corpus bursae elon-
gated with two well developed, invaginated, pointed signa.
Types. Holotype male COLOMBIA: Cerro Aguacatal, Rio Sticio,
1600 m, Caldas 21 March 1997 leg. Callaghan. Par. atypes. 3dd same
data as holotype, and 2d¢ same locality, 30 April 1994 leg. J. Salazar
(CJC), and 1¢ 12 km west of Otanché, Boyaca, 700 m (ex collection
E. Schmidt-Mumm), and 1¢ Rio Cali, Valle, 1100 m, 12 October,
1981 leg. Callaghan (CJC). 12 Aguas Claras 100 m, Rio Anchicaya, 6
June 1982, leg. Callaghan, (CJC); 1d Caucathal, 1000 m, (MNHN).
The ee is deposited i in the Museum of the Universidad Na-
cional , Bogota, and paratypes in the author's collection and the
NMNH, W: agitation D.C.
Etymology. The species is named after the Department in
Colombia where the type locality is located.
Diagnosis. The males of H. caldensis differ from H. zeurippa
and H. martia in the greater extent and lighter yellow color of the
distal half of the diene hindwing and the lighter g eround color and
darker maculation on the ventral surface, and om H. lasthenes by
the expanded yellow on the hindwing. On the ventral forewing, the
post median spots below Cul form a continuous line with the end of
the discal cell. The ventral surface is less strongly marked than H.
lasthenes. The genitalia are very close to H. zeurippa but with
slightly shorter valvae.
A female captured at Aguas Claras (Tatabro) Choc6, 100 m is
tentatively described as the ‘Gomalle of this species. The unique mor-
phology of the genitalia (Fig. 23), in particular the fully developed
invaginated sclerotized signae instead of the sclerotized patch does
not neocae it with any other species. It shares certain characters
with H. caldensis males, such as a spot on the tornus of the forewing
and the beginning of the medial band in line with the end of the cell.
However, the specimen was captured in a completely different bio-
type from the males that are not known from the Chocé. Therefore,
until more material can be examined, the specimen will be main-
tained as the female of H. caldensis.
Range and habits. H. caldensis ranges from 700 to 1800 m in
the Cauca and Magdalena river drainage of Colombia, and possibly
to the Chocé. Its habitat is in the coffee zone characterized as Pre-
montane very humid forest with rainfall between 2000 and 4000
mm. (Espial & Montenegro 1977). The species is probably more
widespread than suggested by the few records known. Males perch
on hilltops from 1300 to 1500, where they fly with a bouncing flight
like a small satyrid, resting on leaf dorsal ‘surfaces 1-2 meters above
the ground (Fig. 52). The Rio Cali specimen was found in the woods
by tiie river at 1130.
Hypophylla flora (Staudinger, 1887), n. comb.
(Figs. 42, 43, 44, 45)
Lemonias flora was described from a male captured near the Rio
San Juan, Chocé, in western Colombia. The type is in the Humboldt
Museum, Berlin.
Diagnosis. The male of H. flora can be easily separated from its
congeners by the shape of the orange-yellow spot on the distal half of
the hindwing, which is rounded at the tornus and culminates in a point
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
at M1; ventral surface dark grey, markings indistinct, apex of ventral
hindwing has three black triangular marks bordered basad with white
between the veins. The male genitalia (Fig. 12) has an uncus similar to
H. zeurippa, but the valvae are extremely truncated. The aedeagus ex-
tends far beyond end of valvae, and is same length as left process of the
transtilla, only slightly shorter than the right one, The female has a 4
mm. wide yellow band reaching from the costa to the distal margin.
The band of H. idae is likewise wide, but ends on the inner margin. In
the female genitalia (Fig. 21), the ductus bursae is a narrow sclerotized
tube, wider in the middle and slightly wider at the ostium bursae
where the rim is folded dorsad, with a high dorsal sclerotized flange in
the middle reaching above the rim. The ductus bursae is slightly
broader where the ductus seminalis separates; the signa in the corpus
bursae are two slightly invaginated sclerotized patches.
Range and habits. Hypophylla flora is known from the Chocé
region on the Colombian west coast to western Ecuador (Tinalandia,
Paramba, Palmar, in northwest Ecuador. J. Hall, pers. comm.), from
sea level to 300 m. Seitz’ (1917) assertion that it ranges to Venezuela
was possibly based on mislabelled material. I have observed this
species perching on the underside of leaves along forest trails and
along streams in primary rainforest. It is rare.
Material examined. COLOMBIA: 4d¢ Rio Tatabro (Aguas
Claras), Rio Anchicayd, Valle, 100-200 m 11 Nov. 1989, leg.
Callaghan (CJC); 12 Rio Tatabro, Valle, July 1992, leg. Salazar.
(CJC); 243 Caucathal, 1000 m (MNHN).
Hypophylla idae Callaghan, new species
(Figs. 46, 47, 48, 49)
Description. Male (Figs 46, 47)—forewing length of holotype
and paratypes 16.7 mm (N = 5). Forewing costa slightly curved to
apex, apex pointed, distal margin slightly convex, hindwing with
costa straight, distal margin rounded. Dorsal wing ground color red-
dish brown with dull purple and yellow-orange scaling. Forewing
discal cell with orange scaling and black bars at end, middle and
base, and two more cell Cu2-1A+2A below; costa and post medial
area above M3 and distad of cell end to apex and to tormus dull pur-
ple, with a row of faint marginal spots; base to slightly beyond end of
cell below M3 reddish-brown with some orange scaling distad, and
bordered distad with a black bar; postmedian area between M3 and
1A+2A with a yellow-orange spot; below 1A+2A to base reddish-
brown, fringe black. Hindwing base, discal area reddish-brown with
black markings at end, middle and base of discal cell, two black spots
on costal margin and a medial row of spots between RS and 1A+2A,
the first four in a straight line, the last two offset basad; apex and
fringe to tornus black, postmedial area to margin yellow-orange;
fringe black. Ventral surface ground color dark grey, maculation as
on dorsal surface, dorsal yellow-orange areas ventrally light pink.
Forewing with marginal row of black spots between the veins bor-
dered basad with white, that continues along the hindwing margin,
the two spots at the apex being the largest.
Head, thorax and abdomen reddish-brown dorsad, mixed white-
brown scaling ventrad, appendages white, antennae brown dorsad,
ventrad with white scales between sections, club weak, orbit white,
frontoclypeus brown; palpi scaled, protruding beyond face when
viewed dorsally.
Genitalia (Fig. 13)—uncus deeply bifurcated, lobes squared; vincu-
lum thin, slightly wider in middle; saccus rounded, valvae tips rounded,
transtilla with two very long, thin processes, left slightly shorter than
right; aedeagus long, pointed with tip tumed up, and internal cornuti;
pedicel rounded, posterior margin of 8th stemite slightly bifurcated.
Female (Figs. 48, 49)—forewing length 16.8 mm. Forewing elon-
gated, height to length 1:1.52. Dorsal wing ground color light brown
oath ventral maculation appearing faintly. F orewing with 5 mm wide
uniform yellow band from costa to just before tornus and inner angle,
enclosing a tiny spot at tornus. Ventral forewing apical area dark
brown with two rows of faint white marginal spots and bordered basad
by pale yellow area corresponding to dorsal band, base light grey with
indistinct brown maculation corresponding to that of male. Hindwing
light grey with dark brown apex and brown maculation as on male.
VOLUME 54, NUMBER 4
AB oe 49
Fics. 38, 39. Hypophylla caldensis, Holotype male, dorsal/ventral view. Fics. 40, 41. Hypophylla caldensis, female, dorsal/ventral view
Fics. 42, 43. Hypophylla flora, male, dorsal/ventral view. Fics. 44, 45. Hypophylla flora, female, dorsal/ventral view. Fics. 46, 47. Hy-
pophylla idaii, Holotype male, dorsal/ventral view. Fics. 48, 49. Hypophylla idae, female, dorsal/ventral view.
Head, thorax and abdomen dark brown dorsad, white scaling
ventrad; palpi sexually dimorphic, longer than male.
Genitalia (Fig. 22) - Ductus bursae as a narrow, long funnel, nar-
rowing slightly at the pointed opening, with the top doubled over
and a small internal sclerotized flange, ductus seminalis joining at
the left side of base; the signa on the corpus bursae are represented
by two very weakly sclerotized patches.
Types. Holotype male: COLOMBIA, Quebrada Valle Sol, km.
104, Bogoté-Medellin, Antioquia, leg. Keith S. Brown Jr. Paratypes.
12 with same data as holotype; 12 Rio Calima (Calima Dam) 800 m,
Valle, 27 Oct. 1982, leg. C. Callaghan, (CJC); 244, 12 Rio Claro 700
m, 5°50/N 74°52’W, Antioquia, Nov. 4, 1989, leg. Callaghan, (CJC);
1¢ Rio Calderas, 700 m Cocorua, Antioquia, Jan. 4, 1996, leg.
Salazar, (CJC). The Holotype is deposited in the Museo de la Uni-
versidad Nacional, Bogota, and paratypes are deposited in the au-
thor’s collection and the NMNH, Washington DC, USA.
Etymology. This lovely species is named for my wife, whom I
met near the type locality.
Diagnosis. Male H. idae are easily separated from its cogeners
by the orange spot at the postmedian area of the forewing. The gen-
128
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics. 50,51. Hypophylla sudias callaghani, male, dorsal/ventral view. Fic. 52. Hypophylla caldensis male perching on hilltop, Rio Sticio,
Caldas. Fics. 53,54. Hypophylla sudias sudias, male, dorsal/ventral view. Fics. 55,56. Hypophylla sudias sudias, female, dorsal/ventral
view. Fics. 57,58. Hypophylla argenissa, male, dorsal/ventral view. Fics. 59,60. Hypophylla argenissa, female, dorsal/ventral view.
italia differs from H. zeurippa in the squared lobes of the uncus,
longer, thinner process on the transtilla and rounded pedicel. The
female differs in the elongated forewing and 5 mm broad forewing
band that terminates mainly on the inner margin instead of the dis-
tal margin.
Range and habits. The type locality is a disturbed forest hilltop
off the Medellin-Bogoté highway in the Central Cordillera. The veg-
etation is Pre-montane very humid forest and Very humid tropical
forest (Espial & Montenegro 1977). All male specimens were ob-
served perching on hilltops between 1100 and 1200 on the upper
surfaces of leaves with their wings slightly raised. The females were
discovered in nearby woods.
Hypophylla sudias sudias (Hewitson, [1858]), n. comb.
(Figs. 53, 54, 55, 56)
Lemonias sudias was described from a male from Honduras. The
holotype is located in the British Museum (Natural History).
VOLUME 54, NUMBER 4
Diagnosis. The males of H. sudias may be easily separated by
the purple scaling covering the dorsal forewing to Cu2—1A+2A with
the postmedian area of the hindwing yellow-orange reaching from
M3 to the tornus with yellow fringe, and the slightly falcate apex of
the forewing. The ventral surface ground color is grey-brown and
the black markings are weak, not appearing above vein Cul; the yel-
low band appears on the ventral surface and the fringe is yellow.
Some specimens from the gulf coast of México (Popocatéptl, Vera
Cruz) have some yellow scaling on the tornus of the dorsal forewing,
similar to H_. s. callaghani (see below). Specimens form Panama dif-
fer in having a uniform marginal black band on the forewing and
heavier markings on the ventral surface, intermediate to Hs.
callaghani. In the male genitalia (Fig. 15), the uncus is deeply bifur-
cated and the lobes pointed; the valvae are only slightly shorter than
H. zeurippa and the aedeagus is the same length as the right process
of the transtilla. H. sudias females are distinguished by a 5 mm wide
white band on the forewing which reaches from the costa nearly to
the distal margin, then curves into the inner margin narrowing to a
point. In the female genitalia (Fig. 24), the ductus bursae is a short,
wide sclerotized tube, slightly wider at the ostium bursae opening
where the rim is folded dorsad, with a deep V-shaped notch in the
rim, the sclerotized flange in the dorsal middle is reduced, not
reaching the rim. The ductus bursae is only slightly broader where
the ductus seminalis separates; the signa in the corpus bursae are
two barely sclerotized patches.
Range and habits. H. sudias sudias ranges from northern
México (Chihuahua) down the Gulf Coast to Oaxaca and Chiapas,
then south to Panama, inhabiting lowland tropical forest. There is
one record in the Allyn Museum of Entomology from San Quintin,
BCN which needs verification. The species is known from Costa
Rica from a single specimen (DeVries 1997). J. Hall (pers. comm.)
reports H. sudias males at Lake Petén, Guatemala perching one
meter high under leaves inside the forest edge at 1630.
Material examined. MEXICO 1¢, 1° Chiapas, San Filipe, 3000”,
Pasta, 6 Aug. 1974. leg. R.Wind, (CJC); 1d, 12 Monumento Natural
Yaxachilin, Ocosingo, 5 Abr. 1998, Sept. 1948 (MZFC); 24d, 39
Ocosingo, Chiapas, Sept. 1948, June 1940, (MZFC); 1d Sierra de
Juarez, Oaxaca, July 1982, (MZFC); 1d San Juan Chamula Chiapas, 19
July 1993, (MZFC); 1d Cerro del Coconé, Teapa, Tabasco, 20 April
1995, (MZFC); Ojo de Agua, Michoacan, 17 Nov. 1981 (MZFC); 8d,
2° Puerto eligio, Oaxaca, 6 Oct. 1986, 12 Aug.1986, 6 Oct. 1986, 31
Oct. 1987, 11 May, 1980, 10 Aug. 1986 (MZFC); 465 30 July 1982
(MZFC); 34, 82 Popoctépetl, San Andres Tuxtla, Veracruz, July 1982,
24 Sept. 1982, 9 July 1982, 20 Aug. 1982, (MZFC); 1d, 12 Tapalapan, 5
Aug. 1982, (MZFC); 1° Santa Rosa Comitan,Chiapas, Oct. 1982
(MZFC); 39° Metates, San Miguel, Aloapam, Oaxaca, 16 Sept. 1987,
22 July 1979 (MZFC); GUATEMALA: 1d Las Cajas, El Petén, 250 m,
(CJC); PANAMA. 1° Alto de Piedra, Santa Fé de Veraguas 840 m,
23 Feb.1987 (FD).
Hypophylla sudias callaghani
(Constantino & Salazar, 1998), n. comb., n. stat.
(Figs. 50, 51)
The male of H. sudias callaghani was described from Tatabro, Rio
Anchicaya, 100 m, Chocé, Colombia, as a species, Calospila callaghani.
The Holotype is deposited in the Museo de Historia Natural de la Uni-
versidad de Caldas, Manizales. Examination of the phenotype and the
female suggests, however, that it is a subspecies of H. sudias.
Diagnosis. Males are easily distinguished from the nominate
subspecies and other Hypophylla by the yellow-orange spot at the
tornus of the forewing combined with the heavy maculation on both
wing surfaces. The male genitala (Fig. 14) of the two subspecies are
slightly different, with the uncus of H. sudias callaghani more
pointed, and the valvae and processes of the transtilla longer and
narrower. The female of H. sudias callaghani is illustrated in D’Abr-
era (1994) on page 1032 from a specimen in the BMNH from the
Rio San Juan, Chocé, misidentified as the female of H. argenissa.
The females differ from the nominate subspecies in the slightly
more truncated white band on the forewing, and appear to be very
rare.
Range and habits. The habitat of this subspecies is coastal trop-
ical rain forest in the Chocé region of western Colombia. Males can
be encountered rarely on hilltops after 1530 where they fly in circles
with a bouncing flight like a small satyrid, resting on leaf dorsal sur-
faces 2-3 meters above the ground.
Material examined. COLOMBIA: Id Quebrada Bendicién, Valle
7 July 1987 leg. Keith Brown Jr. (CJC); 263 Aguas Claras, Rio An-
chicaya, 100 m, 16 June 1982 leg. Callaghan (CJC); 1d Aguas Claras,
Jan. 17, 1982, leg. Callaghan (CJC); 1d Quibd6, Chocé, June 17, 1993,
leg. K.B.Brown Jr.
Note: The locality Aguas Claras is also known as Tatabro, located
on the Rio Anchicayé east of Buenaventura.
Hypophylla argenissa (Stoll, [1790]), n. comb.
(Figs. 57, 58, 59, 60)
=petronius (Fabricius, 1793) (Hesperia)
=staudingeri (Godman, 1903) (Lemonias)
Stoll described Papilio argenissa from material supposedly from
Surinam. This may be in error, as to my knowledge this species has
not been recorded east of the Cordillera Occidental in Colombia.
This led Godman (1903) to describe Lemonias staudingeri from a
misidentified figure based on a western Colombian specimen in
Staudinger’s Exotische Schmetterlinge, which was in fact P. ar-
genissa. The type of L. staudingeri is in BMNH. I was unable to lo-
cate Stoll’s type during my searches at the museum at Leiden, The
Netherlands, so it is apparently lost. However, the distinct morphol-
ogy of this butterfly precludes the necessity of designating a neotype.
Diagnosis. This species can be confused with no other. The
ground color of the male dorsal surface is uniform dark blue with
black markings. On the ventral surface, the ground color is dark
brown with black markings and infusion of white scaling on the
hindwing. In the male genitalia (Fig. 16) the uncus is bifurcated with
a narrow notch and the lobes are pointed, the valvae are long and
narrow, and the adeagus contains two cornuti. The females are easily
distinguished from other members of the genus by the yellow
forewing band with a straight basal border and a distal border that is
curved towards the tornus. In the female genitalia (Fig. 25) the duc-
tus bursae is a long, narrow, funnel shaped, sclerotized tube, bul-
bous where the ductus seminalis separates and wider at the ostium
bursae where the rim is folded dorsad, with a minuscule, sclerotized
flange in the middle and a U-shaped notch; the signa in the corpus
bursae are two elongated sclerotized patches.
Range and habits. Hypophylla argenissa ranges from Costa Rica
to Colombia, where it inhabits west coast (Choc6) from sea level to
about 300 m. In Panama and Costa Rica, this species is found on
both Pacific and Atlantic slopes to 1000 m (DeVries 1997, pers. obs.).
Both sexes perch on hilltops, in treefalls and on forest edges from
1100 to 1500, resting on ventral leaf surfaces with wings outspread.
Material examined. COLOMBIA: 2d¢ Aguas Claras, Rio An-
chicaya, Valle, 17 Jan. 1982, leg Callaghan (CJC); 1° same locality,
July 1990 e Salazar (CJC); 12 same locality, 1 Aug. 1981, leg.
Callaghan (CJC); 12 same locality, May 24, 1982, leg. Callaghan
(CJC); 24d, 12 same locality, 20 Feb.1982, leg. Callaghan (CJC);
PANAMA: 1° Canal Zone, Gattin, 26 April 1989, leg. Callaghan
(CJC); 1d Colon (Santa Rita) 500 m, 7 Jan. 1975, leg. Nicolay (CJC);
243 Gamboa, 17 May 1979 (CJC); 34d, 12 Colén, Rio Cuango, 25
Ost. 1992, 15 Febr, 1995, 20 Sept. 1991, leg. Delgado (FD); 96d
° Colén, Coclesito, 18 June 1988, 27 Aug. 1988, 27 June 1986, 97
Noe 1986, 11 Sept. 1988, 13 March 1989, 30 Nov. 1988, leg. Del-
gado (FD); 12 Alto de Piedra, Santa Fé de Veraguas, 10 Sept. 1981,
leg. Delgado (FD).COSTA RICA:1<, 29° Chilamate, Heredia, 30
June 1992 leg. Callaghan (CJC);
130
8a.
8b.
KEY TO MALES OF HYPOPHYLLA
Dorsal wing surface with orange areas ................. 2
Dorsal wing surface blue with no orange areas .... argenissa
Orange on hindwing limited to a uniform submarginal
Almmam wide band bere acsde err era a ee ee: 3
Orange on hindwing wider thandmm ................. 4
Ventral hindwing post median row of spots between
Rs and Cul inastraightline ................... zeurippa
Ventral hindwing post median row of spots between
Rs and Cul not in asraightline................... martia
Hningelonhin dwn gavel owANn eae eee 5
Fringe on hindwing brown’ .......................... 6
Maculation heavy, tornus forewing always with an
Oran Pelswote rene tk scatter cic iisenes acer me callaghani
Maculation reduced, tornus of forewing rarely
with an OHINGD FOU’ coccrccocaovdgavoaopanncncvant sudias
Orange-yellow markings on disal area of forewing ..... idae
Orange-yellow markings absent from forewing .......... mi
Submarginal black line on forewing thin and
discontinuous; ventral surface ground color dark gray . . flora
Submarginal black line on forewing wide near apex,
thinner to tornus; ventral surface ground color light gray .. 8
Yellow orange area on hindwing wide, reaching nearly
halfway from margin to base; black marks between
forewing veins Cul and 1A+2A form continuous line
WITHTENGROL CELIO ara yee eres tine emma ote <2 ara naar caldensis
Yellow orange area on hind wing narrower, reaching
only a third of the distance from margin to base; black
marks between forewing veins Cul and 1A+2A do not
form continuous line with end of cell ........... lasthenes
KEY TO THE FEMALES OF HYPOPHYLLA
Dorsal forewing band yellow ......................... 2
Doxsalitorewineibandiwhiteymes erent ie teers sudias
Dorsal forewing band narrow, not exceeding 4mm ....... 6
Dorsal forewing band wide, greater then4dmm.......... 3
Dorsal forewing band more or less constant width ........ 4
Dorsal forewing band convex distad from costa, ending
in a point, followed by an irregular, variable patch of
yellow scaling before distal margin ............. argenissa
Dorsal forewing band ends opposite distal margin .... florus
Dorsal forewing band ends opposite tornus and
HAS? WNETEBIN ooocoacc0ggqocg000090ans00R0a000000006 5
Ventral forewing band more extensive, filling cell and
reaching base; forewing elongated ................. idaii
. Ventral forewing band reaches base only along costal
margin; forewing not elongated ................ lasthenes
Ventral ground color gray-brown, forewing band
bordered basad by line at end of cell ............Zewrippa
. Ventral ground color gray or white, forewing band not
bordered basad by line at end of cell................... 7
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
7a. Ventral hindwing post median row of spots between
Rs and Cul ina straightline .................. caldensis
7b. Ventral hindwing post median row of spots between
Rs and Cul not ina straight line ................. martia
ACKNOWLEDGMENTS
I am indebted to the curators of the museums visited, Mr. Philip
Ackery (BMNH) and Dr. Wolfram Mey (ZMHU) for allowing me to
study and photograph types, and to Dr. Jacques Pierre (MNHN),
Drs. Jorge Llorente and Armando Martinez (MZFC), México for ac-
cess to the collections under their care, and to Dr. Francisco Del-
gado, Panama, for the loan of material. To Drs. Robert Robbins and
Jason Hall of the USNM, Dr. Gerardo Lamas (MUSM) and an
anonymous reviewer my appreciation for helpful comments on the
manuscript. Dr. Lamas also kindly provided a photo of the type of H.
zeurippa.
LITERATURE CITED
D’aBrRERA, B. H. 1994. Butterflies of the Neotropical region, part iv
Riodinidae. Hill House, pp. 880-1096.
DEVRIES, P. J. 1997. The butterflies of Costa Rica and their natural
history. Vol. Riodinidae. Princeton. 288 pp.
EspIAL, L. & E. MONTENEGRO. 1977. Zonas de vida o formaciones
vegetales de Colombia. Memoria explicativa sobre el Mapa
Ecélogico. 2d Ed. Instituto Geografico Agustin Codazzi, Bo-
gota, 238 pp.
GopMaNn, F. 1903. Notes on some Central American and South
American Erycinidae, with descriptions of new species. Trans.
Ent. Soc. Lond. 5:529-550.
Gopman, F. & O. SALvIN. 1885. Biologica Centrali-Americana. In-
secta. Lepidoptera-Rhopalocera. London, Dulan & Co.,
Bernard Quaritch.
HEMMING, F. 1967. The generic names of the butterflies and their
type species (Lep. Rhop.) Bul. Brit. Museum of Nat. Hist.
Suppl. 9 London. 509 pp.
HeEwitTson, W. C. 1852-1877. Illustrations of new species of ex-
otic butterflies, selected chiefly from the collections of W.
William Saunders and William C. Hewitson. London.V.
Voorst. 5(5).
. 1870. Descriptions of five new species of diurnal Lepi-
doptera from Chontales, Nicaragua and one from Minas Gerais.
Ent. Mo. Mag:3-9, =
HoupripcE, L.R. 1947. Determination of world plant formation
from simple climatic data. Science 105:367—-368.
Kxots, A. 1970. Lepidoptera, pp. 115-129. In Tuxen, S. L. Taxon-
omist’s glossary of Genitalia in insects. Munksgaard, Copen-
hagen. 357 pp.
MILLER, L.D. 1969. Nomenclature of wing veins and cells. J. Res.
Lep. 8:37-48.
Seitz, A. 1917. Grossschmetterlinge der erde. V.5. Stuttgart, Alfred
Kernan Verlag. 657 pp.
STAUDINGER, O. & E. ScuHatz. 1888. Exotische Schmetterlinge.
Furth, G. Lowensohn V.1.
STICHEL, H. 1911. In Wytsmann, Lepidoptera Rhopalocera Fam.
Riodinidae. Genera Insectorum 112(B):239-452.
. 1930. Riodinidae I In W. Junk, ed. Lepidoptorum catalo-
gus, v. 40:113-544. Berlin.
Received for publication 13 September 1999; revised and accepted 1
March 2001.
Journal of the Lepidopterists’ Society
54(4), 2000, 131-136
SUITABILITY OF FOUR FAMILIES OF FLORIDA “BAY” SPECIES FOR
PAPILIO PALAMEDES AND P. GLAUCUS (PAPILIONIDAE)
J. MARK SCRIBER,
NICOLAS MARGRAF,!
AND
TAMMY WELLS
Department of Entomology, Michigan State University, East Lansing, Michigan 48824, U.S.A.
ABSTRACT. We tested the suitability of four Florida “bay” plant species for larval growth and adult oviposition preferences for two swal-
lowtail butterfly species, P. palamedes and P. glaucus. Much confusion exists about the host plant records for these butterflies in the literature.
We confirmed that of the four bay species tested, only red bay (Persea borbonia) of the Lauraceae was suitable to support larval survival and
growth of P. palamedes. All P. palamedes larvae offered sweetbay (Magnolia virginiana of the Magnoliaceae), Loblolly bay (Gordonia lasianthus
of the Theaceae) or Southern Bayberry (Myrica cerifera of the Myricaceae) died as neonates. Conversely, only sweet bay (Magnolia) was suit-
able for supporting survival of neonate P. glaucus larvae, with red bay, loblolly bay and bayberry unacceptable or toxic. Oviposition preferences
(individually assessed in a revolving four-choice arena) were strongly in favor of the most suitable host for each species: sweet bay received 93.9%
of the total P. glaucus eggs and red bay received 54.2% of the total P. palamedes eggs. The generally low level of adaptation of the Lauraceae
specialized spicebush swallowtail, Papilo troilus, to red bay was evident in that all nine Florida females refused to oviposit on any of the four
“bays” (including red bay).
Additional key words: _herbivore-plant interactions, tiger swallowtail butterfly, palamedes swallowtail, spicebush swallowtail, Lauraceae,
Magnoliaceae, Theaceae, Myricaceae.
Largely because of its distinctive plant species and literature are especially unclear about the Florida
geographic isolation as a peninsula, Florida harbors hosts since the use of the term “bay” (Mitchell & Zim
some unique and rare butterfly and moth species 1964, Scriber 1984) could refer to several species in
(Minnow & Emmel 1992; Emmel 1995). The phyto- four different families: 1) Red bay (Persea borbonia
chemical constraints and ecological opportunities af- (L.) Spreng) of the Lauraceae family, 2) Sweet bay or
fecting host selection and use by various Lepidoptera white bay (Magnolia virginiana (L.) of the Magnoli-
(Feeny 1995) are of general interest. Some of these aceae family, 3) Loblolly Bay (Gordonia lasianthus
unique biochemical, physiological, behavioral and eco- (L.) Ellis) of the Theaceae family, 4) or Southern Bay-
logical adaptations of Lepidoptera to Florida's local berry (Myrica cerifera L.) of the Myricaceae family.
host plants have been documented for the sweet bay For example, it has been stated that “Papilio
silkmoth, Callosamia securifera (Peigler 1976, Scriber palamedes. . . larvae feed on magnolias in the Bay
1983, Johnson et al. 1996) and three different species Tree hammocks of the Everglades” (Young 1955).
of swallowtail butterflies (Scriber 1986, Nitao et al. While loblolly bay occurs throughout Highlands
1991, Scriber et al. 1991, 1995, Lederhouse et al. 1992, County and in each Florida county north of Lake
Bossart & Scriber 1995a, 1995b, Frankfater & Scriber Okeechobee and into Georgia; red bay, sweet bay, and
1999a, 1999b). We have examined population fluctua- southern bayberry occur in every county of Florida in-
tions and the relative densities of Papilio palamedes, P. cluding the southern ones down to the Keys (Little
troilus, and P. glaucus in central Florida hammocks 1978, Nelson 1994).
and bay forests for the past decade (Scriber et al. In a study of latitudinal and geographic variation in
1998a). Field observations and lab studies suggest host plant records for the 560+ species of swallowtail
close affinities in preference for either Magnoliaceae butterflies, Scriber (1973, 1984) lists numerous cita-
or Lauraceae, but not both. tions that report both the Magnoliaceae and Lau-
Nonetheless, one of the most confusing examples of raceae families as host plants for the Papilio troilus, P.
uncertainty in host plant records for Lepidoptera ex- palamedes, and P. glaucus butterfly species. No spe-
ists for Papilio troilus, P. palamedes, and P. glaucus. cific records of Thay berry (or other Myricaceae) nor
Early food plant references for Papilio species in the Gordonia (or other Theaceae) were listed as hosts or
foods for these Papilio. However, since all four “bay”
‘Present address: Institute of Zoology, University of Neuchatel, species fr equently coexist in swamps, hammocks and
CH-2007 Neuchatel, Switzerland. floodplain forests of southern Florida, we were inter-
—— ee
——— — — —
a
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
4-Choice Papilio palamedes Oviposition
b
o
nN
ro}
Average Percentage Total Eggs
3
Loblolly bay Bayberry Other
Red bay Sweet bay
Plant Species
Fic. 1. The 4-choice oviposition preferences (mean + SE) of
eight Florida P. palamedes females on four “bay” species or “other”
(on paper lining or plastic arena) (red bay, Lauraceae; sweet bay,
Magnoliaceae; loblolly bay, Theaceae; southern bayberry, Myri-
caceae). The average eggs per female was 45 + 16 (SE).
ested in determining which of these four were used (or
capable of being used) for oviposition and/or for larval
food. We conducted our multichoice oviposition and
larval survival and growth studies in Highlands County,
Florida, at the Archbold Biological Station and some
bioassays were conducted with material sent to Michi-
gan State University.
Clarification of uncertain adult oviposition and larval
food plant records is critically important for under-
standing the relationships between insects and plants
in ecological and evolutionary time (Ehrlich & Raven
1964, Feeny 1995). A procedure for reporting host
plants or food plant observations was provided by
Shields et al. (1969), and they explained how misiden-
tification of either the insect or plant species can lead
to errors that get transmitted in textbooks, guidebooks,
and other scientific literature for decades. In our case,
it has been suspected that P. palamedes only feeds on
members of Lauraceae and not on members of Mag-
noliaceae (Brooks 1962, Scriber 1986); however sweet
bay (Magnolia virginiana, previously described as
Magnolia glauca) or Magnoliaceae is listed as hosts for
P. palamedes by numerous authors (Jordan 1907,
Young 1955, Klots 1958, Forbes 1960, Ehrlich &
Ehrlich 1961, Kimball 1965, Harris 1972, Tietz 1972,
Tyler 1975, Pyle 1981, Okano 1983, Beutelspacher-
Baights & Howe 1984, Opler & Krizek 1984, Pyle
1997). While still other authors do not list sweet bay
magnolia as a host for P. palamedes they do list it as a
host for P. troilus (Scudder 1889, Shapiro 1974, Howe
1975, Scott 1986, Tilden & Smith 1986), which is also
very unlikely (Scriber 1986, Scriber et al. 1991, Nitao
et al. 1992).
Our study was conducted with P. palamedes and P.
| 4-Choice Papilio glaucus Oviposition Preferences
Average Percentagae of Total Eggs
a
o
Red bay Sweet bay Loblolly bay Bayberry Other
Plant Species
Fic. 2. The 4-choice oviposition preferences (mean + SE) of five
Florida Papilio glaucus females on four “bay” species (see Fig. 1).
The average total eggs per females was 84 + 36 (SE).
glaucus in order to determine both the adult female
oviposition preferences in 4-choice arenas and the
neonate larval survival abilities of both of these swal-
lowtail butterfly species in no-choice bioassays with
four “bay” species of Highlands County, Florida. While
we intended to include Papilio troilus larvae in these
studies, we were not successful in obtaining oviposition
from the females (n = 9) we did secure.
METHODS
Adult females of Papilio palamedes, P. troilus, and P.
glaucus captured in Florida (Highlands, Levy and Co-
lumbia Counties) were set up in 4-choice oviposition
arenas to assess host preferences via contact chemore-
ception. The arenas are round plastic boxes that re-
volve 10 times per hour on a mechanized platform in
front of incandescent lights. Leaves of each bay species
were supported in water-filled floral aquapics and
draped along the side and bottom of the dish as de-
scribed by Scriber (1993). The four bay species, Persea
borbonia (red bay = RB), Magnolia virginiana (sweet
bay = SB), Gordonia lasianthus (loblolly bay = LB),
and Myrica cerifera (southern bayberry = BB), were
collected from the area west of Istokpoga Lake near
the city of Lake Placid, Florida. The oviposition arenas
were inside the invertebrate biology laboratory of Mark
Deyrup at the Archbold Biological Station. Herbarium
vouchers of these plant species have been deposited in
the Michigan State University research collection (JMS).
Each day, eggs were collected and the number of
eggs placed on each “bay” by individual females was
recorded as were the few eggs sometimes placed on
the side of the plastic dish. Adult females were fed
each day (with the exception of two periods of 48-hour
intervals over our four week study period). Some eggs
of P. palamedes and P. glaucus were sent by overnight
VOLUME 54, NUMBER 4
express mail with leaves of the bay species to our labo-
ratory at Michigan State University. When received,
the eggs were immediately placed in a growth cham-
ber set at 26° and 18 L: 6D photoperiod. The leaves
were placed in refrigeration at about 4°C in order to
maintain their freshness. Subsequent shipments of
leaves were sent via overnight mail, and put under re-
frigeration. A majority of these eggs and neonate lar-
vae of Papilio were maintained in Florida at the Arch-
bold Biological Station (at room temperature) for
parallel and simultaneous bioassays (n = 70 P. pala-
medes and n = 140 P. glaucus). Eggs were checked at
least twice a day so that neonates could be placed on
leaves within hours of eclosion at each location. 150 x
25 mm petri dishes were set up with a circle of paper
towel in the bottom and with an aquapic filled with
water in which to place the leaves to prevent them
from drying out. The maternal source from which the
larvae came was recorded for each bioassay treatment.
When larvae hatched, they were allocated equally
among the four types of bay leaves, three larvae per
dish. Larvae were gently placed on the leaves using a
camelhair brush. All work surfaces and the camelhair
brush used were cleaned with a 5% bleach/95% water
solution before and after uses. Larvae were kept in the
same growth chamber and were monitored daily to de-
termine survival through the first instar. Survivors
were reared to pupation on their host plant treatment.
RESULTS
It was absolutely clear that three of the “bay”
species (sweet bay, loblolly bay and bayberry) were un-
suitable for larvae of P. palamedes. All P. palamedes
larvae placed on these hosts died with little or no eat-
ing and no frass production within a few days during
the first larval instar stage, while survival and growth
was very good on red bay, (Persea borbonia). The red
bay survival for the neonates of six different families at
Michigan State University was: 100%, 100%, 80%,
60%, 44% and 14% (n = 3, 2, 10, 5, 16, and 7 respec-
tively). Similar results were obtained with sibling P.
palamedes larvae from one mother larva bioassayed on
each of the four treatment plant species in Florida at
the Archbold Biological Station: 37 of 40 neonates
reached the second instar on red bay and all neonates
on the other 3 “bay” species died (n = 10 larvae each)
without evidence of eating or feces.
In contrast, Papilio glaucus neonates all died (with
no eating or frass) on red bay as well as on loblolly bay
and bayberry. Sweet bay (Magnolia virginiana) was the
only suitable host for these tiger swallowtail larvae in
the Florida studies. All larvae on red bay (n = 16),
loblolly bay (n = 16), and bayberry (n = 16) refused to
133
eat and died, whereas 77 of 94 larvae (82%) survived
to the second instar on sweet bay. Similarly, the
smaller subset of eggs shipped overnight to Michigan
had a 50% larval survival on sweetbay and 0% survival
on each of the other species.
The 4-choice oviposition preferences of the adult
females of P. palamedes favored red bay over the other
3 “bays” in 7 of 8 females that laid more than 10 eggs.
The other female palamedes oviposited on the dish or
paper lining more than all of the four “bay” choices to-
gether. A total of sixteen other females of P. palamedes
laid fewer than 10 eggs each in their four-choice are-
nas and were excluded from analyses.
In contrast, five adult females of Papilio glaucus se-
lected the sweet bay (the only suitable larval host) for the
majority of their oviposition choices in the 4-choice
arena (four of these selected SB for more than 93% of
their eggs). A total of 28 other female P. glaucus laid
fewer than 10 eggs.
All females (n = 9) of the Florida P. troilus refused
to lay any eggs in the 4-choice oviposition arenas (five
from Highlands County in the south and two each
from Levy County and Columbia County in the
north). While we were unable to bioassay P. troilus lar-
val survival on the four “bay” species in this study, it
was of interest that those females of the spicebush
swallowtail all refused to lay any eggs, even with the
(Lauraceae) red bay presented as one of the choices.
DISCUSSION
It was clear that not even sympatric Florida popula-
tions of the palamedes swallowtail butterfly could sur-
vive on three of the four “bays” of Florida: only red bay
of the Lauraceae supported larval survival and growth.
Neither sweet bay (Magnoliaceae), loblolly bay
(Theaceae), nor southern bayberry (Myricaceae) were
eaten in no-choice bioassays and all neonate larvae
tested from six different families died. It has been
shown previously that sweet bay was toxic to P.
palamedes larvae (Scriber 1986) due to toxic neolig-
nans from Magnolia virginiana leaves (Nitao et al.
1992) but nothing is known about the specific deter-
rent/toxin mechanisms for loblolly bay nor Southern
bayberry.
It was also observed that larvae of Florida tiger swal-
lowtail butterflies, P. glaucus, could not survive the
neonate (first instar) stage on any of the “bays” other
than Magnolia virginiana (sweet bay) of the Magnoli-
aceae. All larvae died on (and refused to even eat)
leaves of red bay, loblolly bay, and southern bayberry
in no-choice bioassays. The toxicity of red bay to P.
glaucus was suggested earlier (Scriber 1986) but the
phytochemical cause is still not known for this plant or
SSS
eS mr ec ee oe ee
ee
ee
Ses Ae eee
134
the other two bay species bioassayed here. It is un-
usual that larvae of this polyphagous species refused to
even nibble on the leaves since it is known that
neonates often eat small trenches in the edge of toxic
plants from many families (Hagen 1986, Scriber 1988,
Scriber et al. 1991, Scriber et al. 1999).
Oviposition preferences of these two different Pa-
pilio were generally for the bay species that was the
only suitable host (red bay for P. palamedes, and sweet
bay for P. glaucus). The few scattered eggs on other
plants is not surprising, perhaps because the experi-
mental 4-choice oviposition arenas do not provide
enough space to prevent co-mingling of volatile chem-
icals (e.g., stimulants and deterrents). Contact chemo-
sensory stimulations are the key cues used by Lau-
raceae-specialized Papilio (Carter & Feeny 1999,
Carter et al. 1999, Frankfater & Scriber 2001) and the
deterrents in the Lauraceae (red bay) for Papilio glau-
cus females (Frankfater & Scriber 1999). These strong
tarsal contact and oviposition stimulation/deterrency
reactions to red bay phytochemicals could explain the
dominant patterns of single host recognition and pref-
erence in our arenas. Adult preference and larval per-
formance in these four bay species seem to be clearly
related for both P. palamedes and P. glaucus for host or
non-host. However, there are many ecological reasons
why such a physiological and behavioral “prefer-
ence/performance” correlation (whether or not genet-
ically based) might not always be expected within a
species (Thompson & Pellmyr 1991, Thompson 1995,
Bossart & Scriber 1999). For example, Florida popula-
tions of P. glaucus survive better and grow faster than
Georgia, Ohio, and Michigan populations on sweet
bay, which reflects significant differences in behavioral
and physiological adaptations of local populations
(Scriber 1986, Scriber et al. 1991). It is interesting
that, despite ecologically significant divergence among
P. glaucus populations of these four states, no de-
tectable genetic divergence in allozyme frequencies
were observed (Bossart & Scriber 1995b).
The fact that P. troilus females refused to lay any
eggs in the 4-choice oviposition arenas (even on red
bay) is interesting, since most palamedes females did.
In southern Florida, the only Lauraceous host plant
for P. troilus appears to be red bay, Persea borbonia.
However, throughout their geographic range north of
Gainesville, P. troilus females prefer sassafras (Sas-
safras albidum) or spicebush (Lindera benzoin). Lo-
cal larval adaptation of southern Florida P. troilus
populations to red bay, their only host species in
Highlands County, has been demonstrated to have a
genetic basis (Nitao et al. 1991, Nitao 1995). Five of
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
the nine P. troilus females were from Highlands
County (four were from the north). Perhaps the
recognition of red bay in the 4-choice oviposition ar-
ray is less well developed for P. troilus in general
compared to P. palamedes, especially since this is the
case for larval survival and growth on red bay (Scriber
et al. 1991, Lederhouse et al. 1992, Nitao 1995). Lar-
val survival on red bay was 77% for P. palamedes (n =
30 families, 562 larvae) and only 47% for P. troilus (8
families, 119 larvae) while larval survival on spice-
bush is basically reversed; 28% for P. palamedes (20
families, 165 larvae) and 86% for P. troilus (7 fami-
lies, 156 larvae; Scriber et al. 1991). For two other
Lauraceae species, larval survival on sassafras (78%
and 79%) and camphor tree (52% and 50%) is basi-
cally the same for P. palamedes and P. troilus, respec-
tively.
Despite their close co-occurrence in Florida
swamps, wetlands, hammocks, and forests, the four
“bay” species from four different plant families ana-
lyzed in our studies are clearly recognized as host and
non-hosts for P. palamedes and P. glaucus. Only red
bay is a host for P. palamedes and only sweet (white)
bay as a host for P. glaucus. When early literature
records are incorrect, mistakes can be passed on from
one to another (Shields et al. 1969). We believe this is-
sue has largely been clarified for Florida “bays” (Young
1955, Mitchell & Zim 1964, Scriber 1984) by our
oviposition and larval bioassays here.
ACKNOWLEDGMENTS
This research was partially supported by the Michigan State Uni-
versity Agricultural Experiment Station (Project #1644). We are
grateful to Dr. Mark Deyrup and the Archbold Biological Station
and the Highlands Hammock State Park for permission to use their
facilities.
LITERATURE CITED
BEUTELSPACHER-BAIGHTS, C. R. & W. H. Howe. 1984. Mariposas
de Mexico. Ediciones cientificas La Prensa Medica Mexicana,
S.A. Mexico, DF.
Bossakrt, J. L. & J. M. SCRIBER. 1995a. Genetic variation in ovipo-
sition preferences in the tiger swallowtail butterfly: interspe-
cific, interpopulation, and interindividual comparisons, pp.
183-194. In J. M. Scriber, Y. Tsubaki & R. C. Lederhouse
(eds.), The swallowtail butterflies: their ecology and evolution-
ary biology. Scientific Publishers, Inc., Gainesville, Florida.
. 1995b. Maintenance of ecologically significant genetic
variation in the tiger swallowtail butterfly through differential
selection and gene flow. Evolution 49:1163-1171.
Bossakt, J. L. & J. M. ScriBer. 1999. Preference variation in the
polyphagous tiger swallowtail butterfly (Lepidoptera: Papilion-
idae). Environ. Entomol. 28:628-637.
Brooks, J. C. 1962. Foodplants of P. palamedes in Georgia. J.
Lepid. Soc. 16:198.
Carter, M. & P. FEENy. 1999. Host plant chemistry influences
oviposition choice of the spicebush swallowtail butterfly, Papilio
troilus. J. Chem. Ecol. 25:1999-2009.
VOLUME 54, NUMBER 4
CARTER, M., P. FEENY & M. HariBaL. 1999. An oviposition stimu-
lant for the spicebush swallowtail butterfly, Papilio troilus (Lep-
idoptera: Papilionidae), from the levels of Sassafras albidum
(Lauraceae). J. Chem. Ecol. 25:1233-1245.
EHRLICH, P. R. & A. H. EnRuicu. 1961. How to know the butter-
flies. William C. Brown Co., Dubuque, Iowa. 262 pp.
EHRLICH, P. R. & P. H. RAVEN. 1964. Butterflies and plants: a study
in coevolution. Evolution 18:586-608.
EMMEL, T. C. 1995. Saving endangered swallowtails—the conser-
vation biology of Papilio aristodemus ponceanus in Florida and
P. homerus in Jamaica, pp. 359-369. In J.M. Scriber, Y. Tsubaki
& R.C. Lederhouse (eds.), Swallowtail butterflies: their ecology
and evolutionary biology. Scientific Publishers, Inc., Gainesville,
Florida.
FEENY, P. 1995. Ecological opportunism and chemical constraints
on the host associations of swallowtail butterflies, pp. 9-15. In J.
M. Scriber, Y. Tsubaki, and R.C. Lederhouse (eds.), The swal-
lowtail butterflies: their ecology and evolutionary biology. Sci-
entific Publishers, Inc., Gainesville, Florida.
Forbes, W. T. M. 1960. Lepidoptera of New York and neighboring
states. Part IV. New York State Agric. Expt. Sta. Mem. No. 371.
FRANKFATER, C. R. & J. M. SCRIBER. 1999. Florida red bay (Persea
borbonia) leaf extracts deter oviposition of a sympatric general-
ist herbivore, Papilio glaucus (Lepidoptera: Papilionidae).
Chemoecology 9:127— 132.
FRANKFATER, C. R. & J. M. ScriBER. 2001. Contact chemorecep-
tion guides oviposition of two Lauraceae-specialized swallowtail
butterflies (Papilionidae). Hol. Lepid. (in press).
HaceEN, R. H. 1986. The evolution of host-plant use by the tiger
swallowtail butterfly, Papilio glaucus Ph.D. Dissertation. Cor-
nell University. Ithaca, New York. 297 pp.
Harris, R. H. 1972. Butterflies of Georgia. Norman: University of
Okahoma Press. 326 pp
Howe, W. H. 1975. The butterflies of North America. Doubleday
and Co., Garden City, New York. 633 pp.
Jounson, K. S.,J.M. Scriper & M. Narr. 1996. Phenylpropanoid
phenolics in sweetbay magnolia as chemical determinants of
host use in Saturniid silkmoths (CALLOSAMIA spp.). J. Chem.
Ecol. 22:1955-1969.
JorDaAN, K. 1907. Papilio. Macrolepidoptera of the world. The
American Rhopalocera 5:11—45 In (A. Seitz, ed.). Alfred Ker-
nen, Stuttgart.
KIMBALL, C. P. 1965. The Lepidoptera of Florida: an annotated check-
list. State of Florida Dept. of Agric., Gainesville, Florida. 363 pp.
Kiots, A. B. 1951. A field guide to the butterflies of North
America east of the Great Plains. Houghton Mifflin Co. 349 pp.
LEDERHOUSE, R. C., M. P. Ayres, J. K. Nivao & J. M. SCRIBER.
1992. Differential use of lauraceous host by swallowtail butter-
flies, Papilio troilus and P. palamedes (Papilionidae). Oikos
63:244-252.
LitTLe, E. L. JR. 1978. Atlas of United States Trees. Vol. 5. Florida
U.S. Forest Service. Misc. Publ. 1361. Washington, DC.
Minnow, M. C.& T. C. EMMEL. 1992. Butterflies of Florida Keys.
Scientific Publishers. Gainesville, Florida.
MITCHELL, R. T. & H. M. Zim. 1964. Butterflies and Moths.
Racine.
NELSON, G. 1994. The trees of Florida. Pineapple Press, Inc., Sara-
sota, Florida.
Nia, J. K. 1995. Evolutionary stability of swallowtail adaptations
to plant toxins, pp. 39-52. In J. M. Scriber, Y. Tsubaki & R. C.
Lederhouse (eds.), The swallowtail butterflies: their ecology and
evolutionary biology. Scientific Publishers, Inc., Gainesville,
Florida.
Ni7ao, J. K., M. P. Ayres, R. C. LEDERHOUSE & J. M. ScriIBER. 1991.
Larval adaptation to lauraceous hosts: geographic divergence in
the spicebush swallowtail butterfly. Ecology 72:1428-1435.
NiTao, J. K., K. S. JouNnson, J. M. ScrIBER & M. G. Nair. 1992.
Magnolia virginiana neoligin compounds as chemical barriers
to swallowtail butterfly host use. J. Chem. Ecol. 18:1661-1671.
135
OKANO, K. 1983. The revision of classification on the genera of Pa-
pilionidae in the world (Preliminary report) Part I: with de-
scription of a new genus: Tokurana (Acta Rhopalocerologica).
No. 5, pp. 1-75.
OPLER, P.A. & G.O. KrizeK. 1984. Butterflies east of the Great
Plains. Johns Hopkins Univ. Press. Baltimore, Maryland. 299 pp.
PEIGLER, R. S. 1976. Observations in host plant aes and
larval nutrition in Callosamia (Satumiidae). J. Lepid. Soc.
30:184—-187.
Pye, R. M. 1981.Field guide to North American butterflies. Chan-
ticleer Press, Knopf, Inc. New York.
PYLE, R. M. 1997. National Audubon Society field guide to the North
American Butterflies. Chanticleer Press, New York, New York.
Scotr, J. A. 1986. The butterflies of North America: A natural his-
tory and field guide. Stanford Univ. Press. Stanford, Connecticut.
SCRIBER, J. M. 1973. Latitudinal gradients in larval feeding special-
ization of the world Papilionidae. Psyche 80:355-373.
. 1983. The evolution of feeding specialization, physiologi-
cal efficiency, and host races in walasied Papilionidae and Sat-
urniidae, pp. 373-412. In R. F. Denno & M. S. McClure (eds.),
Variable plant and herbivores in natural and managed systems.
Academic Press, New York.
. 1984. Larval foodplant utilization by the world Papilion-
idae (Lepidoptera): Latitudinal gradients reappraised. Toku-
rana (Acta Rhopalocerologica) 2:1—50.
. 1986. Origins of the regional feeding abilities in the
tiger swallowtail butterfly: ecological monophagy and the Pa-
pilio glaucus australis subspecies in Florida. Oecologia
71:94-103.
1988. Tale of the tiger: Beringial biogeogaphy, bionomial
dlastReatton, and breakfast choices in the Papilio glaucus com-
plex of butterflies, pp. 240-301. In K.C. Spencer (ea ), Chemi-
cal Mediation of Coevolution. Academic Press, New York.
. 1993. Absence of behavioral induction in oviposition pref-
erence of Papilio glaucus (Lepidoptera: Papilionidae). Great
Lakes Entomol. 26:81-95.
. 1996. Tiger tales: natural history of North American swal-
lowtails. American Entomologist 42:19-32.
SCRIBER, J. M., R. C. LEDERHOUSE & L. CONTARDO. 1975. Spice-
bush, Lindera benzoin (L.), a little known foodplant of Papilio
glaucus (Papilionidae). J. Lepid. Soc. 29:10-14.
SCRIBER, J. M., M. D. DEERING, L. FRANCKE, W. WEHLING & R. C.
LEDERHOUSE. 1998a. Notes on the butterfly population dy-
namics of 3 Papilio species in south central Florida (Highlands
County). Hol. Lepid. 5:53-62.
SCRIBER, J. M., R. C. LEDERHOUSE & R.V. DOWELL. 1995. Hy-
bridization studies with North American swallowtails, pp.
269-281. In Scriber, J.M., Y. Tsubaki & R.C. Lederhouse (eds. ),
The swallowtail butterflies: their ecology and evolutionary biol-
ogy. Scientific Publishers, Inc., Gainesville, Florida.
SCRIBER, J. M., R. C. LEDERHOUSE & R. H. HAGEN. 1991. Food-
plants and evolution within the Papilio glaucus and Papilio
troilus species groups (Lepidoptera: Papilionidae), pp. 341-373.
In PW. Price, T.M. Lewisohn, G.W. Fernandez & W.W. Benson
(eds.), Plant-animal interactions: evolutionary ecology in tropical
and temperate regions. John Wiley, New York.
ScRIBER, J. M., K. Wer, D. Parry & J. DEERING. 1999. Using hy-
brid and backcross larvae of Papilio canadensis and P. allan to
detect induced chemical resistance in hybrid poplars experi-
mentally defoliated by gypsy moths. Entomologia exp. & appl.
91:233-236.
ScuDDER, S. H. 1889. The butterflies of the Eastern United States
and Canada. Vol. 2: 1219-1364. Scudder, Cambridge.
SHapPiRO, A. M. 1974. Butterflies and skippers of New York State:
Search 4: 1-59. Cornell Univ. Expt. Sta. Publ.). Ithaca, New
York.
SHIELDS, O., J. F. EMMEL & D. E. BREEDLOVE. 1969. Butterfly lar-
val foodplant records and a procedure for reporting foodplants.
]. Res. Lepid. 8:21-36.
136
THOMPSON, J. N. 1995. The origins of host shifts in swallowtail but-
terflies versus other insects, pp. 195-203. In J. M. Scriber, Y.
Tsubaki, and R. C. Lederhouse (eds.), The swallowtail butter-
flies: their ecology and evolutionary biology. Scientific Publish-
ers, Inc., Gainesville, Florida. 459 pp.
THompsoN, J. M. & O. PELLMyR. 1991. Evolution of oviposition
behavior and host preference in Lepidoptera. Ann. Rev. Ento-
mol. 36:65-89.
TiETz, H. M. 1972. An index to the described life histories, early
stages, and hosts of the Macrolepidoptera of the continental
U.S. and Canada. 2 Vol. Allyn Museum. Florida.
TILDEN, J. W. & A. C. SmirH. 1986. A field guide to Western But-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
terflies. The Peterson Field Guide Series. Houghton Mifflin,
Boston, Massachusetts. 370 pp:
TyLer, H. A. 1975. The swallowtail butterflies of North America.
Naturegraph. Healdsburg, California. 192 pp.
TyLer, H. A., K.S. BRown & K. H. WILSON. 1994. Swallowtail but-
terflies of the Americas. Scientific Publ., Gainesville, Florida.
378 pp.
YOUNG, BN 1955. Notes on collecting in southem Florida. Lepid.
News 9:204—212.
Received for publication 28 October 1999; revised and accepted 15
January 2001.
Journal of the Lepidopterists’ Society
54(4), 2000, 137-144
ON THE USE OF ULTRAVIOLET PHOTOGRAPHY AND ULTRAVIOLET WING PATTERNS IN
BUTTERFLY MORPHOLOGY AND TAXONOMY
HELGE KNUTTEL!
KONRAD FIEDLER
Lehrstuhl Tierdkologie I, Universitat Bayreuth, D-95440 Bayreuth, Germany
email: Helge. Knuettel@uni-bayreuth.de
ABSTRACT. In a series of feeding experiments we found that, depending on the larval food plant species or part of food plant ingested,
individuals of the blue butterfly Polyommatus icarus (Lycaenidae) exhibit broad variation of wing patterns in the ultraviolet (UV) range of wave-
lengths which is invisible to humans. Such intraspecific variability in UV wing patterns has been underestimated thus far due to the rather de-
manding approach needed to study these patterns. We discuss methodological problems with the assessment of butterfly UV wing patterns by
UV photography. Given proper standardization, UV photography is a suitable method to qualitatively assess UV wing patterns for possible use
in morphology or systematics. Spectrophotometry should preferably be used as quantitative method when considering UV wing pattems in a
communication context. No higher value should be attached to UV wing patterns as compared to human visible wing patterns.
Additional key words: Polyommatus, ultraviolet light, visual communication, color, phenotypic plasticity.
Ultraviolet (UV) wing patterns have been widely
used in butterfly systematics. Differences in UV re-
flectance patterns of butterfly wings proved helpful in
the revision of otherwise morphologically very similar
taxa, such as the genera Colias (Ferris 1973, Silber-
glied & Taylor 1973, 1978, Kudrna 1992) and
Gonepteryx (Nekrutenko 1964, Kudrna 1975, Brunton
et al. 1996). Differing UV wing patterns were sus-
pected to act as “isolating mechanisms” between
closely related species, e.g., by Meyer-Rochow (1991)
in Lycaena, or shown to be involved in mate choice of
several species (e.g., Silberglied & Taylor 1973, 1978,
Rutowski 1977, 1981).
Butterflies, in general, perceive UV light and UV vi-
sion is an integral part of their visual capabilities
(Eguchi et al. 1982, Silberglied 1984, Lunau & Maier
1995, Tovée 1995, Kelber & Pfaff 1999). The same is
true for many visually guided butterfly predators such
as birds, lizards, and robberflies (Menzel & Backhaus
1991, Jacobs 1992, Fleishman et al. 1993, Tovée 1995).
Therefore, UV coloration of butterfly wings has to be
considered as an essential part of overall wing patterns
in the spectral range of 300 nm to 700 nm, i.e., the en-
tire range of visual communication (Endler 1990,
Cuthill & Bennett 1993, Bennett et al. 1994b). Human
observers cannot perceive UV light directly. This may
be the reason why many researchers implicitly or ex-
plicitly attached an extraordinary meaning to the col-
oration of butterfly wings in the UV range of wave-
lengths as compared to the human visible range. In the
New Zealand Lycaena salustius (Lycaenidae) species
complex, for example, distinction of species based on
‘Present address: Universititsbibliothek Regensburg, D-93042
Regensburg, Germany.
human-visible wing patterns and morphology of geni-
talia is possible (Gibbs 1980), yet the discovery of
marked differences in UV reflectance patterns was
taken even as evidence to suggest the existence of UV
wing pattern-based isolating mechanisms (Meyer-
Rochow 1991; for another case study in the genus Ly-
caena see Schaider (1988) versus van Oorschot & de
Prins (1989)). Human lack of UV perception, and the
processing of visual stimuli in other animals by nervous
systems which are completely different from our own,
make it very hard for the researcher to imagine what
the world may look like for other animals. But it seems
as if the lack of UV vision in humans and many other
mammals is the exception rather than the rule in the
animal kingdom (cf. Tovée 1995).
In this paper we compare UV photography and
spectrophotometry as methods for assessing butterfly
UV wing patterns. In particular, we discuss some
methodological problems with the assessment of but-
terfly UV wing patterns by UV photography. Finally,
we point out an underestimate of individual variability
in UV wing patterns which may result from such
methodological problems as well as from the neglect of
environmentally driven phenotypic plasticity.
MATERIALS AND METHODS
Feeding experiments. Mated females of Polyom-
matus icarus (Rottemburg, 1775) (Lycaenidae) were
caught in summer 1997 at two locations in Norther
Bavaria, Germany and allowed to lay eggs. Caterpillars
used in this study were from the F 1 or F2 generations of
these females. All larvae were reared in the same climate
chamber (25°C, 18 h light, 6 h dark). We kept the larvae
in transparent plastic boxes (125 ml) lined with moist pa-
138
per tissue. Fresh food plant material was available ad
libitum and was supplied at least every two days. Food
plants were either grown outside in a garden (Med-
icago sativa L. leaves) or collected locally from natural
populations (flowers of Trifolium repens L., flowers
and leaves of Lotus corniculatus L.) (all Fabaceae). We
killed the emerging butterflies with either HCN or by
deep freezing. We measured spectral reflectance of
the wings of these specimens and took UV pho-
tographs.
Ultraviolet photography. UV photographs were
taken with a Pentax Asahi Ultra-Achromatic-Takumar
85 mm/f 4,5 UV transmitting lens and Novoflex Auto-
Bellows on Agfapan APX 100 film. To exclude all but
ultraviolet light a combination of 3 mm Schott UG1
and 2 mm Schott BG38 filters was placed in front of
the lens for UV photographs. Illumination was pro-
vided by two Metz Mecablitz 45 CT1 flash lights in
manual mode so that the same amount of light was
available for every exposure. Emission spectra of these
flash lights reach far enough into the UV range for the
purpose of this study (data not shown). Photographic
processing and development of film material was stan-
dardized as much as possible. Films were developed
for 5 min in 10% Agfa Neutol and fixed for 4 min in
Tetenal fixing agent at 21°C.
To examine the effect of using different grades of
paper on the appearance of prints to be used for com-
paring UV patterns, we produced prints on different
grades of Agfa Broviro Speed glossy paper. These
prints were then developed for 3 min with 10% Agfa
Neutol and fixed for 15 min with Tetenal fixing agent
ate AUG,
For purposes of comparison we took color pho-
tographs of all objects on Kodak Elite II 100 slide film.
We used the same equipment as above but without the
filters and with one flash light only.
Spectrophotometry. We measured wing re-
flectance with a L.O.T. Oriel spectrometer system (In-
staSpec II diode array detector, MS 125 spectrograph
with 400 l/mm grating, sighting optic) equipped with a
Zeiss Ultrafluar 10/0.20 UV transmitting objective at a
right angle to the wing surface. Measuring spot diam-
eter was 0.2 mm, numerical aperture of the measuring
beam was 0.14. The measuring spot was illuminated at
an angle of 45° to the wing surface via liquid light
guide by an Osram XBO 75 W/2 OFR lamp powered
by a L.O.T. Oriel 68806 power supply. Numerical
aperture of the illumination was 0.08. Wings were ori-
ented so that they were all illuminated from the same
apical direction. With this setup we were able to
record spectral reflectance of individual wing spots in
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
the range of 300 nm to 700 nm with a resolution of ap-
prox. 0.5 nm. A Spectralon™ 99 reflectance standard
was assumed as having 100% reflectance. For further
details of methods see Kniittel and Fiedler (2001).
RESULTS
UV patterns strongly varied in both sexes of the Eu-
ropean common blue butterfly, Polyommatus icarus
(Figs. 1, 2). We found consistent differences of UV re-
flectance among individuals that were fed different
plant species or plant parts during their larval stages.
Reflectance in the UV was much lower for animals
reared on flowers of Trifolium repens and flowers or
leaves of Lotus corniculatus, as compared to animals
reared on leaves of Medicago sativa (Figs. 1, 2). These
differences were most pronounced in the white spots
(as seen with human eyes) but were apparent in the
underside ground coloration, too (Fig. 1). Overall,
judging from the UV photographs (Fig. 1), one might
be tempted to assign highly UV-reflecting specimens
reared on M. sativa foliage to a different ‘species’. No
differences in UV reflectance were found for the up-
persides, the orange spots, and the black spots (Kniit-
tel & Fiedler 1999).
Altering photographic processing had a strong effect
on the appearance of the resulting prints, a phenome-
non well known to any photographer. The influence of
using photographic paper of differing grades is illus-
trated in Fig. 3. Even if processed from the identical
negative using the same chemicals, the resulting prints
of UV photographs may be quite different. We there-
fore included a calibrated gray scale, made from thick
chromatography paper and dyed with various dilutions
of black India ink, in every photograph. Spectral re-
flectance of the steps of the gray scale is illustrated in
Fig. 4. Parts of a given photographic print that are of
similar brightness, compared to the gray scale, will
have a similar reflectance value (Figs. 1 and 3).
The differences in wing patterns in the UV range,
among individuals reared on different plant species or
plant parts, emerged from UV photographs and spec-
trophotometric measurements alike. However, more
subtle or gradual host plant-dependent color differ-
ences could better be visualized in the reflectance
spectra (Fig. 2). For example, only by studying the
spectra is one able to identify the wavelength ranges
where individuals reared on M. sativa foliage converge
into the variation seen in individuals fed other food
plants. Moreover, the small but consistent differences
between food treatments in the human-visible range
were also only noticeable using spectrophotometric
measurement data.
VOLUME 54, NUMBER 4 139
L. comiculatus leaves”
LE. Se flowers T. repens (ones
oC #2 o #227
L. corniculatus leaves M. sativa leaves
Q #251 Q #132
Fic. 1. UV photographs of the undersides of Polyommatus icarus reared on different food plants. Differences in UV patterns seen in these
individuals are representative for larger series. Individuals were reared on leaves of Lotus corniculatus (upper left), leaves of Medicago sativa
(upper right), flowers of Lotus commculots (lower left), and flowers of Trifolium repens (lower right). Upper photograph: males, lower photo-
graph: females.
140
DISCUSSION
1. Underestimate of individual variability in
UV coloration
Butterfly systematists are well aware of the large ex-
tent of intraspecific variation in wing patterns and col-
oration within the human visible spectral range. In
contrast, intraspecific variability in UV reflection pat-
terns has been underestimated so far. We feel that this
mainly arises from the difficulty in studying UV wing
patterns, making the comparison of large series of
specimens more laborious and demanding compared
to patterns seen in the human visible range. Since in
most studies UV photography applied to small samples
was used to assess UV wing reflectance, subtle differ-
ences in UV reflectance of individuals may have fre-
quently been missed.
Though only studied for a small range of butterfly
species thus far, intraspecific variation of UV patterns
may be as pronounced as, or even larger than that in
other ranges of wavelengths visible to humans (Brun-
ton & Majerus 1995, Kniittel & Fiedler 1999, 2001,
this work). Yet, minor differences in UV patterns have
been sometimes taken as evidence for erecting new
species or subspecies (e.g., Nekrutenko 1968, Schaider
1988, but see van Ooorschot & de Prins 1989), or as a
later confirmation of taxonomic hypotheses originally
proposed on the grounds of other data (e.g., Meyer-
Rochow 1991, Coutsis & Ghavalas 1996).
Differences in UV wing patterns may be due to ge-
netic or environmental reasons, but only genetically
determined UV wing patterns are of systematic impor-
tance. We demonstrated that high intraspecific varia-
tion in UV wing patterns in Polyommatus icarus can be
caused by different larval food plants under otherwise
identical rearing conditions among individuals from
the same parents.
Flavonoids are a class of secondary plant compounds
that highly absorb UV light (Harborne 1991, 1999).
Some Polyommatus species sequester flavonoids from
their larval food plants, and these pigments are de-
posited in the wings during metamorphosis (Wilson
1987, Wiesen et al. 1994, Geuder et al. 1997, Korn-
maier 1999). Using artificial diets which only differed
in their flavonoid content but were otherwise identical
in their chemical composition, Kniittel & Fiedler
(1999, 2001) showed that flavonoids sequestered by
the larvae alter wing reflectance mainly in the UV
range. In the polyphagous P. icarus the types and
amounts of flavonoids that are taken up and stored by
the larvae vary strongly depending on the larval food
plants (Wiesen et al. 1994, Burghardt et al. 1997, 2001,
Schittko et al. 1999). Therefore, it seems very likely
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
100
ot - — -— Medicago leaves
Sess Lotus leaves
80 Lotus flowers
—-- — Trifolium flowers
SS
c
Ae)
16)
®
oD see
oc 0 saa] (BS
300 350 400 450 500 550 600 650 700
Wavelength [nm]
100 -— -— -— Medicago leaves
QO or Lotus leaves
Lotus flowers
80 — .. = Trifolium flowers
60
3 40
Oo
ca 20
®
oO
oc
0
300 350 400 450 500 550 600 650 700
Wavelength [nm]
Fic. 2. Spectral reflection of the white spots of the undersides of
the hindwings of Polyommatus icarus reared on different food
plants. Each curve is the mean of the measurements of 5 to 10 spots
of a hind wing of one of the individuals illustrated in Fig. 1. Individ-
uals were reared on leaves of Lotus corniculatus ( _ . ), leaves of
Medicago sativa ( _ _ ), flowers of Lotus corniculatus ( __ ), and
flowers of Trifolium repens (__..). Upper part: males, Lower part:
females. Conventions as in Fig. 1.
that flavonoids are involved in mediating variation in
UV wing patterns of P. icarus feeding on different food
plants in nature.
It is important to emphasize that intraspecific vari-
ability in UV reflectance in P. icarus appears to be
caused by chemical variation in the host plants, while
VOLUME 54, NUMBER 4
variation in UV reflectance may be caused by structural
colors in other species (e.g., Colias eurytheme (Brun-
ton & Majerus 1995)). UV pattern variation in P. icarus
therefore must be regarded as a host-plant derived,
environmentally shaped form of phenotypic plasticity,
although heritable components cannot be fully ex-
cluded. As different food plant species or plant parts
are of varying value as a food source to P. icarus, it
seems likely that UV wing patterns may be used in in-
traspecific visual communication, indicating other food
plant-derived properties of individuals, such as nitro-
gen content (Burghardt et al. 2001). Males of P. icarus
discriminate between flavonoid-rich and flavonoid-
free female dummies, preferring UV-absorbing,
flavonoid-rich dummies (Burghardt et al. 2000, Kniit-
tel & Fiedler 2001).
The differences shown here in the UV photographs
(Fig. 1) and quantitatively demonstrated in the accom-
panying reflectance spectra (Fig. 2) very much resem-
ble the differences in UV reflectance claimed by Cout-
sis & Ghavalas (1996) as characters separating
Polyommatus icarus and the recently described P. an-
dronicus Coutsis & Ghavalas, 1995. As differences of
such magnitude can be found within one species and
even among offspring of the same parents, they are
unlikely to be sufficient to differentiate between
species. No quantitative data on spectral wing re-
flectance are available for P. andronicus, and the range
of individual variation has not been documented statis-
tically. Therefore we cannot presently assess whether
significant differences in UV patterns may exist be-
tween P. andronicus and P. icarus. However, based on
the UV photographs in Coutsis & Ghavalas (1996) it
seems unlikely that UV reflectance in P. andronicus
falls outside the range observed in the highly variable
species P. icarus.
2. Problems related to the technical visualization of
UV patterns
As humans cannot see ultraviolet light, UV wing
patterns must be translated into a form of information
that is accessible to us. This must be accomplished by
appropriate technical means. UV photography or UV
videoviewing was chosen in most studies of UV wing
patterns known to us (e.g., Ferris 1973, Rutowski
1977, 1981, Bowden & Kay 1979, Meyer-Rochow
1991, Kudrna 1992, Coutsis & Ghavalas 1996). Both
methods yield comparable spatial pattern information
but almost no spectral information. Ultraviolet light
from a broad range of wavelengths is reduced to a
single brightness value for every point or pixel in the
picture. Usually the spectral response of the picture-
generating system is unknown. Alternatively, wing re-
141
flectance can be measured by spectrophotometry
(Ghiradella et al. 1972, Endler 1990, Brunton & Ma-
jerus 1995).
Both UV photography and spectrophotometry have
advantages and disadvantages in their practical use.
When selecting a method to study UV wing patterns
the first step must be to answer the questions “What is
the purpose of the study? What is it that UV patterns
should actually tell me?” Not all studies have ade-
quately addressed these questions. However, different
conclusions may have to be drawn from the use of dif-
ferent methods. Therefore it is important to be clear
about the purpose of the study before choosing the
method.
UV patterns may be considered as a morphological
feature like any other character. UV patterns result
from wing areas that differ from each other in UV re-
flectance due to their physical and chemical constitu-
tion. There is no conceptual difference to the reflec-
tions or colors in the human-visible range. Therefore,
UV wing patterns may be used as regular morphologi-
cal characters in systematics, if they are assessed ap-
propriately. For example, if individuals within a
species exhibit substantial variation in UV wing pat-
terns, such as we found in P. icarus, then UV wing pat-
ters may not be appropriate systematic characters. UV
photography done in the right way (see below) seems
a perfectly acceptable means for the description of UV
wing patterns in this context.
On the other hand, UV wing patterns may serve as
signals in a behavioral context. They may be important
in mimetic or aposematic coloration (Os Beccaloni
1997) or in sexual selection (e.g., Brunton & Majerus
1995), to give examples. But it is not sufficient to sim-
ply assume that UV patterns do have a function, for ex-
ample in mate recognition. This has to be proven in
separate studies reaching farther than assessing differ-
ences in UV reflectance only. Wheh considering the vi-
sual physiology of butterflies (e.g., Eguchi et al. 1982)
or other visually guided species interacting with but-
terflies (e.g., Bennett et al. 1994a) it seems likely that
UV light is important in the species’ interactions. But
this is so only because UV sensitivity is an integral part
of these species’ visual systems and is not a conse-
quence of some putative special quality of UV light or
vision in the UV range. The mere possibility that UV
patterns serve a function in communication gives them
no special or “higher” value in systematic reasoning
(see e.g., Meyer-Rochow 1991, Brunton et al. 1996).
The same is true when comparing UV patterns to hu-
man-visible color patterns.
To emphasize this point: There is no reason at all to
assign a higher value to UV patterns than to human-
Fic. 3. UV photograph of the underside of one female Poly-
ommatus icarus reared on leaves of Lotus corniculatus. Both prints
were produced from the same negative but on photographic paper
of differing grades and accordingly illuminated for different time pe-
riods. They illustrate the influence of a minor change in photo-
graphic processing. A calibrated gray scale (cf. Fig. 4) included in
the photographs allows for a comparison up to a certain degree de-
spite the different appearance of the prints. Width of a step of the
gray scale is 5 mm. Upper print: Illumination for 5.5 sec, aperture
5.6 on grade 1 paper. Lower print: Illumination for 10 sec, aperture
5.6 on grade 5 paper.
visible wing patterns. And there is no reason to pre-
suppose a special function of UV wing patterns as a
signal in visual communication.
More elaborate techniques must be applied when
studying UV wing patterns as signals used in visual com-
munication. Only when there are very strong differ-
ences, without intermediates in UV reflectance, will UV
photography be useful in such a context. This might be
the case when comparing wing patterns with and with-
out strongly reflecting structural colors. However, even
then UV photography will provide rather coarse qualita-
tive results only and individual variability of UV re-
flectance may be missed (cf. Endler 1990, Brunton &
Majerus 1995, this study). Variation in spectral informa-
tion within the UV range that may be important in com-
munication will also not be apparent with UV photogra-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
100
80
60
3 40
6
rs 20
co)
oO
am
0
300 350 400 450 500 550 600 650 700
Wavelength [nm]
Fic. 4. Spectral reflection of the steps of the gray scale shown in
Figs. 1 and 3. The higher the reflectance values the brighter the ar-
eas appear in the photographs. The curve for the darkest step almost
falls in line with the abscissa. Mean spectral reflection for any given
range of wavelengths may easily be obtained from the very flat, al-
most horizontal curves. The UV range is from 300 nm to 400 nm and
the visible range is from 400 nm to 700 nm. Abscissa: Wavelength in
nanometers. Ordinate: Reflection in %, i.e., the amount of light re-
flected at a given wavelength as compared to a white standard as-
sumed to reflect 100%.
phy. For these reasons, in the study of communication
or sexual selection (Bennett et al. 1994b), the method of
choice is reflectance spectrophotometry. The whole
range of 300 nm to 700 nm needs to be covered by the
measurements, that is from the ultraviolet to the red.
Comparison of obtained spectra can be done by appro-
priate statistical procedures (Endler 1990, Cuthill et al.
1999, Kniittel & Fiedler 2001). So far, for only very few
species is information available on the physiology of
photoreceptors and associated neurons. For these
species a physiological model closer to the processes oc-
curring in the organisms may allow to calculate a classi-
fication of colors. More details on the assessment of col-
ors in animal communication systems may be found in
the excellent works of Endler (1990), Cuthill and Ben-
nett (1993), and Bennett et al. (1994b).
UV photography provides an easy method to assess
the spatial distribution of areas of differing UV re-
flectance. Yet, in the majority of studies qualitative
rather than quantitative results were obtained. This
means that wing areas were mostly classified as UV-
reflecting vs. not UV-reflecting. However, reflectance
is a continuous measure that may not easily be as-
sessed in discrete steps (cf. Fig. 2).
Moreover, comparisons between different pho-
VOLUME 54, NUMBER 4.
tographs and/or studies may be difficult without
proper standardization. Brightness and contrast in
photographic prints depend on a number of parame-
ters, not all of which might be under the control of the
investigator. Important parameters that contribute to
variation are amount and spectral composition of illu-
minating light, spectral transmission of photographic
lenses and of UV transmitting filters, the film type, and
all kinds of influences on photographic processing in-
cluding printing during publication. Variation in the
appearance of UV photographs may arise from a minor
difference in photographic processing as illustrated in
Fig. 3. Therefore, a detailed description of optical in-
strumentation, processing and film material should be
given and, as a minimum standard, a gray scale of
known UV reflectance should be part of every UV
photograph. Such a gray scale will allow comparisons
between photographs up to a certain degree because it
provides a set of reflectance standards revealing inten-
tional and unintentional differences in brightness or
contrast between photographs (Figs. 1 and 3). This
method is beautifully described in the pioneering work
of Daumer (1958) on the UV patterns of flowers.
UV spectrophotometry yields very accurate quanti-
tative data but requires expensive equipment not avail-
able to most systematists and a fair amount of com-
putational data processing. Spectrophotometry is
superior whenever spectral information will be re-
quired to answer biological questions. However, in
contrast to UV photography, spectrophotometry will
not provide easily comprehensible spatial pattern in-
formation. Hence, for taxonomic purposes where the
recognition and documentation of qualitative similari-
ties and discrepancies in wing patterns is usually the
most important goal, properly standardized UV pho-
tography will continue to be the preferred method,
though at the cost of loss of quantitative information.
ACKNOWLEDGMENTS
We are indebted to Prof. Dr. Klaus Lunau who generously per-
mitted us to use his UV lens. Dr. Susanne Hanika, Prof. Dr. Klaus
Lunau, Prof. Dr. Deane Bowers and two unknown reviewers made
helpful comments on an earlier version of the manuscript. We thank
Annick Servant for technical assistance. HK was supported by a
grant from the Konrad-Adenauer-Stiftung.
LITERATURE CITED
BECCALONI, G. W. 1997. Ecology, natural history and behaviour of
ithomiine butterflies and their mimics in Ecuador (Lepi-
doptera: Nymphalidae: Ithomiinae). Trop. Lepid. 8:103-124.
BENNETT, A. T. D. & I. C. CUTHILL. 1994a. Ultraviolet vision in
birds: what is its function? Vision Res. 34:1471-1478.
BENNETT, A. T. D., I. C. CurHiLt & K. J. Norris. 1994b. Sexual se-
lection and the mismeasure of color. Am. Nat. 144:848-860.
Bowben, S. R. & O. N. Kay. 1979. Ultra-violet photography of
Lepidoptera. Nota lepid. 2:27-30.
143
BRUNTON, C. F. A. & M. E. N. Majerus. 1995. Ultraviolet colours
in butterflies: intra- or inter-specific communication? Proc. R.
Soc. Lond. B 260:199-204.
BRUNTON, C. F. A., P. J. C. RusseLt & M. E. N. Majerus. 1996.
Variation in ultraviolet wing patterns of brimstone butterflies
(Gonepteryx: Pieridae) from Madeira and the Canary Islands.
Entomologist 115:30-39.
BURGHARDT, F. 2000. Stoffwechsel und dkologische Funktion von
Flavonoiden im Bléuling Polyommatus icarus (Lepidoptera:
Lycaenidae). iv, 192 pp. Ph.D. Dissertation, Bayerische Julius-
Maximilians-Universitat W’ tirzburg, Germany.
BURGHARDT, F., K. FIEDLER & P. ProKscH. 1997. Uptake of
flavonoids from Vicia villosa (Fabaceae) by the lycaenid butter-
fly, Polyommatus icarus (Lepidoptera: Lycaenidae). Biochem.
Syst. Ecol. 25:527-536.
BURGHARDT, F., H. KNUTTEL, M. BECKER & K. FIEDLER. 2000.
Flavonoid wing pigments increase attractiveness of female com-
mon blue (Polyommatus icarus) butterflies to mate-searching
males. Naturwissenschaften 87:304—307.
BURGHARDT, F., P. PRoKSCH & K. FIEDLER. 2001. Flavonoid
sequestration by the common blue butterfly Polyommatus
icarus: quantitative intraspecific variation in relation to larval
hostplant, sex and body size. Biochem. Syst. Ecol.
29:875-889.
Coutsis, J. G. & N. GHavaLas. 1996. Ultra-violet reflection pattem
in Polyommatus andronicus Coutsis & Ghavalas, 1995 and Poly-
ommatus icarus (Rottemburg, 1775) (Lepidoptera: Ly-
caenidae). Phegea 24:167-169.
CuTHILL, I. C. & A. T. D. BENNETT. 1993. Mimicry and the eye of
the beholder. Proc. R. Soc. Lond. B 253:203-204.
CuTHILL, I. C., A. T. D. BENNETT, J. C. PARTRIDGE & E. J. MAIER.
1999. Plumage reflectance and the objective assessment of
avian sexual dichromatism. Am. Nat. 153:183-200.
DauMER, K. 1958. Blumenfarben, wie sie die Bienen sehen. Z.
vergl. Physiol. 41:49-110.
EGUCHI, E., K. WATANABE, T. HARTYAMA & K. YAMAMOTO. 1982. A
comparison of electrophysiologically determined spectral re-
sponses in 35 species of Lepidoptera. J. Insect Physiol.
28:675-682.
ENDLER, J. A. 1990. On the measurement and classification of
colour in studies of animal colour patterns. Biol. J. Linn. Soc.
41:315-352.
Ferris, C. D. 1973. A revision of the Colias alexandra complex
aided by UV reflectance photography with designation of a new
subspecies. J. Lepid. Soc. 27:57-73.
FLEISHMAN, L. J., E. R. Logw & M. LEAL. 1993. Ultraviolet vision
in lizards. Nature 365:397.
GEUDER, M., V. Wray, K. FIEDLER & P. PROKSCH. 1997. Seques-
tration and metabolism of host-plant flavonoids by the lycaenid
butterfly Polyommatus bellargus. J. Chem. Ecol. 23:1361-1372.
GHIRADELLA, H., D. ANESHANSLEY, T. EISNER, R. E. SILBERGLIED &
H. E. HINTON. 1972. Ultraviolet reflection of a male butterfly:
interference color caused by thin-layer elaboration of wing
scales. Science 178:1214—1217.
Gisps, G. W. 1980. Reinstatement of a New Zealand copper but-
terfly, Lycaena rauparaha (Fereday, 1877). N. Z. J. Zool.
7:105-114.
HaRBORNE, J. B. 1991. Flavonoid pigments, pp. 389-429. In G. A.
ROSENTHAL & M. R. BERENBAUM (eds.), Herbivores, their in-
teractions with secondary plant metabolites. Vol. 1: The chemi-
cal participants. Academic Press, San Diego.
(ed.). 1999. The flavonoids. Chapman & Hall, London. XII.
676 pp.
Jacoss, G. H. 1992. Ultraviolet vision in vertebrates. Amer. Zool.
32:544-554.
KELBER, A. & M. PraFr. 1999. True colour vision in the orchard
butterfly, Papilio aegeus. Naturwissenschaften 86:221-224.
KNUTTEL, H. & K. FIEDLER. 1999. Flavonoids from larval food
plants determine UV wing patterns in Polyommatus icarus
(Lepidoptera: Lycaenidae). Zoology 102 (Suppl. 2):83.
SS ERO RTE SS IS EE
144
KNUTTEL, H. & K. FIEDLER. 2001. Host-plant-derived variation in
ultraviolet wing patterns influences mate selection by male but-
terflies. J. Exp. Biol. 204:2447-2459.
KORNMAIER, B. 1999. Hinlagerung von Flavonoiden in die Fliige-
lanlagen wihrend der Puppenphase bei Polyommatus icarus
(Lepidoptera: Lycaenidae). Unpublished diploma thesis, Uni-
versitat Bayreuth, Germany. IV, 120 pp.
Kuprna, O. 1975. A revision of the genus Gonepteryx Leach
(Lep., Pieridae). Entomologist’s Gaz. 26:3-37.
. 1992. On the hidden wing pattern in European species of
the genus Colias Fabricius, 1807 (Lepidoptera: Pieridae) and its
possible taxonomic significance. Entomologist’s Gaz. 43:167—-176.
Lunau, K. & E. J. Mater. 1995. Innate colour preferences of
flower visitors. J. Comp. Physiol. A 177:1-19.
MENZEL, R. & W. BAckHaus. 1991. Colour vision in insects, pp.
262-293. In P. Gouras (ed.), Vision and visual dysfunction: per-
ception of colour. Vol. 6. Macmillan, Houndsmills, UK.
MEYER-RocHow, V. B. 1991. Differences in ultraviolet wing pat-
terns in the New Zealand lycaenid butterflies Lycaena salustius,
L. rauparaha, and L. feredayi as a likely isolating mechanism. J.
R. Soc. N. Z. 21:169-177.
NEKRUTENKO, Y. P. 1964. The hidden wing-pattern of some
Palaearctic species of Gonepteryx and its taxonomic value. J.
Res. Lepid. 3:65-68.
. 1968. Phylogeny and geographical distribution of the
genus Gonepteryx Leach (1815). Kiev. 128 pp., 20 pls.
OorscHot, H. vAN & W. O. DE PRINS. 1989. Some critical remarks
on the paper “Unterschiede von Lycaena hippothoe und can-
dens im UYV-Licht (Lep., Lycaenidae)” by P. Schaider (1988).
Phegea 17:53-54.
Rutowski, R. L. 1977. The use of visual cues in sexual and species
discrimination by males of the Small Sulphur Butterfly Ewrema
lisa (Lepidoptera, Pieridae). J. Comp. Physiol. A 115:61—-74.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
. 1981. Sexual discrimination using visual cues in the check-
ered white butterfly (Pieris protodice). Z. Tierpsychol.
595:325-334.
SCHAIDER, P. 1988. Unterschiede von Lycaena hippothoe und can-
dens im UV-Licht (Lep., Lycaenidae). Atalanta 18:415-425.
SCHITTKO, U., F. BURGHARDT, K. FIEDLER, V. Wray & P. PROKSCH.
1999. Sequestration and distribution of flavonoids in the com-
mon blue butterfly Polyommatus icarus reared on Trifolium
repens. Phytochemistry 51:609-614.
SILBERGLIED, R. E. 1984. Visual communication and sexual selec-
tion among butterflies, pp. 207-223. In R. I. Vane-Wright & P.
R. Ackery (eds.), The biology of butterflies. Academic Press,
London.
SILBERGLIED, R. E. & O. R. TayLor, JR. 1973. Ultraviolet differ-
ences between the sulphur butterflies, Colias eurytheme and
C. philodice, and a possible isolating mechanism. Nature
241:406-408.
. 1978. Ultraviolet reflection and its behavioral role in the
courtship of the sulfur butterflies Colias ewrytheme and C.
philodice (Lepidoptera, Pieridae). Behav. Ecol. Sociobiol.
3:203-243.
Tove, M. J. 1995. Ultra-violet photoreceptors in the animal king-
dom: their distribution and function. Trends Ecol. Evol.
10:455-460.
WIESEN, B., E. KRUG, K. FIEDLER, V. Wray & P. Prokscu. 1994.
Sequestration of host-plant derived flavonoids by lycaenid but-
terfly Polyommatus icarus. J. Chem. Ecol. 20:2523-2538.
WILSON, A. 1987. Flavonoid pigments in chalkhill blue (Lysandra
choridon Poda) and other lycaenid butterflies. J. Chem. Ecol.
13:473-493.
Received for publication 3 December 1999; revised and accepted
June 25 2001.
GENERAL NOTES
Journal of the Lepidopterists’ Society
54(4), 2000, 145-146
LARVAL DESCRIPTION AND HOSTPLANT RECORD FOR ITHOMIA DIASIA HEWITSON
(NYMPHALIDAE: ITHOMIINAE) IN PANAMA
Additional key words: pupa, hippocrenis, Witheringia, Solanaceae.
Despite the fact that butterflies in the genus Ithomia Hiibner
participate in well documented mimicry rings (e.g., Beccaloni 1997)
and are therefore important players in community level interactions
that characterize the Neotropics, information on their larval biology
is scarce. All known Ithomia immatures feed on Solanaceae (Table
1, Brown and Freitas 1994, G. Beccaloni pers. comm.). We found no
published host plants records for the remaining Central American
species of Ithomia, and brief descriptions of immatures are available
only for I. patilla and I. heraldica (see DeVries 1987).
On 18 February 1995 (dry season) we collected a fourth instar
larva of Ithomia diasia hippocrenis Bates on Witheringia aster-
otricha (Standl.) (Solanaceae) on the Atlantic slope just below the
Continental Divide near E] Cope, Panama (800 m elevation). The
range of I. diasia hippocrenis includes Panama and Costa Rica, and
in Costa Rica it is distributed from sea-level to 1,400 m on the At-
lantic slope, reaching greater abundance in the dry season (DeVries
1987). On 2 March 1995, an adult male emerged from the pupa, and
was subsequently compared to specimens in the collection of the
Natural History Museum, London (BMNH). The emerged male
was similar to two specimens from highland Panama except for the
black (rather than brown) triangular marking from the forewing
costal margin to the distal end of the discal cell in the ventral surface
of the forewing (see photograph of I. diasia in DeVries 1987). Of the
20-30 specimens of I. diasia hippocrenis at the BMNH, only one
from Costa Rica also had the same black triangular marking, and we
expect that this represents a rare variant of the phenotype.
The host plant, Witheringia asterotricha, occurs in moist upland
forest, and the plants have a purplish appearance caused by purplish
dendroid hairs on stems and leaves (visible in Fig. 2) and a bluish
cast to the epidermis (D’Arcy 1973). D’Arcy (1973) considered
Witheringia asterotricha to be a variety, of W. solanaceae. Hybrids of
these two species are reported to occur in disturbed areas (D’Arcy
1973).
In the laboratory the larva was reared in a plastic container at ca.
23°C, and fed ad libitum on leaves that were kept in a plastic bag in
the refrigerator. Note that developmental time in the laboratory is
TABLE 1. Host plant records of Ithomia from
Central America.
Species Host plant References
I. patilla Witherengia solanacea Drummond & Brown 1987
Witherengia sp. Drummond & Brown 1987
Lycianthes multiflora Drummond & Brown 1987
I. xenos W. cuneata Drummond & Brown 1987
Acnistus arborescens | Drummond & Brown 1987
Cuatresia riparia Drummond & Brown 1987
I. iphianassa_ C. riparia Drummond & Brown 1987
C. morii Drummond & Brown 1987
A. arborescens Drummond & Brown 1987
I. celemia C. riparia Drummond & Brown 1987
I. diasia L. heteroclita DeVries 1985
W. solanacea Drummond & Brown 1987
W. asterotricha Srygley & Penz 2000
Fic. 1. Fifth instar larva of Ithomia diasia from Panama. Fic. 2.
Pupa of Ithomia diasia from Panama.
probably distinct from that in nature due to differences in tempera-
ture regime.
Larva. Fourth instar (4 days, n = 1). Head light gray with a black
transverse stripe that starts above front and ends below stemmata,
stemmata black, mandibles brown; body translucent gray with
broad, opaque, light gray markings across segments resulting in
banded pattern; broken, longitudinal yellow spiracular line com-
posed of color patches of uneven size; spiracles black; ventrally
transparent (tracheal system visible); thoracic legs and prolegs
translucent gray. Fifth instar (Fig 1, 3 days, n = 1). Head same as
fourth instar although more translucent in color; body black with
faint banded pattern on the posterior portion of each segment, three
wrinkles across segments T2—A8; broken, longitudinal yellow spirac-
146
ular line composed of color patches of uneven size; anal cap gray;
ventrally translucent gray; thoracic legs and prolegs translucent gray.
Pupa (Fig. 2, 5 days, n = 1). Short, slightly compressed laterally,
translucent green with patches of iridescent gold. Head with two small
gold bumps just above eye; sutures of mouthparts and antennae dark
brown; antennae with dark brown spots on each segment; T2 with gold
dorsal keel and a pair of brown, dorso-lateral spots; gold patches later-
ally on T1-3; brown spots at the base and in the center of wing pad;
four brown spots near distal margin of wing pad; T3—A8 with a pair of
brown dorso-lateral spots; abdomen with gold dorsal midline band;
large lateral gold patches on A1—8; spiracles brown; cremaster brown.
The broad black frontal stripe present in I. diasia fifth instar also
occurs in other species of Ithomia; the broken sublateral band in the
fifth instar and partially bent pupa are characteristic of the Ithomiinae
tribes Ithomiini, Napeogenini and Oleriini (A. Freitas pers. comm.).
We thank G. Beccaloni for helping with identification of the
adult, and G. Beccaloni and A. Freitas for commenting on the man-
uscript. Autoridad Nacional del Ambiente (ANAM) graciously
granted permission to collect butterflies in the Republic of Panama.
The adult voucher has been deposited in the Museo Fairchild of the
Universidad de Panama. This research was supported by the Na-
tional Geographic Society (to RBS), a Smithsonian Tropical Re-
search Institute (a pre-doctoral fellowship to CMP), and the Na-
tional Science Foundation (NSF DEB-9806779 to CMP).
LITERATURE CITED
BECCALONI, G. W. 1997. Vertical stratification of ithomiine butter-
fly (Nymphalidae:Ithomiinae) mimicry complexes: the relation-
ship between adult flight height and larval host-plant height.
Biol. J. Linn. Soc. 62:313-341.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Brown, K. S. JR. & A. V. L. Freitas. 1994. Juvenile stages of
Ithomiinae: overview and systematics (Lepidoptera: Nymphali-
dae). Trop. Lepid. 5 (1):9-20.
D'Arcy, W. G. 1973. Solanaceae. In Flora of Panama. Ann. Mis-
souri Bot. Gard. 60:573-780.
DEvRIES, P. J. 1985. Hostplant records and natural history notes on
Costa Rican butterflies (Papilionidae, Pieridae & Nymphali-
dae). J. Res. Lepid. 24 (4):290-333.
. 1987. Butterflies of Costa Rica and Their Natural History.
Princeton Univ. Press, Princeton, New Jersey.
Drummonp, B. A. & K. S. Brown, JR. 1987. Ithomiinae (Lepi-
doptera: Nymphalidae): summary of known larval food plants.
Ann. Missouri Bot. Gard. 74:341-358.
MERBEL, V. 1982. Host plant finding and oviposition behavior of
Ithomia heraldica Bates (Lepidoptera: Ithomiidae) at Mon-
teverde. Brenesia 19/20:369-372.
Rosert B. SryGLEy, Department of Zoology, South Parks Road,
University of Oxford, Oxford OXI 3PS United Kingdom and
Smithsonian Tropical Research Institute, Apdo. 2072, Balboa,
Republic of Panama, AND CARLA M. PENZ, Department of
Invertebrate Zoology, Milwaukee Public Museum, 800 West Wells
Street, Milwaukee, Wisconsin 53233, USA and Curso de Pos-
Graduagdao em Biociéncias, Pontifcia Universidade Catolica do Rio
Grande do Sul, Av. Ipiranga 6681, Porto Alegre, RS 90619-900,
Brazil.
Received for publication 28 October 1999; revised and accepted 15
March 2001.
BOOK REVIEWS
Journal of the Lepidopterists’ Society
54(4), 2000, 147
INSECTORUM SIVE MINIMORUM ANIMALIUM THEATRUM: THE BUTTER-
FLIES AND Motus, by George Thomson. Published in 2000. Page size
7 3/4 by 11 1/2 inches. (2) + 66 pages, with numerous line illustrations
and two colored plates pasted in. Hard covers, with dust wrapper. No
ISBN, price £65 (approx $100). Limited edition, privately published
by the author, and obtainable from Dr George Thomson, 2 Ravenhill,
Lochmaben, Lockerbie, Scotland DG11 1QZ, UK.
Despite its many shortcomings, Thomas Moufet’s Insectorum
Theatrum is important as being the first entomology book to have
been published in Britain, appearing in its original Latin version in
1634, and in English translation in 1658 although the original
manuscript dates from the late 16th century. That the work is
effectively the product of several authors, including Gesner, Penny
and Wotton, is well known and has been widely discussed before,
and we are fortunate that the original manuscript of the book is still
extant in the British Library.
Thomson has here attempted to give an introduction to the book
and its history, with a particular study of the adult Lepidoptera,
which form only a small part of the whole Theatrum. He begins with
a section on the manuscript and its origins, an account of Thomas
Moufet and the other contributors, and a brief note on the
illustrations. These are followed by the text of the English
translation of the “Butterflies” chapter of the book, with the
illustrations inserted in their correct places. Thomson has made
useful comparisons of the mostly rather crude published woodcuts
with the often very accurate and beautiful original paintings that are
pasted in the margins of the manuscript. Each illustration is
annotated with its modern scientific and common name, with a note
on its occurrence in the original manuscript, the Latin text and the
English text. After a brief bibliography a facsimile of the Latin
version of the “De Papilionibus” chapter occupies 22 pages, and the
book ends with a note about a butterfly specimen found between
two pages of the original manuscript.
Although Thomson mentions that two chapters of Moufet'’s book
contain many descriptions of lepidopterous larvae he does not
include them, which is a sad omission as many could have been
easily identified. He rightly states that the adults are not given
names, but omits to state that many of the larvae are named, and
they provide a useful insight into the origins of scientific names later
given to some species.
The 82 forms described in the original work comprise 56 species,
consisting of 30 moths, 25 butterflies, and one unidentifiable taxon.
Five of the species are not found in Britain, although four of these
occur elsewhere in Europe. The exception is the well-known North
American Papilio glaucus, and a colored reproduction of John
White's watercolour painting of this species forms a frontispiece to
Thomson's book.
The historical sections of this book are all based on secondary
sources, such as Lisney (1960) and Raven (1947), and, excellent
though these books are, they are no substitute for more detailed
works such as the study of Thomas Penny by W. Gardner (1930. A
Lancashire entomologist in the time of Queen Elizabeth. Transactions
of the Lancashire and Cheshire Entomological Society 1931: 31-52)
which is not mentioned here. Thomson's paraphrasing of secondary
sources has led to ludicrous errors, such as stating that Moufet
“accompanied Peregrine Bertie and Lord Willoughby to Elsinore”
without realizing that these “two” men are one and the same!
Having described Rowland’s English translation (published by
Topsell) of the Theatrum as “somewhat inaccurate” and “rather
poor’ we might take this to mean that Thomson could have
improved on it; however, he gives the translation in full without
comment and we have to assume that his critical remarks on
Topsell’s edition are merely quoted from other sources.
This is a privately published book of 500 copies, well printed on
good quality paper in an attractive binding, and it is inevitable that
such a limited edition book will sell for a high price. However, readers
expect a certain quality of content for their money and unfortunately
this book falls into the familiar trap of many privately printed works in
that it would clearly have benefited from professional editing. There
are too many errors to list in this review; some are minor
typographical mistakes but others cannot be so easily ignored and are
very prominent. For example, the section on Theodore Mayerne
bears the heading “Sir Thomas Mayerme”, a name by which he was
never known (this even appears in the contents list!). The two pages
outlining Thomas Moufet's life have at least nine mistakes, and even
the transcription of the Topsell translation contains a great many
errors, with several misspelled scientific names (Aglais is spelled three
different ways) and even the wrong authorship of one species
(Euphydryas aurinia, mis-spelled aurinea, is attributed to Linnaeus
rather than Rottemburg). Altogether I have noted over 60 errors in 42
pages of text (not including the facsimile section) which is
unacceptably high in such an expensive book.
The final brief note on a butterfly found between the pages of the
manuscript is accompanied by a colored illustration of the specimen.
Thomson speculates on the possibility that the specimen may be
contemporary with the manuscript, and he makes comparisons of
the style of preservation with some known early collections such as
that of James Petiver. Unfortunately he seems to be unaware of the
published work on these collections, e.g., M. Fitton and P. Gilbert
(1994. Insect collections. In A. MacGregor (ed.), Sir Hans Sloane.
British Museum Press, London: pp. 112-122).
When I first picked up this book I had high hopes that it would
be an original and stimulating work, but sadly those hopes have not
been fulfilled. If the author had confined himself to describing the
Lepidoptera in the Theatrum then this book would have had some
value, but even then more care should have been taken with
accuracy, and the inclusion of the larval chapters would have
enhanced it considerably. But as a supposed historical study it has so
many errors that it will be a disappointment to the expert, and
potentially misleading to the novice.
P. C. BARNARD, Department of Entomology, The Natural History
Museum, Cromwell Road, London SW7 5BD, UK.
148
Journal of the Lepidopterists’ Society
54(4), 2000, 148
MICROLEPIDOPTERA OF EUROPE, VOLUME 2, SCYTHRIDIDAE, by
Bengt A. Bengtsson. 1997. Published by Apollo Books, Stenstrup,
Denmark. 301 pp, 14 color plates, 419 text figures. Hard cover, 25 x
17 cm, ISBN 87-88757-11-0. Available from Apollo Books ApS,
Kirkeby Sand 19, DK-5771, Denmark, fax +45 62 26 37 80,
www.apollobooks.com. 550 Danish Kroners + postage.
Fifteen years ago, Bengtsson published a monograph of the
Scythrididae of northem Europe (1984, Fauna Entomologica
Scandinavica, v. 13), which treated a limited fauna of 33 species.
With this new book, the second in the excellent series Micro-
lepidoptera of Europe, Bengtsson has expanded his coverage to
include the entire European fauna east of the Urals, the
Mediterranean basin, Turkey, and the Canary Islands. The
Scythrididae have thus been the object of two major works in the
span of 15 years, both by the same author, which is rather remarkable
for a family of (figuratively and actually) small and obscure moths.
The present treatment includes 237 species in 7 genera, with 40
species described as new. A number of new synonyms are also
introduced. The book begins with a general introductory section
with parts about Scythrididae morphology, practical hints for
identification, genitalia preparation and collecting, biology, as well
as brief comments on classification and geographical distribution
of the family. This is followed with a checklist of all taxa treated
with full synonymy, and then the systematic treatment, which
makes up the bulk of the work. Individual species treatments
include a diagnosis, a brief description of genitalia of both sexes,
and a summary of known geographical distribution and biology.
The host plant and biology of very few species is known. The
geographical distribution of all species is conveniently summarized
in a dense table placed at the end of the systematic section and
lists occurrences by country; the table is useful albeit somewhat
difficult to read with its 53 columns and 237 lines. The book
concludes with an extensive reference list and an index to taxon
names.
The most remarkable feature of the book is the 14 color plates,
which present 258 superb watercolor paintings of adults (body and set
right wings), painted by the author. Scythrids are rather dull-colored
moths, many have dark brown coloration with a metallic sheen whose
hue varies with the species and with the lighting, making these moths
a real challenge to illustrate. The fine rendering of all subtle shades of
greys, browns, beiges, off-whites as well as metallic hues is simply
remarkable and attests to the great artistic talent of Bengtsson. I can
only marvel at the countless hours that were required to produce all
these watercolors, a benedictine labor of love.
The genitalia of all species are illustrated with line drawings (419
of them!). Many scythrid species are confusingly similar moths with
strikingly different genitalia; conversely several species display a
great amount of color variation depending on seasonality and region.
Thus genitalia examination will be essential for species identification
in many cases, unless one has a fair amount of experience with these
moths and their subtle color differences. Curiously, however, the
line drawings in this book are of a lesser quality than those produced
by Bentgsson in his 1984 work on the northern European species.
Generally, the drawings do not render the impression of depth and
structural complexity exhibited by the genitalia of scythrids,
although they display diagnostic features sufficiently well for species
recognition, including arrows pointing at specific features on many.
Most illustrations of male genitalia were drawn from standard
flattened, dorso-ventral slide mounts, which are often inadequate for
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
preserving the highly tri-dimensional, and fused scythrid genitalia
because they result in a fair amount of distortion making
interpretation of parts difficult.
A conspicuous omission in this otherwise fine work is an
identification key. This is a significant deficiency considering the large
number of species treated (237), most of which belong to the genus
Scythris Hiibner (204 spp.). Species are arranged into species-groups
for which diagnoses are provided, but the user wanting to identify
specimens has little choice but to wade through the entire set of
illustrations to try to match specimens at hand, then go to the
diagnoses to read diagnostic features. In some species-groups where
similarities among species are high, the process is laborious.
The arrangement of the taxa is somewhat perplexing. On p. 14,
paragraph 2, it is stated that species are arranged alphabetically
which, although debatable in its value, is understandable. However,
this is contradicted largely by the broad arrangement of the genus
Scythris, by far the largest genus, for which the author states that a
“logical order” was used to arrange the species. Actually, the
alphabetical order is found only within species-groups. Bengtsson
explains his so-called “logical order” as follows (bottom of p. 13): “In
order to arrange species in a logical order, the author has based the
classification on male genital structure together with the female
genital structure and other characters when known.” It is not clear
what “logical order” means in biological classification.
The short section entitled “Systematics and classification” on pp.
13-14 explains the arrangement of taxa in the book. One is left with
the impression that the author could have either expanded some of
the points mentioned or omitted them altogether. For example, the
statement that (p. 14) “Some of the remaining genera will probably
be synonymized in the future when the whole world fauna can be
analysed” is difficult to appreciate by the reader because there is
little in the book itself to help evaluate this. Bengtsson also discusses
his undocumented and unsatisfactory attempt at cladistic analysis:
“In addition, species-groups are introduced, based on assumed
synapomorphies or similarities of uncertain phylogenetic value. The
author has used the computer program Hennig86 for analysis of the
phylogeny, including a limited number of species at each evaluation.
Starting with a complex and detailed data matrix the author has
successively simplified the matrix until the most parsimonious tree
structure approximately agreed with the intuitive opinion. Even so,
the number of possible trees produced is very high.[. . .] . . . the
phylogeny of the Scythrididae is extremely complicated. Instead of
trying to utilise a systematic sequence in the present work, I have
dealt with the species in alphabetic order.” Because none of the data
are presented, we are left with no basis to appreciate how the
classification was developed. However, this is an academic point
unlikely to bother the average user.
The English style is not always standard, being a little awkward in
places and occasionally muddling the intended meaning or clarity,
however, I did not find that this reduced the usability of the book.
The book is of high quality, printed on glossy paper, clearly typeset,
and has an attractive cover. The Smythe-sown binding allows the
book to lay flat-open and will contribute to its durability.
Despite the criticisms expressed above, I can only praise
Bengtsson for his accomplishment. I recommend unconditionally the
book to any serious lepidopterist, if only because it provides a
comprehensive treatment of a family for a large geographical region of
the world and is part of a series (Microlepidoptera of Europe) that is
akin to a counterpart of the Moths of North America (MONA) series.
JEAN-FRANGOIS LANDRY, Agriculture and Agri-Food Canada, 960
Carling Ave, Ottawa, Ontario K1A 0C6, Canada.
Journdl of the Lepidopterists’ Society
54(4), 2000, 149-150
ADELPHA
cocala, 97
mythra, 97
syma, 97
adult diet, 91
Aiello, A., 97
allometry, 111
“Antigonus” Genus Group, |
Asteraceae, 97
Austin, G. T., 1
Balough, G. J., 102
Barnard, P. C., 147
behavior, 78, 88, 96
Behemothia, 41
biology, 97
body size, 111
Boreal, 47
Brazil, 1, 97
Brower, A. V. Z., 110
Brown, K., 97
Brown, R. L., 111
Burns, J. M., 52
bursa copulatirix, 83
Callaghan, C. J., 119
Carrhenes
recurva, 22
caterpillar behavior, 77, 145
Catocala
marmorata, 107
Charlton, R. E., 96
cladistics, 41
Colombia, 119
color, 137
Cook, R. P., 33
Costa Rica, 119
Crambidae, 72
Danaus
plexippus, 29
distribution, 52
diurnal feeding behavior, 96
Doleschallia tongana, 33
early stages, 98, 107, 145
ecology, 33
Ecuador, 88
egg load, 83
egg maturation, 83
Epargyreus
clarus, 77
Fabaceae, 72, 77
fecundity, 91
Fiedler, K., 91, 137
Fischer, K., 91, 107, 145
food plants, 97, 107, 145
Freitas, A. V. L., 97
INDEX FOR VOLUME 54
(new names in boldface)
Gall, L. F, 111
genitalia, 1, 44, 48, 50, 52, 72
Gentili, P., 72
geographic distribution, 52
Gerardo, N. M., 88
Greeney, H. F., 36, 88
Guatemala, 1, 119
Hall, J. P. W., 41
Hawaii, 29
herbivore-plant interactions, 131
Hesperiidae, 1, 52, 77
Hodges, R. W., 39
Holarctic, 47
Honduras, 119
host plant, 97, 131, 137, 145
Hypophylla
argenissa, 119
caldensis, 119
flora, 119
idea, 119
lasthenes, 119
martia, 119
sudias, 119
sudias callaghani, 119
zeurippa, 119
immature stages, 88, 96, 111
Indiana, 111
Ithomia diasia hippocrenis, 145
Ithomiinae, 145
Jalava, J., 47
Japan, 83
Jones, M. T., 77
Kansas, 96
Kasuya, M., 29
Kniittel, H., 137
Kopper, B. J., 96
Landry, J.-F., 104
larva, 88, 96, 97, 145
larval feeding, 131
Lauraceae, 131
leaf folder, 77
leaf shelter, 77
life history, 33, 97, 111
Limenitidinae, 97
Lind, E. M., 77
Long, J. D., Ui
longevity, 91
Lycaena
hippothoe, 91
phlaeas, 83
Lycaenidae, 83, 91, 137
Magnoliaceae, 131
marbled underwing, 111
Margolies, D. C., 96
Margraf, N., 145
mate-avoidance, 83
Matthews, D. L., 36
Mexico, 119
Miller, W. E., 47, 111
monarch butterfly, 29
monogamous, 83
morphology, 41, 72, 119
Mylon
argonautarum, 15
cristata, 17
simplex, 11
Myricaceae, 131
Nearctic, 47
Neotropics, 1, 41, 88, 97
new genera, 4]
new species, 1, 73, 119, 126
new synonymy, 47
Nicaragua, 119
Nicolay, S. S., 76
Nishimura, M., 83
nitrogen, 111
Noctuidae, 107
Notocelia
rosaecolana, 111
trimaculana, 111
nutritional ecology, 91
Nymphalidae, 29, 33, 88, 96, 97, 145
Nymphidiini, 41
Olethreutinae, 111
Omiodes
pseudocuniculalis, 72
ontogenetic changes, 77
oviposition behavior, 88
oviposition preference, 132
Palaearctic, 47
palamedes swallowtail, 131
Panama, 119, 145
Papilio
palamedes, 131
glaucus, 131
Papilionidae, 131
parategumental sclerites,
Peacock, J. W., 107
Penz, C. M., 145
perching behavior, 119
phenology, 1
phenotypic plasticity, 137
phytochemical toxicity, 131
polymorphism, 29
Polyommatus icarus, 137
polyphagous species, 131
population dynamics, 91
Populus heterophylla, 107
Protonymphidia, 43
7
150
pupa, 88
purple-edged copper, 91
Pyraloidea, 72
Pyrginae, 1, 52
Pyrgus
albescens, 52
communis, 52
Pyrrhogyra
crameri, 88
otolais, 88
range extension, 33
regal fritillary, 96
reproduction, 91
reproductive output, 83
Riodinidae, 41, 119
Rosaceae, 97
Rubiaceae, 97
Salicaceae, 107
Samoan Archipelago, 33
Sapindaceae, 88
Scriber, J. M., 131
Serjania, 88
Shapiro, A. M., 38
silver-spotted skipper,
small copper, 83
Solanaceae, 145
Solis, M. A., 72
South America,
speciation, 52
spermatophore, 83
v7
(
(1
72
Speyeria
idalia, 96
spicebush swallowtail, 131
Srygley, R. B., 146
stenophagous species, 131
Stimson, J., 29
systematics, 52
tallgrass prairie, 96
taxonomy, 1, 119, 137
Date of Issue (Vol. 54, No. 4): 10 December 2001
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
temperature, 91
Theaceae, 131
Theopeiti, 41
tiger swallowtail, 131
Tortricidae, 47, 111
Trans-Andean region, 119
Tuck, K., 111
ultraviolet photography, 137
Underwing moths, 107
Vargo, D., 33
variation, 1, 52
visual communication, 137
Watanabe, M. 83
Weiss, M. R., 77
Wells, T., 131
Western Hemisphere, 72
white morph, 29
Witheringia, 145
EDITORIAL STAFF OF THE JOURNAL
M. Deane Bowers, Editor
Entomology Section
University of Colorado Museum
Campus Box 218
‘University of Colorado
Boulder, Colorado 80309, U.S.A.
~ email: bowers@spot.colorado.edu
- Associate Editors:
casas LAmas (Peru), Kenetm W. Pamir (USA), Rosrrt K. Rossms (USA),
Fexix A. H. Speruine ( USA), Davin L. Wacner (USA), Curister WikLunp (Sweden)
rai Seats NOTICE TO CONTRIBUTORS
ae eet may dedi with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Technical Comments,
5, Obituaries, Feature Photographs, and ois Illustrations. Obituaries must be authorized by the President of the Society. Requirements
he manuscripts: concerning new state Boats current events, and notices sbeuld nae sent to the News, Phil Schappert, Dept. Zoology, Univ.
: 78712. Requests to review ee for the ae and pone review isp aettlts should be sent to M. Alma Solis, Syst. Entomol. Lab.
An iicrmiative abstract siould precede the text of Articles and Profiles.
y words: Up to five key words or terms not in the title should accompany Articles, Profiles, General Notes, and Technical Com-
~ Contributors should write with precision, clarity, and economy, and should use the active voice dea first person whenever appropriate. Titles
, descriptive, and as short as possible. The first mention of a plant or animal in the text should include the full scientific name with
ily. Measurements should be given in metric units; times in terms of the 24-hour clock ed h, not 9: Peel AM). Underline only, where
ca
959, ES caleclion and vaueae 2nd ed. Hutchinson, London. 209 pp:
e contributions to population genetics resulting from the study of the Lepidoptera. Ady. Genet. 10:165-216.
“Only half of symmetrical objects such as adults with wings spread should be illustrated, unless whole illustration is crucial. Pho-
s should be mounted on stiff, white backing, arranged in the desired format, allowing (with particular regard to lettering) for re-
a Journal page. Illustrations larger than letter-size are not acceptable and should be reduced photographically to that size or smaller. The
ure e numbers a as cited i in the text ioe be printed on od back of each illustration, Figures, both line Rea and hrtoseel
ae hols be Oe:
Spee amsen: When appropriate manuscripts must name a oe cae: where specimens ge identity of iar can be
yr reason should oe to the editor at the time of Siti for. a reduced rate or free pu bliewtion! aches of Book Reviews a
ies are exempt from. page charges.
eae BUSES all matters relating to the Journal to the editor.
- PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A,
ls 3
| Suman OF FOUR. AMILIES OF see ‘say SPECIES FOR.
cribei Nicolas Margraf, and Tammy Wells ues
| eae! Bitsy Ont THE USE oF “ULTRAVIOLET PHOTOGRAPHY AND ULTRAVIOLET wine ATTERN .
My ee
\
‘
ie : i
fs
ay
|
%
2)
Fave
; tes ee x Number 1
V7 assiv 0024-0966
rly by THE LEPIDOPTERISTS’ SOCIETY
LA SOGMTEDESLEPIDOPIBRISTES 9). 8) Nt
| : HAF ER EPIDOPT HOGOCEN (os, ac
CIEDAD DE LOS LEPIDOPTEROL ae
‘THE LEPIDOPTERISTS’ SOCIETY. noe
EXECUTIVE Counen.
Joun W. Brown, President ~~ cA Gates Sega Se Bom Vice President
Micnae J. Smira, Immediate Past President = ~Mirna M. CasaGRanbe, Vice President — a, é
Manuex A. Barcazar-Lara, Vice President Davyip Sp TrTNer, Treasurer
Ernest H. WituiAMs, Secretary :
aah ee : Members at large: ates
Ronald L. Rutowski ~ M. Deane Bowers George Es Balogh”
Felix A. H. Sperling ss Ron Leuschner) ~ Andrew V, Z. Brower —
. Andrew D. Warren gee ae Pagetns Brian Scholtens
Epirortat Boarp>
Repos K. Rossins (Chairman), Joun W. Brown (Member at large)
~ Carva M. Penz (Journal)
WituiAM E. Minter (Memoirs)
—Parur J. ScHarPertT os
Honorary Lire MEMBERS OF THE Society
: are L. Re tinectON (1966), E.. G. Munror (1973), IAn F. B. Caran | (1987).
Joun G. F RANCLEMONT (1988), Lincoun P. Brower (1990), Douc.as C. Fercuson (1990),
Hon. Miriam RoriscHitp (1991), Craupe Lemaire: O92) F REDERICK ay RinpcE ae
The object of The Lepidopterists’ Society: which was ferineda in May 1947 and ae ciagaaads in Decne 1950
mote the science of lepidopterology in all its branches, . . . to issue a periodical and other publications on Lepidoptera, t (
the exchange of specimens and ideas by both the professional worker and the amateur in the joes to secure aga
sures’ ’ directed towards these aims.
gether with their full name, address: and es ee inter ests. In aftemate years alist af members of the S
sued, with addresses and special interests.
Active Henbene ee heed Ae $45.00 hae the U.S., $50. 00 sl i vu. S.
Affiliated members—annual dues $10.00 within the U.S., $15.00 outside the U. eee
- Student members—annual dues $20.00 within the U.S., $95, 00 outside the U.S.
Sustaining members—annual dues $60.00 within the U.S., $65.00 outside the U. Se
Life members—single sum $1,800.00
Institutional subscriptions—annual 60.00:
Airmail postage for the News $15.00
‘path
Send remittances, payable to The Lepidopterists’ Society, to: Kelly M. Richers, Asst. Treasurer 9417 Carvalho Court, a ersfiel
CA 93311; and address uae to: i eas E Donahue, Natural nga Museum, 900 eg Blvd., Los fee CA eee :
field, CA 93311.
The additonal cost for members outside the U.S, is to cover mailing costs.
ee Ee The Lepidantorisrs: Society (ISSN 0024-0966) is pital quarterly by The Tote Snoiely: > Los oe Ae
County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angele
and at additional mailing offices. POSTMASTER: Send address ee to The Peron Society, “fo Natural ee Muse
900 Exposition Blyd., a Angeles,’ CA 90007- 4057. ;
t
Cover illustration: Sphinx frankii ecient (Sphingidae), female genitalia. Pes and ink ms laine R. Ss. §. Hodges From: Hodges
R. W. 1971. Moths of America North of Mexico. Sphingoidea, Fascicle 21. cokes .
JOURNAL OF
ire WEPIDOPTERISTS’ SOCIETY
Volume 55
2001 Number 1
Journal of the Lepidopterists’ Society
55(1), 2001, 1-7
PRESIDENTIAL ADDRESS, 2000: NOMENCLATURAL NONSENSE—
FLYING IN THE FACE OF A FARCICAL CODE
JOHN W. BROWN
Systematic Entomology Laboratory, PSI, ARS, USDA
c/o National Museum of Natural History,
Washington, DC 20560-0168
e-mail: jbrown@sel.barc.usda.gov
[The following is a modified transcript of the presidential address
delivered at the annual meeting of the Lepidopterists’ Society,
Wake Forest University, Winston-Salem, North Carolina, 29 July
2000. The original title was “Everything You Always Wanted to
Know About Nomenclature, But Were Afraid to Ask.” You may re-
create the mood of the banquet by waiting until late in the evening,
eating a large meal, and drinking several glasses of wine before
reading. |
The presidential address typically represents one of
those points in the meeting when time... seems. . . to
. stand . . . still, especially for spouses and others
who have been coerced into attending the meeting, or
at least the banquet. And you know who you are. Well,
Yl try my best to be mercifully brief. But I'll warn you
right now, you're going to have to pay attention be-
cause there are a couple quizzes during the talk and
there’s a test at the end.
Well, if you ask any biology student what the most
boring and mundane topic is that he or she has had to
endure as part of his or her undergraduate or graduate
career, most will answer with little hesitation that
nomenclature and/or taxonomy are absolutely the
worst. I mean, what could be more boring than study-
ing the rules, regulations, and recommendations goy-
erning the formation and use of scientific names . . . in
Latin? Well, this evening I hope to demonstrate to you
that although the study and practice of taxonomy and
nomenclature may seem boring, it actually may be joy-
ous, intriguing, fascinating, and entertaining . . . or at
least not as boring as it seems. So, if I can have the first
carousel we'll get started. Don't worry, there’s only one
carousel. Actually, there’s only about 50 slides; so, if you
want, you can keep track of how near we are to the end.
Well, before I get started, allow me to digress . . .
but just briefly, of course. Well, nowadays everybody
uses a software package called PowerPoint® to make
spiffy slides for presentations (Fig. 1), and I’m no dif-
ferent. And when I’m preparing my slides for a talk,
the first thing I do is try to match the subject matter of
my talk and the type of audience with the appropriate
background pattern or color scheme, and this can be
quite challenging because PowerPoint gives you a ton
of snappy templates upon which to build your presen-
tation. So, for example, if my talk is going to be real
sciency, I might use a template like Fig. 2, matching
the intellectual quality and scholarly content of the
presentation. To me this slide just reeks “Trust me, I’m
a doctor, I know what I’m talking about.” If my talk has
a more evolutionary, ecological, or biogeographic
bend, then I might use something like Fig. 3. Here
we ve got these green and yellow eco-colors going for
us; and we've got this faint silhouette of a tree in the
background. This template says “I’m concerned with
the environment; I’m eclectic; I think globally.” If my
subject matter is going to be more high-tech, maybe
using mathematical modeling or statistics (as if), I
might use a template like Fig. 4—simple but contem-
porary. What I’m looking for here is a slide that says
“Hey, I got 1600 on the math part of my SATs and I
know a lot more about statistics than you do.” Well, fi-
nally, if I'm just going to give a regular old talk to a di-
verse audience, I might choose a template like Fig.
5—sort of plain and unpretentious, kind of under-
stated. Well, after carefully reviewing these and other
templates, I selected Fig. 6. Here we've got this little
bo
bald guy up in the corner, obviously apprehensive
about the subject matter of the talk, but we also have
this confetti action going on here, indicating that we're
going have a good time. Okay, now back to the talk.
Well, T. S. Eliot must have been a great lover of
cats, as illustrated by his book Old Possum’s Book of
Practical Cats (Eliot 1939). And this is the first stanza
of a poem from that book entitled “The Naming of
Cats.” And I'll read it to you.
The naming of cats is a difficult matter,
It isn’t just one of your holiday games;
You may think at first 'm mad as a hatter
When I tell you a cat must have three different names.
Well if old T. S. had been a lepidopterist rather than
a cat-lover, this poem may not have been that much
different, and it might have gone something like this:
The naming of moths is a difficult matter,
It isn’t just one of your holiday games;
You may think at first 'm mad as a hatter
When I tell you a moth must have two different names.
Actually, he might have left it as three if he had
worked on butterflies . . . but we won't go there.
The beginning of the “modern era” of scientific
nomenclature is typically defined by Linnaeus’ classic
treatment, Systema Naturae 10th edition, published in
1758, long before the time of T. S. Eliot. Linnaeus’
consistent use of Latin binomials—that is two names,
a genus and a species—for all organisms in Systema
Naturae established it as the “starting point” for our
modern taxonomy. If you think about it, its really
pretty remarkable to have such a well defined mile-
stone for any advancement in science, literature, or
art. And probably because of this, Linnaeus has been
dubbed “the father of modern biology’ —so this bino-
mial thing was really a pretty big deal.
But as you can imagine, it took a while for everyone
to get on-board with this two-name taxonomy; and it
wasn't until 1905 that a group of systematists drafted
the first set of rules to guide the use of scientific
names: [Fanfare] The International Code of Zoological
Nomenclature. Over the past 100 or so years, these
rules have become more standardized and rigorous
through successive editions of the Code, four in all. A
new and improved version of the Code was published
just last year. It’s a little larger than the previous edi-
tion, and the cover is a little greener. I'm not exactly
sure what the significance of the change in color is, but
you can bet that it was a hotly debated issue, as are all
issues associated with changes in the Code. We now
have this complete Code clearly describing what con-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
stitutes a valid name; defining priority, synonymy, and
homonymy; detailing what constitutes publication; and
addressing a host of other complications that may be
encountered. There is actually a Commission of Zoo-
logical Nomenclature that reviews proposals and
makes decisions regarding specific cases of usage when
controversy arises.
Scientific names are supposed to be Latin or at least
“latinized,” which is fine for those with a classic educa-
tion that included Latin. But for some of us cretins
whose only experience with Latin is pig-Latin (in
grade school), conformance with this tradition may be
a major chore. Fortunately, over the years our nomen-
clature has become contaminated with names of vari-
ous origins, including Greek, Spanish, English, and so
forth, some of which are described, even by their au-
thors, as “arbitrary combinations of letters” and by
their critics as just plain nonsense. These authors have
paved the way for those of us with limited skills in, and
knowledge of Latin to propose new names for animals
that may not be ideal, but are recognized as valid,
nonetheless. Well, finally we get to the purpose of this
address, and that is to provide you with a brief glimpse
into the rules and recommendations that apply to the
naming of animals, not just cats, relying primarily, of
course, on Lepidoptera. We're going to examine three
areas: patronyms, synonyms, and inappropriate names.
So here we go. [Slide of a playground slide] Hmmm.
Well this is obviously the wrong slide.
Here we are. Let's start with Recommendation 25C:
Responsibility of authors forming new names. “Au-
thors should exercise reasonable care and considera-
tion in forming new names to ensure that they are cho-
sen with their subsequent users in mind and that, as
far as possible [and this is the good part], they are ap-
propriate, compact, euphonius [i.e., pleasant to the
ear], memorable, and do not cause offence.” Its this
last phrase that I want you to remember for the test.
Okay, here comes the meat.
Wuat Is A PATRONYM?
A patronym is a scientific name that honors a person
by incorporating that person’s name into the name of a
genus, species, or subspecies. Here’s one of the rules
you need to follow. Article 31.1.2. “A species-group
name . . . is to be formed by adding to the stem of that
name ‘i’ if the personal name is that of a man, ‘orumy if
of men or man (men) and woman (women) together,
‘ae’ if of a woman, and ‘arum’ if of women . . .” This is
one of the easiest ways to come up with latinized
names for species, and I used it liberally when I
started describing Lepidoptera about 20 years ago. For
example, I named Habrodais poodiae for my wife
VOLUME 55, NUMBER 1 3
— | AY dame onored Metliod for,
EVERYTHING YOU | Sircss| Reduction mm
ALWAYS WANTED TO | Fost-Reproductive Lepidoptensis
KNOW ABOUT |
NOMENCLATURE.....
BUT WERE AFRAID TO ASK se
Bemic Mi St. John
Ine statistical rrovi
Parry A. Howell
Best Western
The Book-of-the-Month
Club:
Writing One, That Is
Fics. 1-6. PowerPoint slides illustrating templates for various types of talks (see text for explanation).
Poody Brown, Mitoura thornei for one of my early named for Donald and Sandra Duckworth. (Just sort
mentors (Fred T. Thorne), and Ewphyes vestris har- of on the side, if your last name was Duckworth, would
bisoni (hmm, three names, must be a butterfly) for an- you name your son Donald? Isn’t that a little like hav-
other of my early mentors—Charles Harbison. In later ing the last name of Butterworth and naming your
years I even became clever enough to use the “orum” daughter Mrs.? Or having the last name of Wonder-
form, so this species, Cuproxena duckworthorum, is land and naming your daughter Allisen?) Anyway .. .
Vp
IEEE %
=
————
; it 2ZZ Zz
Wy, We SMe LG
Fic. 7. Straw man on house of cards beating a dead horse with a
red herring.
As your run-of-the-mill taxonomist, even I make a
contribution to the study of tortricid moths from time
to time. And once in a while some of our contributions
are recognized by others in our field and they name
a species after you. And here it is, my very own
patronym—Phtheochroa johnibrowni Razowski, 1991—
solid gold! This species was named after me by Josef
Razowski—a Polish tortricid worker, wouldn't you
know it. Actually, this is a pretty goofy-looking species
name. Remember, you add an “i” to the end of a man’s
name, so with a last name like Brown, you shouldn't
expect too many patronyms, if you get my drift.
Well, if you're one of those scientists who make lots
of significant contributions, several people may name
species after you. So for example, here’s some of the
Lepidoptera species named for Jerry Powell (Table 1),
who is in our audience this evening. There are geo-
metrids, and pyralids, and tortricids and all sorts of
things. Well, if youre one of those scientists who
makes lots of significant contributions and you're also
really dead, there’s virtually no end to the number of
patronyms you may receive. Here’s (Table 2) just the
tortricid species named for Alex Diakonoff, a Dutch
microlepidopterist whose work spanned the period
from about 1940 to about 1990; he published over 250
papers on Lepidoptera, and he has a ton of things
named after him.
Actually, [ll bet there are 15-20 folks here tonight
with species named after them. I know there’s one or
more leuschneri (for Ron Leuschner), and we saw
there are lots of powelli, and there’s an epsteini and a
poguei, but I think they're names of biting flies (cerat-
apogonids) rather than Lepidoptera, and there’s a
millerorum for Lee and Jackie Miller, and a burnso-
rum for John and Sarah Burns, and probably a whole
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
TABLE 1. A few of the many Lepidoptera patronyms for Jerry
Powell.
Coptodisca powellella Opler (Heliozelidae)
Gyros powelli Munroe (Pyralidae)
Stegea powelli Munroe (Pyralidae)
Pterotaea powelli Rindge (Geometridae)
Dorithia powellana Brown (Tortricidae)
Clepsis powelli Razowski (Tortricidae)
Henricus powelli Razowski (Tortricidae)
bunch more that I don’t know about. Okay, so we’ve
got the concept of patronym nailed. So let’s move on.
But first, here’s our first quiz. This quiz is for those
young ladies in the audience 16 years or younger. Who
is this devilishly handsome young lad? [Slide of
Leonardo DiCaprio] [The voice of Astrid Caldas
shouts out from the back of the room—Leonardo Di-
Caprio]. Okay. Any idea of his Latin bnomen? How
about Homo sapiens? Good.
Wuat Is SYNONYMY?
When a species of animal has been described or
named more than once, the names are said to be syn-
onyms—that is, both (or all) names refer to the same
species. This can happen in a variety of ways. For ex-
ample, it can happen when different scientists name
the same species because they are unaware of each
other's work. But it also can happen when the same
scientist names a species more than once from differ-
ent specimens because he doesn’t recognize that they
represent the same species. And this typically happens
when species are real variable, that is, no two individ-
uals look alike, or when they exhibit strong sexual di-
morphism, that is, males and females look different.
Most of you are probably familiar with the California
dog face butterfly—the male has been called the “fly-
ing pansy” and the female is a plain yellow butterfly, so
they are remarkably distinct. Remember, a lot of us
work on dead, pinned bugs in a museum, so we sel-
dom get the chance to see interaction between the
sexes (the Lepidoptera sexes, that is).
One of our greatest authors of synonyms in Lepi-
doptera was Francis Walker. And this is obviously a
very dubious honor. Walker was paid by the British
Museum to catalogue their Lepidoptera collection,
and when he came across species that he did not rec-
ognize, to describe them. Actually, he was paid by the
species. Well, apparently Walker did not have that
great of an eye for species because he described many
of them multiple times. For example, Mike Pogue tells
me that in the noctuid genus Spodoptera, an ugly
bunch of cutworms, Walker described 48 different
species, placing them in 10 different genera. Of these
VOLUME 55, NUMBER 1
TABLE 2. The Tortricidae patronyms for Alex Diakonoff.
Bactra diakonoffi Amsel
Eucosma diakonoffi Gibeaux
Metaselena diakonoffi Horak & Sauter
Sycacantha diakonoffi Kawabe
Penthostola diakonoffi Kawabe
Statherotis diakonoffi Kuznetsov
Eboda diakonoffi Razowski
Tortricibaltia diakonoffi Skalski
Diakonoffiana Kogak
Diakonoffiana Kuznetsov
48 species, only 8 are recognized as valid today, so
Walker is responsible for creating 40 synonyms in
Spodoptera alone! Another example of Walker's keen
eye is the species Epiphyas postvittata (Walker), the
light brown apple moth, a leafrolling pest in many
parts of the world. Granted, its pretty darn variable,
and males look different from females. Walker de-
scribed this species 9 times in three different genera—
§ times in the same catalogue! All these names refer to
the same species. And since a species can have only
one unique name, only one is the correct name and
the rest are synonyms, extra names that clutter the lit-
erature and cause confusion.
Now for a slightly more twisted example of syn-
onymy, Id like to tell you a little story about Edward
Meyrick and William Kearfott. We'll start with Kear-
fott. William Kearfott was a physician who worked on
American Tortricidae around the turn of the 20th cen-
tury. And the names he proposed for new species are
among those that are, well, shall I say, less than schol-
arly. Actually, Kearfott’s names stand as a tribute to
whimsy, whether intentionally or not. When faced with
a large number of new species, most of us soon ex-
haust our imagination for names, leaning on old stan-
dard prefixes such as pro-, neo-, pseudo-, eu-, and so
forth. Not Kearfott. Kearfott approached his new
names in a very orderly fashion, apparently leaning
heavily on his very thorough knowledge of the alpha-
bet (you know, a, b, c, d. . .) and his keen ear for a
good rhyme. Here are some real Kearfott species
names (see Table 3): bobana, cocana, dodana, fofana,
gogana, hohana . ..—stop me when you see a pattern.
Well, for this set of names, Kearfott started a species
name with every letter of the alphabet, except vowels,
j, q, w, and x. So he got a lot of mileage from this one
pattern—16 names. Here are more Kearfott names
(Table 3): fandana, gandana, handana, kandana . . .;
and who could forget the concise, euphonious, and
memorable (Table 3) gomonana, tomonana, vomo-
nana, womonana, zomonana, or baracana, caracana,
daracana, faracana, haracana, maracana, naracana,
raracana, and yaracana.
TABLE 3. A few of the many tortricid species names proposed by
William Kearfott.
bobana dandana baracana gomonana dana
cocana fandana caracana tomonana __ fana
dodana gandana daracana vomonana kana
fofana handana faracana womonana lana
gogana kandana haracana zomonana mana
hohana landana maracana tana
kokana mandana naracana vana
lolana nandana yaracana wana
momana pandana zana
nonana randana
popana sandana
rorana tandana
sosana vandana
totana wandana
vovana
zozana
Because Kearfott’s (1904, 1907a, b, c) names were
published in widely distributed scientific journals and
his species were adequately described and diagnosed,
his names are as valid as anyone's. Well I like Kearfott’s
names. Actually, they remind me of that song from the
1960's, by Shirley Ellis. [The voice of Don Harvey
shouts out from the side of the room: “The Name
Game.” Yes, exactly! And If I remember correctly, the
first verse went something like: Shirley, Shirley, bope-
early, banana, fana, fope-early, me, my, moe, merly,
Shirley . . . or something like that. [Don nods in agree-
ment. |
Well in contrast to Kearfott was Edward Meyrick, a
no-nonsense, British school master that was a contem-
porary of Kearfott. Meyrick was quite the Latin scholar
and probably the most prolific describer of microlepi-
doptera ever, describing over 14,000 species (Clarke
1955), all with well formed Latin binomials. You can
just imagine his outrage and incredulity upon seeing
the Kearfott names in a published journal. He surely
must have thought that these unwashed, godless hea-
thens in the colonies have no right naming new species
if they can't do it correctly. Well, Meyrick responded to
Kearfott’s work with a paper called “On some impossi-
ble scientific names in Micro-Lepidoptera,” published
in 1912. In this paper Meyrick (1912a) described the
Kearfott names as “. . . openly and obviously based on
a barbarous and unmeaning gibberish.” I like that. It
kind of reminds me of something I’ve seen in reviews
of my papers . . . .and at least one of those anonymous
reviewers is probably in this room this evening.
Meyrick totally rejected Kearfott’s names and pro-
posed new “appropriate” Latin names to replace them.
Unfortunately, because the Kearfott names are valid,
Meyrick did nothing more than create a ton of syn-
onyms—new names for species that already have
names. For Meyrick (1912b) the concept of priority,
TABLE 4. Names on final examination.
Eubetia Brown, 1999—valid
Eubetia bigaulae Brown, 1999—valid
Eubetia raz Brown—tejected
Eubetia boop Brown, 1999—valid
Phryganidia Packard, 1864—-valid
Phryganidia steinbrenneri Miller—rejected
Polywana Brown—rejected
Polywana krakar Brown—rejected
Jerapowellia Miller, 1995—valid
Jerapowellia burnsorum Miller, 1995—valid
Dyaria Neumoegen, 1893—valid
Cephise nuspesez Burns, 1996—valid
Doa Neumoegen & Dyar, 1894—valid
that is, recognition of the oldest name as the valid
name, was nothing more than a fetish of certain taxon-
omists of the time. So instead of saving nomenclature
from the gibberish of Kearfott, Meyrick only cluttered
it with useless names of his own. Okay. So that’s the
deal with synonymy. Time for a quick quiz. ’'m going
to show you the life history of a lepidopteran; and
you'd better bask in it because they're the only pho-
tographs of Leps in the entire talk. As soon as you
know the family, the genus, or the species, shout it out.
Here's the egg; the first instar; the fifth instar; the
pupa; and here’s the adult. Oh, no....wrong adult!
Here’s the real adult. Everybody got Papilio thoas?
Okay, our next and last topic.
Wuat Is AN INAPPROPRIATE NAME?
Wuat Is TAUTONOMY?
Per the Code, inappropriate names are those that
convey false information about a species or genus; for
example, something like the name gigantea for the
smallest member of the genus. Article 18 states: “The
availability of a name is not affected by inappropriate-
ness or tautonymy.” So, here you can see that the Code
does not dismiss these names just because they are
stupid. Here's a few examples that sort of portray this
concept.
Philotes sonorensis (Felder & Felder), the Sonoran
blue butterfly. You might suspect that this butterfly is
from Sonora. Nope—California. Well maybe it occu-
pies the Sonoran Zone. Nope—it ranges from the
coast to the mountains. How about Ethmia arc-
tostaphelella (Walsingham). You might suspect that the
larva of this feeds on Arctostaphylos. Nope—Eriodic-
tyon. Simmondsia chinensis (Link) C. K. Schneid. This
is the scientific name of jojoba, the plant that provides
that fancy oil used in gucchi shampoos, which I use, of
course (I thought it would be okay to use one plant
name). From the name chinensis, you might suspect
that it is from China. Nope—its native to Chile and Cal-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
ifornia, not China. And Decodes fragariana (Busck).
Fragaria is this genus of strawberry, so maybe this thing
is a strawberry pest. Nope—its larvee feed on oaks.
So how about tautonymy. Well, that’s when the
genus and species both have the same name. Its like if
there was a man with the last name of William and he
named his son William—he’d be William William, but
I guess you could call him Bill, so that’s a little differ-
ent. Well here are a few tautonomous names: Ozotun-
cus ozotuncus, a tortricid moth described by the same
Polish tortricid worker mentioned before. Apus apus,
I haven't a clue what this is, but its always used as an
example in the Code. Rattus rattus is one of those
pesky European rats. And here’s my favorite—Bison
bison bison (three names; no its not a butterfly). It’s
not just a binomial tautonomy, it’s a trinomial because
there is a subspecies of bison in Europe. I really like
this name because I can just image the first mammal
taxonomist out there on the American prairie, creep-
ing along on his hands and knees, peeking over a ridge
and seeing this endless sea of American buffalo, and
thinking to himself, BISON! BISON! BISON!
CONCLUSION
Well, there’s just a few more sections of the Code
that we haven't talked about, but I'll bet you now know
plenty to take the test. And this is an oral examine, so
you don’t need a paper and pencil. Here’s what I’m go-
ing to do. I'm going to present a bunch of proposed
names, some of which are published and valid, and
others of which have been rejected by manuscript re-
viewers for one reason or another. And you need to tell
me which are which (See Table 4 for answers).
Here’s our first question: the genus Eubetia. The
Latin derivation is obvious, the “eu” means true or
real, the “bet”, Latin for wager or gamble, and the “ia”
just for good measure. Anybody see a problem with
this genus? Of course not, it’s a fine generic name. So
here are three potential species names in this genus:
How about Eubetia bigaulae? Yes, this is a valid name.
How about Eubetia raz? A sort of half-baked (i.e.,
one-cheek) or abbreviated patronym for the Polish tor-
tricid worker Josef Razowski. No, offensive according
to an anonymous reviewer . . . who happens to be in
this room. How about Eubetia boop? Sure; although
boop is not Latin, we can merely say that it is an arbi-
trary combination of letters; its short, euphonious, etc.,
and valid.
Now the next question: Phryganidia. I just love this
one; it reminds me of something you might hear some
taxi driver in New York shouting at you as you cross the
street in front of him— Hey, get outta da way, ya Phry-
ganidia!” Anybody got a problem with Phryganidia as a
VOLUME 55, NUMBER 1
valid name? Reasonable? Sure, its valid. So what if an
avid baseball fan in New York wants to describe a
patronym for George Steinbrenner and names it Phry-
ganidia steinbrenneri? Would that be okay? No, I'm
afraid this one was found unacceptable by a reviewer.
How about the genus Polywana? This name was
proposed for a new genus in the tortricid tribe Polyor-
thini, a group that exhibits a Gondwanan distribution.
Get it, Poly(orthini) (Gond)wana? However, the new
genus would be represented by the single species Poly-
wana krakar. Acceptable? No, both names were found
unacceptable by a co-author with no sense of humor.
How about this genus: Jerapowellia. Here the au-
thor of the new genus has used both the first and last
name of the honoree to make sure that no other Jerry
or no other Powell can think that he is the person hon-
ored by the name. Actually, the animal is a non-
descript little orange moth that nobody would want
their name associated with anyway. Is this an accept-
able genus name? Sure. How about if you add the
species name burnsorum? How cow, now there's a
frightening combination: Jerapowellia burnsorum—
two Berkeley graduates united for perpetuity in the
name of a tortricid moth. Acceptable? Yes, but in very
poor taste (depending on your taste).
How about a new genus honoring the work of Har-
rison Dyar ... . Dyaria? But what if it was intended to
be pronounced “diarrhea”? Sure. Good name.
Well say you've got a new species of skipper butter-
fly, and all the good names in the genus already are
used up. Could you name the new species “nuspesez”?
Yes, and the culprit who perpetrated this atrocity also
is in the audience this evening.
How about the genus D-O-A (Doa)? Sounds like
every moth in my collection. [The voice of Ron
Leuschner could be heard chiming in—*That’s also a
good name for a family.”] Yes, this is a valid genus and
actually the type genus for the family Doidae.
Well, I hope you've seen from this exercise that con-
cepts like concise, euphonius, memorable, and offen-
sive are really pretty subjective. And sometimes it
~l
seems as though the Code merely provides reviewers
and editors with a justification to reject names that
they don't like personally. And my interpretation is that
some rules of the code are like this (see Fig. 7)... and
this is called “Straw man on a house of cards beating a
dead horse with a red herring.” Well, there's little
doubt that our Code will continue to evolve over time,
let’s just hope it evolves faster than the species for
which it is intended to provide stable nomenclature.
Thank you.
ACKNOWLEDGMENTS
I sincerely thank my fellow lepidopterists who allowed me to
poke a little fun at them. I also thank Poody Brown for years of pa-
tience and understanding; Jerry Powell for my graduate training;
John Burns for hosting me as a post-doc; Howie Wier for rescuing
me from poverty; Martie Clemens for the dead horse; and the
USDA Systematic Entomology Laboratory for providing me with a
place to indulge in that which fascinates me the most. Linda
Lawrence arranged Figs. 1-6. Natalia Vandenburg and David Smith,
Systematic Entomology Laboratory, National Museum of Natural
History, Washington, D.C., and William Miller, University of Min-
nesota, St. Paul, provided comments and corrected errors in the text
(but they were unable to resolve the errors in logic).
LITERATURE CITED
CLARKE, J. F. G. 1955. Catalogue of the type specimens of Mi-
crolepidoptera in the British Museum (Natural History) de-
scribed by Edward Meyrick. Trustees of the British Museum.
332 pp.
EioT, T. S. 1939. Old Possum’s Book of Practical Cats. Harcourt
Brace & Co., New York.
Kearrort, W. D. 1904. North America Tortricidae. Trans. Am. En-
tomol. Soc. 30:287-299.
Kearrott , W. D. 1907a. New North American Tortricidae. Trans.
Am. Entomol. Soc. 33:1-98.
. 1907b. Microlepidoptera from the Black Mountain Region
of North Carolina, with descriptions of new species. Bull. Am.
Mus. Nat. Hist. 23:153-167.
. 1907c. New Microlepidoptera. Canad. Entomol. 34:1-9,
53-60, 77-84, 121-128, 153-160, 211-212.
LINNAEUS, C. 1758. Systema Naturae 10th ed. Holmiae.
Meyrick, E. 1912a. On some impossible specific names in Mi-
crolepidoptera. Entomol. Month. Mag. 48:32-36.
. 1912b. Correction of impossible names. Entomol. Month.
Mag, 48:32-36.
Received for publication 16 November 2000.
REE SST OS
Journal of the Lepidopterists’ Society
55(1), 2001, 8-14
INFLUENCE OF MOSQUITO CONTROL CHEMICALS ON BUTTERFLIES (NYMPHALIDAE,
LYCAENIDAE, HESPERITDAE) OF THE LOWER FLORIDA KEYS
MarkK H. SAtvaTo!
Department of Entomology and Nematology, University of Florida,
Building 970, Hull Rd., Gainesville, Florida 32611
ABSTRACT. A 14-month survey was conducted in the rock pinelands of south Florida (Long Pine Key) and the Lower Florida Keys (Big
Pine Key) to determine the status of three potentially threatened butterfly species. Populations of the Florida leafwing, Anaea troglodyta flori-
dalis F. Johnson & Comstock, and Bartram’s hairstreak, Strymon acis bartrami (Comstock & Huntington), were monitored in areas that receive
year-round chemical applications for mosquito control and in those without such treatment. Anaea troglodyta floridalis maintained significantly
higher adult densities during both years of the survey on transects where chemical applications were restricted. No significant differences were
found in A. troglodyta floridalis larval densities among transects in either year; however, the overall larval density was significantly higher in the
1998 sampling period. Strymon acis bartrami showed consistently high adult and larval densities at all Big Pine Key transects, but was not ob-
served in Long Pine Key. A third species, the rock-land grass skipper Hesperia meskei (W. H. Edwards), was not found on any of the survey tran-
sects despite a high density of its host grass, Aristida purpurascens Poir. (Poaceae). Experiments to test the potential toxicity of mosquito con-
trol chemicals on various surrogates of the above species showed naled and permethrin to be most toxic, with lethal dosage (LD,,) values of 1.0
ug or less of AI per gram of body weight for the species and stages tested. LD,, values of 48.1 ug or less AI per gram of body weight were found
for surrogates treated with malathion. Given the susceptibility of these butterflies in all their life stages to the mosquito control insecticides
presently in use, these chemicals should be considered a major factor in the populational declines and fluctuations of the butterflies studied.
Additional key words: Anaea, chemical pesticides, non-target arthropods, Strymon, Hesperia.
Human activity in south Florida and the Florida
Keys has increased dramatically in recent decades.
With year-round mild climate and scenic beauty, the
area became an appealing place to live and visit. It was
only a matter of time until this region’s unique flora
and fauna would feel the effect of human population
growth. Among the first to be negatively influenced
were the native butterflies. Although normally more
appreciated for their aesthetic appeal, butterflies are
also an extremely good indicator of an ecosystem’s sta-
bility (Erhardt 1985, Longley & Sotherton 1997).
Anaea troglodyta floridalis, the Florida leafwing F.
(Johnson & Comstock) (Nymphalidae), Strymon acis
bartrami, the Bartram’s hairstreak (Comstock & Hunt-
ington) (Lycaenidae) and the rock-land grass skipper,
Hesperia meskei (W. H. Edwards) (Hesperiidae) have
enjoyed relatively large historic ranges, occupying the
pinelands that once covered most of southern Florida
and the Lower Keys (Minno & Emmel 1993, Smith et
al. 1994). Their rapid demise in recent decades
(Baggett 1982, Schwartz 1987, Hennessey & Habeck
1991, Schwarz et al. 1995, Salvato 1999) is representa-
tive of many species in the region and can be attrib-
uted in large part to habitat loss and mismanagement.
Another possible contributor to the decline of these
butterflies is the use of chemical pesticides meant to
control mosquitoes but with collateral effects on non-
target arthropod species. The lethal effect of second-
generation organophosphate pesticides, such as naled
and fenthion, on non-target Lepidoptera was particu-
‘Mailing address: 3415A Carlton Arms Drive, Bldg. 3, Tampa,
Florida 33614.
larly well noted initially in south Florida and the Keys,
with the demise of the Schaus’ Swallowtail, Papilio
aristodemus ponceanus Schaus (Emmel & Tucker
1991, Eliazar 1992). This species’ dramatic decline in
the early 1970s coincided with the expanded use of
chemical pesticides by the Monroe County Mosquito
Control District (MCMCD) on the northern Keys.
When spraying was halted during two periods (1987
and 1989-1992), the species began to recover. Its im-
mediate decline when applications resumed clearly
suggested the adverse effect that chemical pesticides
were having on non-target species. Baggett (1982) sug-
gested that the rapid decline in A. troglodyta floridalis
and S. acis bartrami populations on the Lower Keys
was directly attributable to mosquito control insecti-
cide applications.
Studies conducted by Hennessey et al. (1991, 1992)
illustrated the presence of spray residue long after ap-
plication in the habitat of the Schaus’ Swallowtail and
several other threatened butterflies, including A.
troglodyta floridalis and S. acis bartrami. This re-
search followed a joint agreement between U.S. Fish
& Wildlife and MCMCD in 1987 on areas which were
to be designated “no-spray” zones. Dade County did
not spray insecticides for mosquito control during that
time period. Thus the only chance for chemical con-
tact on the southern mainland for these butterfly
species occurs in the residential areas east of Long
Pine Key, in Everglades National Park, where
resmethrin is sprayed occasionally. As of 1989 the fol-
lowing areas in the Florida Keys were designated no-
spray zones by agreement between U.S. Fish &
Wildlife and MCMCD: in the north, a strip of land
VOLUME 55, NUMBER 1
east of Crocodile Lake National Wildlife Refuge, El-
liott Key, and several of the smaller keys of Biscayne Na-
tional Park; and in the Lower Keys, the small outlying
areas of the National Key Deer Wildlife Refuge. All of
Big Pine Key except Watson's and eastern Cactus Ham-
mocks is sprayed with the chemical pesticides naled,
permethrin and Bacillus thuringiensis var. israeliensis.
However, no-spray does not mean a lack of chemical
intrusion. These areas were established with the un-
derstanding that there was to be no use of insecticides,
and any residues detected within them would be unac-
ceptable. When these zones were created, pesticide
drift downwind into them had not been documented.
Hennessey et al. (1992) documented naled residues
from the edge of Watson’s Hammock on o-cellulose
pads up to six hours after its application. The highest
EEC recorded was 0.009 + 0.0001 g/cm? as com-
pared to 0.011 + 0.001 Ug/cm®? in the target area.
Residues were detected 750 m into the no spray zone
at Watson’s Hammock, 150 m at Cactus Hammock and
30 m into the Schaus’ hardwood hammock habitat on
Key Largo’s Crocodile Lake. Truck-applied ultra-low-
volume (ULY) fenthion, sprayed primarily in residen-
tial areas, did not appear to drift into non-target areas.
This study indicated that naled remained in the habi-
tat well into midday, posing risk to diurnally active
non-targets such as the Florida leafwing, rock-land
grass skipper and Bartram’s hairstreak.
Eliazar (1992) conducted intensive testing on the ef-
fects of the chemical pesticides naled and fenthion on
several south Florida non-target nymphalid and papil-
ionid species. His results indicated that chemical pes-
ticides and their field application rates, particularly
those of naled, were indeed extremely toxic to non-tar-
get Lepidoptera, and were being administered in the
field at levels above the dosage required to kill target
Aedes mosquitoes. Eliazar’s naled experiments in-
cluded several butterfly species likely to be found in
the Lower Keys, including nymphalid species similar
to the Florida leafwing. Among these were the gulf
fritillary, Agraulis vanillae Michener, and the zebra
longwing, Heliconius charitonius Comstock & Brown.
The potential influence of the pyrethroids, such as
permethrin, which are currently used in the Lower
Keys has only been evaluated for two butterfly species
previously, Papilio cresphontes Cramer (Papilionidae)
and Vanessa caudui L. (Nymphalidae) (Eliazar 1991,
1992). Furthermore, toxicity tests have never been
performed to determine the effect of any chemical
pesticide upon lycaenids, such as Bartram’s hairstreak,
or on hesperiids such as the rock-land grass skipper.
Part of any recovery plan for these species must in-
clude evaluation of pesticide effects on populations. I
conducted experiments on various life stages of non-
threatened butterflies (congeneric to the three focal
species here), to obtain lethal toxicity levels for expo-
sure to naled, malathion and permethrin. Further-
more, I monitored populations of Florida leafwing,
rock-land grass skipper and Bartram’s hairstreak in the
field, and examined the possible impact of chemicals
being sprayed for control of adult mosquitoes in the
Lower Keys.
MATERIALS AND METHODS
Population survey. A 14-month survey was con-
ducted in south Florida and the Lower Florida Keys
from July 1997 to August 1998. Line transects (N = 9)
were established on Big Pine Key to monitor popula-
tions of Anaea troglodyta floridalis, Strymon acis bar-
trami and Hesperia meskei. The survey employed a
combination of several butterfly count methods (Pol-
lard 1977, Gall 1995). These were adapted to accom-
modate the sparse populations of butterflies associated
with a consistently occurring host on Big Pine Key.
Each transect was 400 m in length x 5 m in width (area
0.2 ha) (437 x 6 yards, or 0.5 acres); each had evenly
distributed amounts of Croton linearis Jacq., the sole
host plant for both A. troglodyta floridalis and S. acis
bartrami. Croton linearis (N = 100) of varying sizes
were randomly chosen within each transect and
marked with flagging tape for larval inspection. These
plants were surveyed at every visit during the course of
the study. When plants died, new ones were included.
Larvae and adults viewed at the fringes of designated
transects were noted but not included in transect
counts. Aristida purpurascens, the only known host
plant for H. meskei, was inspected for larval activity.
Transects were visited twice daily during late spring to
early fall (April-August 1998, July-September 1997),
and once daily the rest of the year (October-Decem-
ber 1997, January-March 1998) when daylight was
more limited. This procedure allowed mainland and
key transects to be visited on the same sampling dates.
One transect was established at Gate 4 of Long Pine
Key within Everglades National Park; it had the same
dimensions as described above, but it was interrupted
by a clear-cut area from historical logging times which
mimicked grass savannah at its midpoint, and this
clear-cut area was not considered part of the transect
proper. Gate 4 at LPK was chosen because its pineland
habitat is extremely similar to that of Big Pine Key.
These two areas historically have maintained the
largest populations of A. troglodyta floridalis and S.
acis bartrami as well as moderate levels of H. meskei.
The pinelands of Everglades National Park are not
sprayed with chemical pesticides.
10
Anaea troglodyta floridalis and S. acis bartrami
adults were captured by net and marked with the 1-2-
4-7 numbering system (Ehrlich & Davidson 1960).
Hesperia meskei was not observed within the transects
at any point during the survey despite high density of
its host. Plants which contained earlier stages were
marked with field tape for later inspection; however,
this technique was replaced by use of natural field
markers whenever possible.
Adult and larval densities were transformed to the
square root of (X + 0.5) and analyzed by ANOVA:
Single Factor analysis in a completely randomized
block design with all sampling dates and sites as
sources of variance. Data from 1997 and 1998 were
analyzed separately. All treatment areas were com-
pared against controls (Watson’s Hammock and Long
Pine Key, the areas where insecticide applications are
restricted). Where significant F values were found,
Tukey’s test was used to separate means.
Lethal dosage experiments. To help determine if
mosquito control chemicals have a toxic effect on the
various life stages of these potentially threatened but-
terfly species, lethal dosage levels were determined for
similar, non-threatened butterfly species. The chemi-
cal insecticides used were malathion (O, O-dimethyl
phosphorodithioate of diethyl mercaposuccinate;
molecular weight = 330.4 ), naled (dimethyl-1, 2-di-
bromo-2, 2-dichloroethyl phosphate; molecular weight
= 381) and permethrin ([3-Phenoxyphenyl] methyl-
[+]-cis-trans-3-[2, 2-dichloroethenyl]-2, 2-dimethylcy-
copropane carboxylate; molecular weight = 391.3).
Malathion and naled donated by the Division of Plant
Industry (DPI) in Gainesville and DPI in Winter
Haven, Florida, respectively. Permethrin was pur-
chased from Chemserve (West Chester, Pennsylvania).
The Monroe County Mosquito Control District
(MCMCD) applies naled, a second-generation organo-
phosphate insecticide, by aircraft (DC-3 and heli-
copter) throughout the Florida Keys. Planes/heli-
copters fly at an altitude of 50 m (165 ft) with swath
widths of 61 m (200 ft) (Hennessey et al. 1992). The
field level of naled application by MCMCD is 0.08 kg
(Al)/ha as a 4% mixture with No. 2 diesel fuel (vol/vol).
Likewise, diesel fuel is frequently used by MCMCD in
their ground based ULV fog mixtures of another
organophosphate insecticide malathion and a pyreth-
riod insecticide permethrin, so this substance was used
as the control whenever possible to best simulate ac-
tual application conditions. Acetone was used as an al-
ternate control when No. 2 diesel proved to be toxic to
a test species. No. 2 diesel fuel was obtained from var-
ious gasoline stations and acetone was purchased from
K-mart, both in Alachua County.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Test butterfly species and test procedure. The
gulf fritillary, Agraulis vanillae, and the zebra long-
wing, Heliconius charitonius, were used as the experi-
mental nymphalid butterflies to evaluate toxicity to
topically applied mosquito control insecticides. Both
species share similar habitat with A. troglodyta flori-
dalis throughout south Florida. Larvae of both species
were reared from eggs obtained at the Sanibel-Captiva
Conservation Foundation (SCCF) in Sanibel, Florida,
and from females reared at the Boender Endangered
Species Laboratory in Gainesville, Florida. Develop-
ing larvae were fed a daily diet of Passiflora spp.
leaves. Specimens were reared in plastic cups. Fifth
instars and adults of both species were tested for toxi-
city of malathion and permethrin. Larvae were tested
with treatment levels in logarithmic steps by applying
one microliter (11) of malathion or permethrin solution
to the dorsum of the thorax with a Hamilton dispenser
(PB600-1) fitted with a 25-1] syringe. Upon applica-
tion, treatment groups were returned to their respec-
tive containers, provided fresh food, and monitored for
24 hours. Adult treatments were given in the same
way. Adults were marked on the wing with colored ink
to indicate the dosage used and released into a screen-
enclosed flight cage at the Boender Endangered
Species Laboratory or SCCF. These adults were pro-
vided with live flowering plants as a nectar source.
The Atala hairstreak, Ewmaeus atala Rober, was
used as a surrogate lycaenid butterfly in tests with
mosquito control chemicals. Although E. atala is clas-
sified as a threatened species, large populations of the
butterfly exist in parts of Dade County. One such area
is Fairchild Tropical Gardens in Coral Gables. Here
the larvae are considered a major pest of ornamental
cycads, especially Zamia pumila. Early instars were
obtained with permission on several occasions and
reared to 5th instar for testing with naled and perme-
thrin with the same techniques as those for the
nymphalids. After treatment, adults were placed in
small indoor cages for the test period, with each treat-
ment in a different, labeled cage. Experimental sur-
vivors were brought to the SCCF for use in their but-
terfly enclosure exhibit.
The long-tailed skipper, Proteus ae (L.), and
the tropical checkered skipper, Pygrus oileus (L.), are
two hesperid species common throughout the
pinelands of southern Florida and the Lower Keys.
These species were tested as surrogates to determine
the potential toxicity of naled and malathion toward
Hesperia meskei. Larvae and adults of P. urbanus were
collected from areas throughout Gainesville, Florida,
where its host, beggar’s tick, Desmodium spp., occurs.
Additional P. urbanus specimens were collected in
VOLUME 55, NUMBER 1
TaBLE 1. Comparison of mean adult Anaea troglodyta floridalis
& Strymon acis bartrami densities per hectare at all transects for
1997 (29 sampling dates)—1998 (34 samples).
A. troglodyta floridalis S. acis bartrami
Transect 1997 1998 1997 1998
Nature Conservancy 0.5 1.0 6.2 6.9
Lytton’s Way 0.0 0.2 2.6 2.6
Watson's Blvd. 0.0 0.2 4.8 2.3
Blue Hole 0.0 0.2 3.6 9.4
Watson’s Hammock QD 5.0 17 2.5
Key Deer Refuge North 0.3 0.6 5.0 46
Key Deer Refuge South 0.3 0.9 8.6 5.0
Coconut Palm 0.0 1.9 BD 3.7
Ixora Drive 0.5 1.5 17 8.4
Long Pine Key, Gate 4 2.4 2.9 0.0 0.0
Alva, Florida at sites along State Route 78. Pygrus
oileus larvae and adults were obtained from hollyhock
in July and August 1998 from northeastern Newberry,
Florida. The larvae of both species roll a silken tent on
the leaves of their respective hosts, making them easily
located. Larvae were tested by the above procedures.
As with the lycaenids, adult hesperids were placed in -
individual cages after treatments.
Lethal dosage analysis. Determination of lethal
dosage values for various surrogate species and stages
of south Florida Lepidoptera were derived using an in-
novative experimental design created by Peter J. Eli-
azar (Department of Entomology and Nematology,
University of Florida) (Eliazar 1992, Eliazar in press).
Prior to each test, replicate larvae/adults were weighed
to obtain an average instar/imago weight for each
species before treatment. Mortality data were pooled
for each species and stage tested, to provide a larger
sample for analysis. Experimental LD,, values were
then determined with a probit analysis program run
through an Apple Macintosh Microsoft Excel Spread-
sheet. This probit analysis program, created by Dr.
James L. Nation (Department of Entomology and Ne-
matology, University of Florida), was derived from the
equations and discussion found in Finney (1964).
Table values were taken from Busvine (1971). LD,,
values of each species were divided by the average in-
star or adult weight of that species to derive an “LD.,
per gram of body weight” value. For direct comparison
of lethal dosages between larvae and adults, and for
comparison of LD,, values for other lepidopteran
species, the “percent volume-to-volume concentration
per gram of body weight” value was converted into
“micrograms of active ingredient per gram of body
weight” (g/g). The levels of active ingredient (AI) are
1510, 1186 and 26.9 micrograms per microliter of con-
centrate for naled, malathion and permethrin, respec-
tively. See Eliazar (1992) and Salvato (1999) for a more
1]
detailed description of the testing protocol used in
these experiments.
RESULTS
Adult survey results. A total of 131 adult Anaea
troglodyta floridalis was marked and released during the
survey period. Means per hectare at each of the ten tran-
sects for the 1997 portion ranged from 0.0 at four Big
Pine locations (Lytton’s Way, Watson’s Blvd., residential
sites, Blue Hole and Coconut Palm within the refuge)
to 2.4 within the Everglades. The Watson’s Hammock
site maintained the highest density on Big Pine with 2.2
per hectare (Table 1). There was no significant differ-
ence (F-test, p = 0.05) in A. troglodyta floridalis density
between the control sites. There were significant differ-
ences, however, between both these control sites and
four treatment areas for 1997 (Tukey's test, p = 0.05),
with lower butterfly density in these sprayed locations.
A total of 97 adult A. troglodyta floridalis was
recorded on dates between 18 January and 29 August
1998 (Table 1). Means during this period ranged from
0.2 individuals per hectare at Lytton’s Way to 5.0 at
Watson’s Hammock (Table 1). There was no significant
difference between control transects (F-test, p = 0.05);
however, there were differences between both con-
trols and three of the treatment sites in 1998 (Tukey's
test, p = 0.05).
A total of 407 Strymon acis bartrami adults was
marked and released, 232 during the 1997 and 175 in
the 1998 sampling periods. During the sampling dates
of July and August 1997, far more individuals were ob-
served than could be marked by a single surveyor; this
combined with the few recaptures illustrates the large
density of S. acis bartrami present. The 1997 sampling
period revealed per-hectare population means ranging
from 0.0 (at Long Pine Key) to 8.6 on the southern
Key Deer Refuge transect (Table 1). This low S. acis
bartrami density at Long Pine was significantly differ-
ent from the Watson’s Hammock transect (Tukey’s-
test, p = 0.05). Two other sites (Nature Conservancy
and southern Key Deer Refuge, both are insecticide
treated areas) were also shown to have a significant dif-
ference with both control transects, in these cases,
however, butterfly density was much higher on the in-
secticide treated spots than controls.
Again in 1998, the low population for S. acis bar-
trami was at Long Pine Key where no S. acis bartrami
were recorded, thus being significantly different from
Big Pine Key control site at Watson’s Hammock (Table
1). Ixora Drive had the highest density on Big Pine at
8.4; this site, as well as the Nature Conservancy site,
were both significantly different in hairstreak density
(higher) than either control.
TABLE 2, Comparison of mean larval Anaea troglodyta floridalis
and Strymon acis bartrami densities per hectare at all transects for
1997 (29 sampling dates)—1998 (34 samples).
A. troglodyta floridalis S. acis bartrami
Transect 1997 1998 1997 1998
Nature Conservancy 0.5 1.6 0.0 0.2
Lytton’s Way 0.3 15) rll 0.6
Watson's Blvd 0.0 0.3 0.7 0.2
Blue Hole 0.2 4.9 1.0 0.9
Watson’s Hammock 0.0 Bd 0.5 29)
Key Deer Refuge North 0.2 18 0.0 0.2
Key Deer Refuge South 0.7 1.3 0.5 0.3
Coconut Palm 0.7 4.1 LQ 1.0
Ixora Drive 0.2 3.1 0.4 1.2
Long Pine Key, Gate 4 0.0 1.9 0.0 0.0
Adult Hesperia meskei were observed during the
course of this study, but not within the transects. These
sightings, first reported by Dr. Thomas C. Emmel on
Big Pine, represent the first documented observations
of the rock-land grass skipper in more than twenty
years. A female specimen was taken by Dr. Jaret C.
Daniels and the author on Big Pine on 2 June 1998.
Larval survey results. Anaea troglodyta floridalis
larvae became increasingly more common at all tran-
sects as the survey progressed. A total of 168 larvae of
various stages were marked in the field, 145 of these
on Big Pine Key. Sixteen were marked in 1997
(July-December), all at Big Pine locations (Table 2).
Per hectare mean estimates for this sampling period
ranged from 0.0 at three locations (Watson’s Blvd.,
Watson’s Hammock and Long Pine) to 0.7 at both the
southern Key Deer Refuge and Coconut Palm tran-
sects (Table 2). The remaining 152 larvae were marked
in 1998 (January—August). Densities ranged from
0.3/ha on the residential Watson’s Blvd. to 4.9/ha at
Blue Hole (Table 2). The highest larval density oc-
curred on 18 May 1998 at Blue Hole, where 18 larvae
were marked. As with the adults, larvae of the Bar-
tram’s hairstreak were plentiful on Big Pine during the
summer of 1997, with a total of 37 larvae marked from
July to December (Table 2). Lytton’s Way maintained
the highest per hectare density at 2.1, while Nature
Conservancy, north Key Deer Refuge and Long Pine
had none. However, in keeping with the decrease in
adult density to mid-1998, the number of larvae also
declined slightly to 31 for the 1998 portion of the cen-
sus. Four locations supported no larval activity (Wat-
son’s Blvd., north-south Key Deer Refuge and Long
Pine). The un-sprayed Watson’s Hammock had the
highest density for 1998 with 2.2/ha. No S. acis bar-
trami larvae were found at Long Pine Key.
There were no significant differences found in larval
density between transects for S. acis bartrami larvae in
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
either survey year. However, there were significant dif-
ferences in 1998 for larvae of the Florida leafwing on
four transects, with these higher densities on areas that
are normally treated with chemicals. Extremely dry
conditions dominated the final two thirds of 1998, this
alleviated the need for insecticide applications.
LD,, results. Following the experimental design of
Eliazar (1992), LD,, values obtained using probit
analysis (Salvato 1999) were divided by the average
weight for the instar/imago of each species to give a
percent volume-to-volume concentration per gram of
body weight value. These values were then multiplied
by micrograms of active ingredient (AI) per microliter
of liquid concentrate to give an LD,, value expressed
as ig/AI per gram of body weight (Table 4). These
treatments showed naled and permethrin to be the
most toxic, with LD,, values of 1.0 g or less of AI per
gram of body weight for the surrogate species tested.
LD,, values of 48.1 g or less of AI per gram of body
weight were found for surrogates treated with
malathion.
DISCUSSION
Adverse effects of pesticides on non-target organ-
isms depend not only on the concentration of chemi-
cals applied, but also on the pesticides’ persistence and
availability to susceptible life stages of the organisms
(Pierce et al. 1989). During these surveys, transects
were established on Big Pine to assess not only poten-
tial differences in butterfly density between treated
and untreated areas, but also to measure and correlate
the potential differences between the impact of drift
and contact from aerial and/or ground ultra-low-vol-
ume applications. If indeed upwards of 70-80% of
aerially delivered insecticides are lost in the canopy,
then the nature of that canopy, the needles of slash
pine or the herbaceous layer itself; as a landing site for
the spray droplets needed to be considered (Fairchild
et al. 1987).
Although these butterfly species are multivoltine,
their numbers experience their largest increase at the
same time as the seasonal increase in the insecticide
applications. The largest adult densities for A.
troglodyta floridalis were in Watson’s Hammock, the
no-spray area, both years. Similarly, 2.7/ha was the
mean density for A. troglodyta floridalis in the Ever-
glades. Chemical insecticide applications, which had
formerly involved resmethrin by airplane, have not
been performed at this location in 30 years. Thus these
sites were considered control areas for comparison
with those on Big Pine that are currently treated.
These remaining insecticide treated transects (N = 8)
were further divided by the insecticide applications
VOLUME 55, NUMBER 1
TABLE 3. Type of mosquito control application with influence to
each transect and mean adult densities (per hectare) for A. troglodyta
floridalis, S. acis bartrami and H. meskei over 63 sampling dates dur-
ing 1997-98 on Big Pine Key and Everglades National Park.
A.
Aerial Ground troglodyta S. acis H.
Transect spray spray floridalis bartrami meskei
Nature Conservancy x — 0.8 6.6 0.0
Lytton’s Way xX x 0.1 2.6 0.0
Watson's Blvd. = xX 0.1 3.5 0.0
Blue Hole x x 0.1 2.9 0.0
Watson’s Hammock = = OH 2.1 0.0
NKDWR (N) — Xx 0.5 48 0.0
NKDWR (S) = Xx 0.6 6.7 0.0
Coconut-Palm x xX 1.0 4.4 0.0
Ixora Drive — x 1.0 5.3 0.0
Gate 4 at LPK — — D7 0.0 0.0
_ that might impinge upon them (aerial swath of ultra-
low-volume or thermal naled fog, ultra-low-volume
spray of truck applied thermal permethrin fog, or
both). A comparison of all three treatment scenarios
revealed no significant difference in adult butterfly
mean densities over the survey period as a result of ap-
plication type (Table 3). In all cases, A. troglodyta
floridalis density failed to exceed 1.9/ha (Coconut
Palm transect 1998) during this study, on chemically
treated transects. Anaea troglodyta floridalis would
appear at greater risk overall from both types of appli-
cation, due to the fact that it flies at all levels of the
canopy, including perching high atop the slash pines.
All surrogate nymphalid species tested proved very
sensitive to chemical insecticides applied for mosquito
control. Heliconius charitonius was found to possess a
high sensitivity towards permethrin with LD,, values
of 0.002 and 0.0004 ug of AI per gram of body weight
for the fifth-instar and adult, respectively (Table 4).
This species appeared less sensitive to malathion. It
must be noted that all values provided for H. charito-
nius indicate the species toxicity solely towards the in-
secticide. These studies, as with those of Eliazar
(1992) found a 100% mortality rate for H. charitonius
towards the No. 2 diesel fuel. Because of this toxicity
to No. 2 diesel fuel, acetone was used as the control
substance for treatments using this species. However,
No. 2 diesel fuel appeared to have a less dramatic ef-
fect on the other nymphalid tested, Agraulis vanillae.
When tested with malathion, A. vanillae indicated
LD,, values of 6.7 and 8.5 ug of AI per gram of body
weight for fifth instar and adult, respectively (Table 4).
These malathion values, while not as low as those
recorded for this species towards naled (from Eliazar
1992) and permethrin, can still be classified as ex-
tremely toxic.
Immature and adult Atala hairstreaks were found to
be equally susceptible to truck applied ULV permeth-
13
TABLE 4. LD. values (in micrograms of active ingredient per
gram of body weight) for all surrogate test species.
Naled Naled Perme-
Species/Stage (acetone) (diesel) — Malathion thrin
Proteus urbanus
3rd instar 0.0699 — 0.2603 =
4th instar 0.0439 = 8.912 —
5th instar 0.0296 0.0889 0.3045 =
adult 0.1892 0.3632 13.458 —
Pygrus oileus
4th instar 1.021 = = =
5th instar 0.304 = _ =
adult 0.0823 = = =
Eumaeus atala
5th instar = 0.0009 = 0.0009
adult = 0.0012 a 0.0036
Agraulis vanillae
5th instar = = 6.572 =
adult = — 8.515 =
Heliconius charitonius
5th instar = = 8.127 0.0015
adult — — 48.087 0.0004
rin as well as the aerially applied naled. Given the ex-
treme sensitivity of this lycaenid to both pyrethroid
and organophosphate insecticides, it is likely that
threatened species, such as Strymon acis bartrami, are
at grave risk to these applications. Due to the fact that
this species commonly aggregates on the low-lying
shrubs along disturbed roadsides, areas frequently tar-
geted by ULV truck spraying, this butterfly would ap-
pear most at risk to insecticide applications of this
type. If this is truly the case, then the potential S. acis
bartrami densities indicated in this survey, especially
during the summer 1997 sampling dates, are only a
fraction of what might have been attained.
The lower density of S. acis bartrami within the
non-chemically treated areas appears to be in response
to an inconsistent burn agenda and a negative effect
on host density due to this fire restriction. Bartram’s
hairstreak larvae are dependent upon fresh host growth
for development, Croton linearis is the single host both
for this butterfly and A. troglodyta floridalis. The adult
hairstreaks appear to require consistently occurring
host for larger densities and successful dispersal (Sal-
vato 1999, in press). Neither of these requirements
seems to be met within the areas of Watson’s Ham-
mock or Long Pine Key used for this study.
Hesperiid species maintain populations in virtually
every ecosystem type found throughout the keys. Both
hesperiid species tested showed the greatest suscepti-
bility to naled, followed by the ULV malathion. Large
densities of Aristida purpurascens are also located
where they are directly threatened by ULV truck ap-
plications, and other large populations of the plant ex-
ist across the open grass savannah areas of Big Pine
within the Refuge. These large clumps of A. purpuras-
a ne
ee
14
cens have allowed H. meskei to be perhaps the most
exposed butterfly of the three rock-pineland occurring
species discussed here to the full array of insecticides
used for mosquito control. Proteus urbanus was found
to be extremely sensitive to naled, in both juvenile and
adult stages. The area in which Hesperia meskei were
located during this survey was one not frequented by
ULV trucks, but one that was still likely exposed to the
aerial applications. The drier conditions on Big Pine
during the last eight months of 1998 resulted in fewer
mosquito control insecticide applications. The new ob-
servations of adult H. meskei and the enormous in-
creases in A. troglodyta floridalis larval and adult ac-
tivity in normally treated areas could likely be the
result of this reduction in chemical spraying. Contrar-
ily, the dry conditions had a negative impact on Croton
linearis blooms and density, which directly affected S.
acis bartrami, over a time frame apparently so favor-
able to the other two butterflies (Salvato 1999, in
press).
The toxicity of permethrin to E. atala and H. chari-
tonius indicated in this study may be underestimates.
Permethrin was prepared in an non-synergized form.
Further studies with this insecticide and these butter-
flies will need to include piperonyl butoxide (PBO),
the synergist added to most pyrethroid insecticides to
increase the potency and residual effects and thus
likely their toxicity to non-target organisms.
According to Matsumura (1990), an insecticide that
produces an LD,, value that is less than 1 ug/g of body
weight is commonly classified as extremely toxic; 1-50
ug/g is highly toxic; 50-500 ug/g is considered moder-
ately toxic and a value between 500-5000 ug/g is only
slightly toxic. Beyond this last level, the chemical is
considered practically nontoxic to relatively harmless.
In regard to the Florida leafwing, Bartram’s hairstreak,
and rock-land grass skipper, the chemicals currently
being applied for mosquito control in the Lower
Florida Keys can be considered extremely to highly
toxic, depending on the surrogate species and stage
tested. Given the results of the Hennessey et al. (1991,
1992), insecticide drift experiments in Watson’s Ham-
mock, it is likely that chemical applications play an im-
portant role in affecting the population size and be-
havior of these species.
ACKNOWLEDGMENTS
The author thanks T. C. Emmel and J. L. Nation (Department of
Entomology and Nematology, University of Florida) for advice and
review of this manuscript, P. J. Eliazar for his many hours of instruc-
tion and assistance in determining the lethal dosage values pre-
sented, D. Serage (Sanibel-Captiva Conservation Foundation) for
rearing and testing facilities, Fairchild Tropical Gardens for E. atala
larvae. The author thanks J. C. Daniels, J. H. Frank, D. M. Griffin
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Ill, K. A. Schwarz, R. A. Worth, M. K. Hennessey and D. H.
Habeck for their encouragement, suggestions and insights on this
research, and the natural history of these organisms. The author
would also like to thank all at National Key Deer Wildlife Refuge,
Nature Conservancy and Everglades National Park for permitting
and various technical support.
LITERATURE CITED
BAGGETT, H. D. 1982. Order Lepidoptera. In R. Franz (ed.), Inver-
tebrates. In P. C. Pritchard (ed.), Rare and Endangered Biota of
Florida. 6:78-81. Gainesville: Univ. Pr. Florida. 131 pp.
BusVINE, J. R. 1971. A Critical Review of the Techniques for test-
ing Insecticides, 2nd ed. Pp. 271-276.
EHRLICH, P. R. & S. E, Davipson. 1960. Techniques for capture-
recapture studies of Lepidoptera populations. J. Lepid. Soc.
14:227-229.
ELIAZAR, P. J. 1992. Effect of two mosquito adulticides, naled and
fenthion, on selected non-target lepidopteran species. M.S.
Thesis. Univ. Florida, Gainesville.
EMMEL, T. C. & J. C. TUCKER (EDS.). 1991. Mosquito control pes-
ticides: ecological impacts and management alternatives.
Gainesville, Florida: Mariposa Press, Scientific Publishers.
105 pp.
ERHARDT, A. 1985. Diurnal Lepidoptera: sensitive indicators of
cultivated and abandoned grassland. J. Appl. Eco. 22:849-861.
FAIRCHILD, W. L., D. C. Eipt & C. A. A. WEAVER. 1987. Effects of
fenitrothion insecticide on inhabitants of leaves of the pitcher
plant, Sarracenia purpurea L. Can. Ent. 119:647-652.
FINNEY, D. J. 1964. Probit Analysis. 2nd ed. Cambridge: Cam-
bridge University Press. 318 pp.
GALL, F. 1985. Measuring the size of Lepidopteran populations. J.
Res. Lepid. 24:97-116.
HENNESSEY, M. K. & D. H. HABECK. 1991. Effects of mosquito
adulticides on populations of non-target terrestrial arthropods
in the Florida Keys. Gainesville: U.S. Fish and Wildlife Service
and the Univ. of Florida Cooperative Wildlife Research Unit
(unpublished final report).
HENNESSEY, M. K., H. N. Nicc & D. H. HABEcK. 1992. Mosquito
(Diptera: Culicidae) adulticide drift in wildlife refuges of the
Florida Keys. Environ. Entomol. 21:714-721.
LONGLEY, M. & N. W. SOTHERTON. 1997. Factors determining the
effects of pesticides upon butterflies inhabiting arable farmland.
Agric. Ecosystems Environ. 61:1—-12.
Matsumura, F. 1985. Toxicology of Insecticides. 2nd ed. New
York: Plenum Press. 598 pp.
MINNO, M. C. & T. C. EMMEL. 1993. Butterflies of the Florida
Keys. Gainesville: Scientific Publ. 168 pp.
PIERCE, H. R., R. C. BRowN, K. R. HARDMAN, M. S. HENRY, C. L. P.
PALMER, T. W. MILLER & G. WICHTERMAN. 1989. Fate of
temophos applied to an intertidal mangrove community. J.
Amer. Mosq. Cont. Assoc. 5:569-578.
POLLARD, E. 1977. A method for assessing changes in the abun-
dance of butterflies. Biol. Conserv. 12:115-124.
SatvaTo, M. H. 1999. Factors influencing the declining populations
of three threatened butterflies in south Florida and the Florida
Keys. M.S. Thesis, Univ. Florida.
ScHWaRrZ, A. 1987. The butterflies of the Lower Florida Keys. Mil-
waukee Public Museum Contribution in Biology and Geology,
No. 73. 34 pp.
SCHWARZ, K. A., R. A. WorTH & T. C. EMMEL. 1995. Conservation
of two threatened south Florida butterflies and their host plant
(Lepidoptera: Lycaenidae, Nymphalidae). Hol. Lepid. 3:59-61.
Situ, D.S., L. D. MILLER & J. Y. MILER. 1994. The Butterflies of
the West Indies and South Florida. Oxford: Oxford Univ. Press.
264 pp. 32 pl.
Received for publication 8 November 1999; revised and accepted 19
July 2001.
Journal of the Lepidopterists’ Society
55(1), 2001, 15-43
BIODIVERSITY OF PYRRHOPYGINE SKIPPER BUTTERFLIES (HESPERIIDAE)
IN THE AREA DE CONSERVACION GUANACASTE, COSTA RICA
JOHN M. BuRNs
Department of Systematic Biology, Entomology Section, National Museum of Natural History, Smithsonian Institution,
Washington, DC 20560-0127, USA
burns.john@nmnh.si.edu
AND
DANIEL H. JANZEN
Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA
djanzen@sas.upenn.edu
ABSTRACT. Twenty-two years of rearing 2192 pyrrhopygine caterpillars (and collecting far fewer adults) show that the Area de Conser-
vacion Guanacaste (ACG) supports at least 15 species, or some 60% of the Costa Rican pyrrhopygine fauna: 3 in (lowest elevation) dry forest-—
Elbella scylla (Ménétriés) [of which E. dulcinea (Plétz) is a synonym], Mysoria ambigua (Mabille & Boullet), and Myscelus amystis hages God-
man & Salvin; 4 in (highest elevation) cloud forest—Pyrrhopyge creon H. Druce, P. aesculapus Staudinger, P. cosyra H. Druce, and Passova
gellias (Godman & Salvin); and 10 in (middle elevation) rainforest—Pyrrhopyge zenodorus Godman & Salvin, P. crida (Hewitson), P. cosyra, P.
erythrosticta (Godman & Salvin) [lectotype designated], Parelbella macleannani (Godman & Salvin), Jemadia pseudognetus (Mabille) [rein-
stated status], another Jemadia belonging to a J. hewitsonii species complex, Myscelus belti Godman & Salvin, M. perissodora Dyar [rein-
stated status], and Passova gellias. No species spans all three ecosystems, and none occurs in both dry forest and rainforest. The caterpillars
are mostly 0.1-3 m above the ground, even when individuals of their foodplants rise 20-40 m. Except for Myscelus and Passova, the caterpillars
are showy because they are ringed, barred, or spotted (with a row of large round dots) in contrasting colors. Pyrrhopygine caterpillars and pu-
pae differ sharply from those of all other hesperiids in being long-haired.
ACG pyrrhopygines are strongly host specific, like other dicot-eating skippers (pyrgines) and certain butterfly and moth families in the ACG.
(However, pyrrhopygines are more ecosystem-specialized than pyrgines.) Although ACG pyrrhopygine foodplants are dotted across the botan-
ical taxonomic landscape (in the families Lauraceae, Clusiaceae, Flacourtiaceae, Cunoniaceae, Myrtaceae, Combretaceae, Meliaceae, Malpighi-
aceae, and Araliaceae), some patterns emerge. The three species of Myscelus plus Passova gellias, which are morphologically similar, all focus
on Meliaceae (mainly Guarea glabra Vahl). Two-thirds of the reared species currently in the large genus Pyrrhopyge are monophagous to
oligophagous on members of the Clusiaceae. At the same time, ACG pyrrhopygine caterpillars are specialists at levels far below that of the plant
family: for each species of caterpillar, there are many ACG species of plants within the plant family fed upon that are not used. Moreover, within
an oligophagous species, percentages of records among its few foodplants do not reflect the relative abundances of those plants. Host specificity
seems to be geographically conservative: relatives of half of the ACG pyrrhopygine species were reared by Moss (1949) in Para, Brazil, on plants
that are usually in the same genera, and always in the same families, as those of their Costa Rican counterparts. The only pyrrhopygine to enter
the USA, Pyrrhopyge arizonae Godman & Salvin [reinstated status], has an unusually drab adult but a gaudily ringed caterpillar, whose food-
plant (Quercus) is in yet another unrelated family (Fagaceae).
Though attacked at times by ichneumonids, braconids, and tachinids, ACG pyrrhopygines are remarkably free of parasitoids. Despite slow lar-
val growth rates (among the longest within the ACG hesperiids), pyrrhopygines are at the low end of the spread of larval parasitization frequen-
cies—not just among skippers but among ACG macrolepidoptera generally: only 7.1% of all wild-caught pyrrhopygine caterpillars were para-
sitized. Most notably, not one of 295 caterpillars in the closely related genera Myscelus and Passova (the Meliaceae eaters) has yielded parasitoids.
Additional key words: caterpillars, foodplants, pupae, parasitoids, ecology, taxonomy.
Not once upon a time, but every day, a protected Hallwachs 2001). Here we treat the distinctive skipper
tropical wildland such as the Area de Conservacién subfamily Pyrrhopyginae, after rearing 2192 pyrrho-
Guanacaste (ACG) in northwestern Costa Rica (Fig. 1) pygine caterpillars of 15 species found over a 22-year
becomes more of an ecological island in an agrarian period in ACG dry forest, cloud forest, and rainforest.
sea. Its biodiversity, derived mostly from what was Though the inventory will not have turned up every
there before the advent of European agriculture, keeps last pyrrhopygine species in the ACG, we think it has
dwindling to a new equilibrium density of species. almost all of them by now.
Some additional species may immigrate to the ACG Pyrrhopygines, whatever their taxonomic rank, are a
or—given enough time—even evolve in situ, owing to monophyletic group (Ackery et al. 1999) of basically
the increasing insularization of the site. An inventory neotropical Hesperiidae ranging from the southwest-
of its present biodiversity offers a baseline against ern USA (southeastern Arizona and adjacent south-
which these changes can be measured. western New Mexico, plus the Big Bend area of Texas)
Skipper butterflies (Hesperiidae)—along with the to northern Argentina (especially Misiones, but also
moth families Saturniidae, Sphingidae, and Notodon- Salta, Formosa, Chaco, Tucuman, La Rioja, Corri-
tidae—have had special attention in the early years of entes, Entre Rios, and Buenos Aires [Hayward 1948]).
the macrocaterpillar inventory of the ACG (Janzen & While only one species extends northward to the USA,
16 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
TABLE 1. Foodplants and parasitization frequency of wild-caught pyrrhopygine caterpillars in the Area de Conservacién Guanacaste (data
source: Janzen & Hallwachs 2001).
Caterpillar species Rearing % caterpillars
Plant family records (ti Bee A Oe attacked acne a aaa ene Total %
Plant species n % Ichneumonidae Braconidae Tachinidae parasitized
Pyrrhopyge zenodorus 541 0 Wil 3.0 6.7
Clusiaceae
Vismia baccifera 99
Vismia ferruginea 1
Pyrrhopyge crida 14 0 0 0 0
Clusiaceae
Vismia billbergiana 100
Pyrrhopyge creon (1 found as pupa,
2 found as larvae not reared out but
presumed to belong to this species) 3 0 (0) 0 0
Araliaceae
Dendropanax gonatopodus 50
Dendropanax querceti 50
Pyrrhopyge aesculapus 4 0 0 0 0
Cunoniaceae
Weinmannia wercklei 100
Pyrrhopyge cosyra 353 WAIL 0.2 5.4 Wan
Clusiaceae
Chrysochlamys glauca 8
Chrysochlamys psychotriifolia
Clusia cylindrica
Clusia minor
Clusia rosea
Clusia 13877
Clusia 14104
Pyrrhopyge erythrosticta 3 0 0 0 0
Clusiaceae
Marila laxiflora 100
Elbella scylla 137 0) 8.0 0 8.0
Malpighiaceae
Banisteriopsis muricata 7
Byrsonima crassifolia 66
Heteropterys laurifolia 2
Hiraea reclinata 13
Combretaceae
Combretum farinosum 12
Terminalia catappa 1
Parelbella macleannani 59 0 0 2.0 2.0
Myrtaceae
Eugenia basilaris 29
Eugenia aff. basilaris 1
Eugenia 13782 1
Eugenia 14017 69
Jemadia pseudognetus 31 0 0 0 0
Lauraceae
Nectandra hihua 45
Nectandra aff. latifolia 16
Nectandra membranacea 16
Ocotea cernua 1
Ocotea insularis 3
Persea povedae 13
Jemadia sp. X 16 0 0 0 0
Flacourtiaceae
Casearia arborea 100
Mysoria ambigua 736 0 0.3 5.4 5.7
_ Flacourtiaceae
Casearia arguta 4
Casearia corymbosa 67
Casearia sylvestris ll
Zuelania guidonia 18
eS ee ee eee SS eee
Ter ONE
VOLUME 55, NUMBER 1
TABLE 1. (continued).
Rearing
Caterpillar species
records
Plant family
Plant species n %
Myscelus amystis hages 54
Meliaceae
Trichilia americana 91
Trichilia glabra 1
Trichilia trifolia 8
Myscelus belti 106
Meliaceae
Guarea bullata 2
Guarea glabra 37
Guarea rhopalocarpa 14
Guarea 14097 47
Myscelus perissodora 1
Meliaceae
Guarea glabra 100
Passova gellias 134
Meliaceae
Guarea glabra 8
Guarea rhopalocarpa
Guarea 13856
Guarea 13860
Guarea 14097
Guarea 14137
PQ DpH Oo
many extend southward into Argentina. Pyrrhopygines
occur from sea level to upper elevation forests. Owing
to their diurnal flight, flower visitation, large size, and
showy patterns, plus the puddling and hilltopping be-
haviors of males, adult pyrrhopygines have been rather
extensively collected; but their early stages are poorly
known. Adults of these robust skippers are distinctive
in having most to all of the antennal club reflexed.
Both their dicot-eating caterpillars and the resulting
pupae are the hippies of the skipper world; they are
well endowed with long hairs—not just on the head
but over the body as well (Figs. 2-28). We hope that
this exposure of the pyrrhopygine caterpillars of the
AGG, and this demonstration of their foodplant speci-
ficity, will stimulate further rearing in other areas.
MATERIALS AND METHODS
The inventory site. The macrocaterpillar inventory
of the ACG (Janzen 1988a, 1993, in press; Janzen &
Hallwachs 2001) began in the dry forest in 1978 and has
gradually intensified and spread to adjacent rainforest
and cloud forest (and all intergrades) (Fig. 1). ACG dry
forest (about 60,000 ha) ranges from sea level to about
600 m across the Pacific coastal plain on volcanic, marine
alluvial, and serpentine soils, and receives about 1.5 m of
rain during the May—December 6-month rainy season,
but essentially none during the dry season. It is a fine-
scale mosaic of different successional stages (5400 years
of age) and scattered bits of old-growth forest (Janzen
7
% caterpillars
attacked by Total %
Ichneumonidae Braconidae Tachinidae parasitized
0 0 0) 0
0) 0 0 0
0) 0) 0) 0)
0) 0 0 0)
1988b, http:/Avww.acguanacaste.ac.cr). The cloud forest
(about 10,000 ha) ranges from about 800 to 2000 m on
three recent volcanos (Oros{f, Cacao, Rincén de la Vieja)
and receives 3-5 m of rain and frequent ground-level
clouds, with distinct seasonality. It is mostly old-growth
forest with some abandoned farms, ranches, and logging
sites on both the Pacific and Caribbean sides of the vol-
canos. The rainforest (about 40,000 ha) ranges from
about 400 to 800 m on the eastem and northern
foothills of the three volcanos, even extending in a nar-
row band at about 800 to 1000 m around the western
sides of the volcanos, and receives 3-4 m of rain, with a
1- to 3-month semi-dry season (February—May). It is a
large-scale mosaic of old-growth forest, lightly logged
forest, secondary succession, and abandoned farms and
ranches (see http://janzen.sas.upenn.edu/caterpillars/
RR/rincon_rainforest.htm for photographs of the cloud
forest and rainforest).
Collection and rearing process. We report
records through 1999 (along with a few critical recent
records) from this ongoing inventory. Many methods
of haphazard and patterned search by highly experi-
enced resident paraecologists and parataxonomists
have been used to locate caterpillars in all ACG habi-
tats and ecosystems. The 2192 pyrrhopygine rearing
records summarized here are part of some 120,000
wild-caught caterpillar rearings. Except in a few recent
cases, pyrrhopygines were not explicitly sought but
were found through general search for all macrocater-
pillars on all species of plants. Wild-caught caterpillars
18
were brought to rearing barns and reared individually,
in foliage-filled plastic bags, at ambient temperatures
(see photographs in Janzen & Hallwachs 2001).
Pyrrhopygines were segregated for this analysis after
completion of the rearing process. Each caterpillar
rearing was recorded, and each species of caterpillar
was usually photographed when first encountered (as
well as later). These individual rearing records and
some of these photographs are in the web site data-
base (Janzen & Hallwachs 2001).
Parasitoid frequency (Table 1) comes from rearing
wild-caught pyrrhopygine caterpillars of any instar in
captivity. It is an underestimate of what occurs in na-
ture because, once caterpillars are captured, they are
protected from parasitoids. Parasitoids that oviposit in
eggs or pupae are not considered here.
Adult pyrrhopygines were identified by Burns, who
studied and compared genitalia of both sexes. This in-
volved not only our reared individuals but also other
museum material of the same species as well as simi-
lar, related taxa. Arriving at the best names for reared
Costa Rican species sometimes required taxonomic
minirevision. Caterpillars that failed to produce adults
were identified by Janzen and an experienced team of
13 resident paraecologists and parataxonomists. In
each case the identifier is specified in the database
(Janzen & Hallwachs 2001). Species of ACG pyrrhopy-
gines reared to date can be identified from their larvae
and pupae (Figs. 2-25).
Series of conspecific adults reared by the macro-
caterpillar inventory are pinned and spread, and many
of the Hesperiidae are deposited as voucher speci-
mens in the National Museum of Natural History
(USNM), Smithsonian Institution, Washington, DC,
USA. Representative series are also being deposited in
INBio (Instituto Nacional de Biodiversidad), Santo
Domingo de Heredia, Costa Rica, and elsewhere.
Adult hesperiids were rather thoroughly but hap-
hazardly collected throughout the ACG by Janzen and
W. Hallwachs from 1978 to the present and, in a total
of about 20 person-years of effort, by INBio and ACG
parataxonomists from 1989 to the present. Further-
more, adult skippers were intensively collected by Dan
L. Lindsley in 1998 and by Paul A. Opler and Evi
Buckner in 2000. All this collecting has yielded no
pyrrhopygine species that we have not reared. On the
other hand, six of our 15 reared pyrrhopygine species
have yet to be found as wild adults in the ACG.
PYRRHOPYGINE CATERPILLARS
Feeding behavior and growth rate. Pyrrhopygine
eggs are laid singly near the margins of mature leaves, on
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
the upper sides. If the foodplant grows in a sun/shade
mix, the leaves chosen are usually fully insolated.
The first instar cuts a 5-7 mm diameter disc nearly
all the way out of the leaf interior, or in from the mar-
gin, and silks the disc initially so as to fold it over on its
hinge, and then so as to form a shallow cup tightly
pressed to the leaf surface. As the caterpillar grows, it
builds and abandons successive nests (or shelters),
each larger than the last.
All instars live solitarily in their leaf/silk nests, ven-
turing out at night to feed. In later instars the nest
varies from a couple of overlapping leaves to several
leaves silked together. The leaf nests of pyrrhopygines
are among the most heavily silked of all ACG hesperi-
ids. Of the ACG pyrrhopygines examined, Pyrrhopyge
cosyra H. Druce and Passova gellias (Godman &
Salvin) make the strongest and densest nests.
Pupation occurs in a leaf/silk nest, which is usually
on the host plant and is usually the nest of the last in-
star. Occasionally a caterpillar makes its feeding and/or
pupation nest with the foliage of a plant other than its
host, generally when the foliage of the two plant
species is interleaved.
Each species of ACG pyrrhopygine caterpillar is
monophagous to oligophagous within a single plant
family—or, in one case, two plant families (Table 1).
(No laboratory trials were conducted to determine if
the caterpillars can feed and develop on species other
than those on which they are found in nature.) Be-
cause the ACG inventory of macrocaterpillars searches
all species of foodplants and has been going on for 22
years, further search probably will not expand the
plant lists for the caterpillar species in Table 1 except
to add a few more closely related species within the
genera or families already listed.
Pyrrhopyginae caterpillars are among the slowest
growing of the ACG Hesperiidae (Table 2). As a
group, they are conspicuously slower than Pyrginae
and Hesperiinae. Their prepupal stage lasts 3-5 days,
also among the longest in the ACG Hesperiidae. In no
instance has a reared pyrrhopygine prepupa become
quiescent or dormant in response to the dry season or
some other inimical time of year (other large ACG
hesperiids, such as the pyrgines Epargyreus and Ty-
phedanus ampyx [Godman & Salvin], do so as one of
their behaviors for passing part of the dry season).
Once pupation has occurred, their rate of develop-
ment (Table 2) is about the same as that of other hes-
periids of similar body weight. In no instance has a
pyrrhopygine pupa become seasonally dormant,
though there are several cases where the pupal stage
has lasted a few weeks more than expected. The cooler
VOLUME 55, NUMBER |
19
TABLE 2. Life table traits for wild-caught pyrrhopygine caterpillars reared at ambient temperatures on the species of foodplants on which
they were found, Area de Conservacion Guanacaste (data source: Janzen & Hallwachs 2001). The prepupal stage lasts about 3 days, but the pre-
pupa is not always located on its first day. A close approximation to pupal duration may be obtained by subtracting two days from the mean num-
ber of days between prepupa and eclosion. Extreme values are not included in the means.
First instar to prepupa (days)
Species Mean Range n
Pyrrhopyge zenodorus 78 61-93 13
Pyrrhopyge crida n/a n/a n/a
Pyrrhopyge aesculapus n/a n/a n/a
Pyrrhopyge cosyra
in Sector Cacao 104 92-137 21
in Sector San Cristobal 125 106-160 8
Pyrrhopyge erythrosticta n/a n/a n/a
Elbella scylla 89 82-96 9)
Parelbella macleannani 141 139-144 3
Jemadia pseudognetus 133 127-139 2
Mysoria ambigua 45 4\-47 3
Myscelus amystis hages 51 51 1
Myscelus belti 81 64-117 5
Myscelus perissodora n/a n/a n/a
Passova gellias 68 51-95 10
the weather, the longer the pupal duration, as evi-
denced by Pyrrhopyge cosyra (Table 2).
Color pattern and hairiness. ACG pyrrhopygine
caterpillars have one of two color patterns. The first is
basically dark with strongly contrasting yellow to or-
ange-yellow “rings” that are ventrally incomplete
(Figs. 2, 3, 7, 8, 9, 12) or lateral, vertical bars (Figs. 5,
6, 10, 11) or, in one case, large, orange lateral discs
(Fig. 4)—sometimes with red on the head and rump
(Figs. 2, 3, 8, 9) and ventrally as well (Fig. 12). These
pyrrhopygine caterpillars are thus part of an array of
presumably aposematic, mimetic ACG caterpillars
that involves hundreds of species, scores of genera,
and virtually all families (see photographs in Janzen &
Hallwachs 2001). The second pattern is dull green/
pink with black to brown/red heads (Figs. 13-15), a
seemingly cryptic color pattern shared with hundreds
of other species of ACG macrocaterpillars. In contrast
to many other ACG hesperiids (see Janzen &
Hallwachs 2001), none of the pyrrhopygine caterpil-
lars has false eye markings on the head.
All ACG pyrrhopygines in all instars have notably
long hairs on both the body and the head. They are the
only ACG skippers with long head hairs—except for
the pyrgine Astraptes fulgerator azul (Reakirt) whose
polymorphic caterpillars include one morph that su-
perficially resembles the yellow-ringed pyrrhopygine
caterpillars figured here (but A. f. azul lacks long body
hairs). Pyrrhopygine pupae are hairy over much of the
body and are often brightly colored (Figs. 16-25). Un-
like the caterpillars, the pupae of several pyrrhopy-
gines have conspicuous false eye markings, which will
Prepupa to eclosion (days)
Mean Range n Extremes
20.8 15-33 106 none
23.5 23-24 2 none
33.7 31-36 3 none
31.1 24-37 9] 52 (n = 1)
23.9 19-27 59 none
18.0 18.0 1 none
19.1 17-22 35 none
20.4 17-28 21 none
25.4 22-98 11 none
19.7 17-28 129 3448 (n = 4)
16.6 15-21 5 none
22.9 16-26 67 35-51 (n = 3)
24.0 24.0 1 none
21.1 15-34 50 none
be treated in a future study of this phenomenon across
all Hesperiidae (see Janzen & Hallwachs 2001).
SPECIES COMMENTARY ON PYRRHOPYGINES OF
THE ACG
In the course of reviewing the skipper fauna of the
world using the matchless holdings of the British Mu-
seum (Natural History), Evans became overly enam-
ored of the polytypic species concept and applied it far
too freely. In the brief introduction to his treatment of
the skippers of Europe, Asia, and Australia, Evans
(1949:xi) observed that
In accordance with modern ideas, as expressed in books such as
Mayr’s [1942] Systematics and the Origin of Species, as wide a view
as is possible has been taken of a species. Whenever a form in one
area can be considered as replacing a form in another area, the two
are presumed to be conspecific, even though the differences in fa-
cies, structure and genitalia appear considerable. The bringing to-
gether of subspecies in this manner presents no great difficulty
through the Malay Archipelago and the South Sea islands; for in-
stance, the variations of Tagiades japetus [(Stoll)] can be traced all
the way from Ceylon to the Solomons and compared with those of
other similar species flying with it. In continental areas, such as
China and Malaya, it is not always easy to decide whether two forms
flying together are species or overlapping sub-species of the same
species. There are difficulties also in the islands. For instance, in
Borneo there seem to be unduly numerous forms of the genus
Telicota, due perhaps to migrations or infiltration from Java, Timor or
the Philippines, and it is difficult to decide whether a particular series,
differing slightly in genitalia and facies from another series, represents
a species or a sub-species or perhaps a migration, which will become
submerged in due course by the dominant form in the island.
Evans used the same philosophy in his subsequent
treatments of pyrrhopygines (Evans 1951) and the rest
of the New World skippers. Although Evans's distribu-
tional data were incomparably rich, they were incom-
a reeeeeeeeeeeeemeemeemmememeeemmemmeememeerremrnr
plete. Because of this, because of his cavalier tolerance
of sympatry between subspecies, and because he
pushed to finish cataloging all the skippers on earth
before he died, he frequently lumped separate species
as subspecies of a single species. On finer analysis, his
polytypic species often turn out to comprise one or more
superspecies (e.g., Burns 1964) or species groups—or
even a distinct genus (Mielke 1995). We know that al-
lopatry, parapatry, or slight sympatry of closely related
species is frequent. What Evans called subspecies are
sometimes nothing more; but, for the reasons given
above, they should be reexamined whenever possible.
Pyrrhopyge zenodorus Godman & Salvin
According to Evans (1951), Pyrrhopyge zenodorus
is one of 10 subspecies of P. phidias (Linnaeus). How-
ever, especially on the basis of genitalic comparison
(Godman & Salvin 1893:pl. 73, fig. 3, Bell 1931:fig. 10,
Evans 1951:fig. A.1.2), P. zenodorus is a species in its
own right, ranging from central Mexico to Costa Rica,
at least.
While the caterpillars of P. zenodorus are moder-
ately common in the ACG rainforest, neither caterpil-
lars nor adults have been found in the dry forest, even
though patches of the food plant, Vismia baccifera (L.)
Triana & Planch. (Clusiaceae), are scattered through it
in moist places. This apparent absence from dry forest
is supported by the fact that the conspicuous caterpil-
lar nests have not been found there either. A V. bac-
cifera leaf is dark green above and bright rusty beige
below so that, when the caterpillar cuts a section and
folds it up and over, the resulting nest is a bold finger-
print that lasts for many months. Even where caterpil-
lars are found, adults are not usually seen. They rarely
have been encountered visiting flowers of Stachy-
tarpheta jamaicensis (L.) Vahl (Verbenaceae) at the in-
tersection of ACG dry forest and rainforest (in the Rio
Gongora region).
The adult of Pyrrhopyge zenodorus (Fig. 2), which
is blue-black, white-fringed, and red-orange on both
its head and rump, looks enough like the unrelated
Passova gellias (Fig. 15) to be indistinguishable from it
in flight (unless one can detect the marginal pale blue
on the dorsal hindwing of P. gellias). In flight,
Pyrrhopyge zenodorus superficially resembles one
other ACG rainforest skipper, the pyrgine Phocides
palemon lilea (Reakirt) (which, however, has a small
red bar at mid-costa of the forewing both above and
below and lacks the red-orange rump that gives
Pyrrhopyge its name).
The caterpillar of PR. zenodorus, which is ringed
brightly with orange-yellow (Fig. 2), superficially re-
sembles those of at least 12 other ACG rainforest skip-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
pers (Janzen & Hallwachs 2001), but none of these
feeds on Vismia baccifera: Pyrrhopyge crida (Hewit-
son) (Fig. 3), P. cosyra (Fig. 6), P. erythrosticta (God-
man & Salvin) (Fig. 7), Parelbella macleannani (God-
man & Salvin) (Fig. 9), Jemadia pseudognetus
(Mabille) (Fig. 10), another Jemadia belonging to a J.
hewitsonii species complex (Fig. 11), Phocides pale-
mon lilea (penultimate instar), Polythrix caunus (Her-
rich-Schiffer), Astraptes fulgerator azul, Nascus
broteas (Cramer), one of two species currently going
under the one name Nascus phocus (Cramer), and
Achlyodes busirus heros Ehrmann. The highly similar
caterpillars of Pyrrhopyge zenodorus and P. crida are
most readily distinguished by knowing what they were
found on (Vismia baccifera and the foodplant of P.
crida, V. billbergiana Beurl., grow within a few meters
of each other, with the former in the sun and the latter
more on the forest edge). Then, too, the purplish-red
hairs on the head and body of the caterpillar are not as
brilliant in P. zenodorus (Fig. 2) as they are in P. crida
(Fig. 3).
Caterpillars of P. zenodorus are oligophagous on
mature leaves of the shrubby treelets V. baccifera and
V. ferruginea Kunth (Table 1). Vismia baccifera is
widespread and common in the early successional
stages of old ACG rainforest pastures returning to for-
est. When the area was entirely old-growth forest, V.
baccifera was probably a much rarer plant restricted to
the early stages of tree-fall succession and continually
disturbed sites such as watercourse edges and land-
slides. A few hours’ search can usually produce at least
a few old nests of P. zenodorus caterpillars on V. bac-
cifera, and currently this plant supports the vast bulk
of the population. Vismia ferruginea is very rare, oc-
curring just in the interface between dry forest and
rainforest (Estacién Biolégica Maritza area). Despite
intensive search, only five caterpillars have been found
on it.
Moss (1949:33) reared other members of the
Pyrrhopyge phidias species complex (to which P. zen-
odorus belongs) without knowing exactly what they
were: “Amongst the commonest of the Pyrrhopyge in
Paré [Brazil], were a number of species closely resem-
bling in their underside coloration [P. phidias hyperici
(Hiibner) |. Unfortunately I took them for one single
species and in consequence did not make detailed ob-
servations on their separate larvae and pupae. Their
tent-like shelters are found on the upper side of the
leaves of Lacre bushes (Vismia guianensis (Aubl.)
Choisy., [Clusiaceae]). They also frequently occur[rJed
on Guava and Aracd (Myrtaceae).”
In later instars, P zenodorus, like P. crida and P. aes-
culapus Staudinger, only lightly silks a pair of leaves to-
VOLUME 55, NUMBER 1
Area de Conservacion Guanacaste
384000 East Lambert
85° 34’ 38" West Longitude
Murcle
we tet
Islas Murciélago —>
Océano
Pacifico
Los Almendros’
Tay
ce edi
NX ge Virgen,
_ *tiberia
aE & Santa Cecilia
| Dry Forest
[-]] Rainforest
es} Cloud Forest
322000 North Lambert
10° 55' 4” North Latitude
ut
x -Caribe
“ ‘
Srebbak *
ae Administrative station
’ % Cities and towns
& Biological station
/\/ Main roads
/s/ International border
Interamerican highway
[__] Protected area
Elaborated by Waldy Medina
Area de Conservacion Guanacaste
Printed August 2000
Fic. 1. Area de Conservacién Guanacaste (ACG) in northwestern Costa Rica and the locations of its three major ecosystems.
gether to make its nest, in striking contrast to the other
pyrrhopygines, most of which make nests with much
more extensive and dense silk linings.
Pyrrhopyge zenodorus is one of the three pyrrhopy-
gine caterpillars with more or less host-specific ta-
chinid fly parasites. Eleven of the records of Houghia
sp. 9 are from this skipper (the other record is from
Phocides nigrescens Bell); and all three records of
Pseudosturmia sp. | are from Pyrrhopyge xenodorus.
The distinctive Apanteles wasp (Braconidae) that at-
tacks P. zenodorus (n = 20) may also turn out to be
host-specific.
Pyrrhopyge crida (Hewitson)
Superficially, the adult of Pyrrhopyge crida (Fig. 3)
does not closely resemble other pyrrhopygines or
other ACG hesperiids. The combination of uniform
black (both dorsally and ventrally) with an oblique,
wide, bright white, hyaline slash tapering across the
forewing from below the mid-costa to above the tor-
nus—along with a narrow, off-white fringe on the
hindwing and an orange head and ruamp—is unique
(though a number of ACG pyrgines are very dark with
a row of white hyaline spots angling across the
forewing).
Pyrrhopyge crida is a rainforest skipper in the ACG.
Knowledge of its presence comes from 14 rearings,
and the capture of a single wild female, by P. Rios and
C. Moraga, parataxonomists at Estacion Biol6gica Pit-
illa in the northeastern ACG. Though clearly a wide-
ranging species, P. crida is rare in collections. For ex-
ample, there are only § specimens in the BMNH
(from Nicaragua, Colombia, and Ecuador) and 5 in the
USNM (from Costa Rica, Panama, and Colombia). It
extends north to southern Mexico (Hoffmann 1941).
The caterpillar of P. crida, unlike that of P. zeno-
22
dorus (which occurs throughout the rainforest on the
widespread Vismia baccifera), is apparently restricted
to V. billbergiana (Table 1), found to date only in the
understory of old secondary successional rainforest at
about 600 m along the entrance road to Estaci6n Bio-
logica Pitilla. The caterpillar of P. crida (Fig. 3) looks
very like that of P. zenodorus (Fig. 2). However, the
hairs on the head and the shorter hairs on the body of
P. crida are more brilliantly purplish-red than they are
in P. zenodorus.
Although Evans (1951) included P. zenodorus in his
second species of Pyrrhopyge and treated P. crida as his
thirty-fifth species (five species groups removed from P
zenodorus), the two seem closer. This is suggested by
similarities not only in the caterpillars, their foodplants,
and how lightly they silk leaves of those plants together
when making a nest, but also in basic genitalic form of
the adults (Godman & Salvin 1893:pl. 73, fig. 3 & pl.
74, fig. 3, Evans 1951 :figs. A.1.2 & A.1.35).
Pyrrhopyge creon H. Druce
The wings of this stunning skipper are iridescent
blue, with black edges and fringes and a red-orange
spot above the tornus of the hindwing (in space Ic);
this prominent spot is expressed on both surfaces, but
is a little smaller dorsally than ventrally. Evans (1951)
treated Pyrrhopyge creon as a monotypic species, rep-
resented in the BMNH by specimens from Costa Rica
and Panama (plus a pair from Colombia). Much later,
Nicolay and Small (1969, 1981) described two sub-
species—superficially quite distinct from each other
and from P. creon creon—from cloud forest on isolated
mountains of Panama and Costa Rica, respectively:
Pyrrhopyge creon lilliana, in which the red-orange
subtornal spot of the hindwing is not only enlarged but
also expanded into a series of three to five submarginal
spots (in spaces 1b to 4); and P. c. taylori, in which the
iridescent blue of both pairs of wings is replaced by iri-
descent purple, while the subtornal spot remains un-
changed.
Pyrrhopyge c. creon ranges along the continental di-
vide from Guanacaste, Costa Rica, to Coclé, Panama;
because it was never found farther east in suitable
Panamanian habitat, despite intensive efforts, records
from Colombia seem doubtful (Nicolay & Small 1981).
Both P. c. taylori (at 1150 m in the Fila Cruces,
Puntarenas, Costa Rica) and P. c. lilliana (at 800 m at
Cerro Campana and La Mesa in a disjunct massif in
the provinces of Panama and Coclé, Panama) are sepa-
rated from typical P. c. creon by relatively low-elevation
gaps of just 30-40 km. Since intergrades are unknown,
these skippers, though strong fliers, are presumably
sedentary (Nicolay & Small 1981).
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Male genitalia, which are moderately asymmetric in
the distal part of the valvae (Godman & Salvin 1893:pl.
73, fig. 11, Bell 1931:fig. 45), are identical in all three
subspecies (Nicolay & Small 1969:fig. 3, 1981:fig. 2). In
comparing the genitalia of 14 d and 3 ° that represent
these taxa, we, too, found no significant differences.
Pyrrhopyge c. creon is the epitome of cloud forest
hesperiids. Adults are occasionally seen flying and
perching on leaves 2-10 m above the ground in the
upper parts of tree falls on sunny/warm to foggy/cool
days on the peak of Voleén Cacao (1200-1500 m).
Their presence in this elevated habitat mirrors their
presence at 1600-2000 m—flying even in heavy fog—
on Volcén Barba and Volcan Pods, further south in
Costa Rica. Pyrrhopyge c. creon flies in fog so heavy
that its wings may be covered with fine beads of mois-
ture. No other diurnal ACG butterflies have been seen
on the wing under these conditions.
In flight in the ACG cloud forest, this large blue-
black skipper can be confounded with P. aesculapus.
When P. c. creon perches, the red-orange subtornal
spot of the hindwing sets it apart.
A pupa found at 1550 m, on the summit of Volcan
Cacao, produced our only reared adult of P. c. creon.
However, two pyrrhopygine caterpillars whose dark
bodies sported large orange dots (Fig. 4)—instead of
bars or rings—were found somewhat lower (at about
1200 and 1000 m) on Dendropanax querceti Donn.
Sm. and D. gonatopodus (Donn. Sm.) A. C. Sm. (Ar-
aliaceae). The first died of apparent disease in the
penultimate instar while the second pupated, only to
sprout a mass of white fungus. Though we have yet to
rear the adult we need for positive identification of
these odd caterpillars, we strongly suspect—from their
elevation and habitat—that they are those of P. c.
creon. But, because we cannot be certain, we have de-
liberately omitted photographs of the presumed adult
from Fig. 4.
Pyrrhopyge aesculapus Staudinger
Pyrrhopyge aesculapus presents no taxonomic prob-
lems at the species level. The adult has iridescent pur-
plish-blue wings with a wide, strongly contrasting, or-
ange fringe on the hindwing but not on the forewing
(Fig. 5) (except for traces in space 1b in some individ-
uals). The hindwing is purpler and darker, whereas the
forewing is bluer. The body is dull because it lacks the
orange or red hot tail for which many pyrrhopygines
are famous; nor is it “hotheaded.” One feature of the
distinctive male genitalia—the elongate caudal projec-
tion comprising the distal half of the valva—can be ex-
posed relatively easily with dry dissection. In lateral
view, this projection looks peculiar because it is almost
VOLUME 55, NUMBER |
uniformly wide, marginally dentate, terminally blunt,
and evenly curved upward from its horizontal begin-
ning to its vertical end (Godman & Salvin 1893;pl. 73,
fig. 9, Bell 1931:fig. 43, Evans 1951 :fig. A.1.46).
Known mainly from Costa Rica and Panama, P. aescu-
lapus apparently ranges south to Colombia and
Ecuador.
In flight in the ACG cloud forest, the large blue-
black P. aesculapus resembles only P. c. creon. At rest,
these species are readily distinguished by their hind-
wings: orange-fringed in the former (Fig. 5), spotted
red-orange above the tornus in the latter.
In this cloud-forest habitat, the caterpillar of P. aes-
culapus, with bright yellow, lateral, vertical bars on a
black background (Fig. 5), resembles the barred cater-
pillar of P. cosyra (Fig. 6) and, to a lesser extent, the
ringed caterpillars of Ridens mephitis (Hewitson), As-
traptes fulgerator azul, and a species of Venada. How-
ever, none of these caterpillars feeds on Weinmannia
wercklei Standl. (Cunoniaceae), the sole known food-
plant of P. aesculapus (Table 1). Weinmannia wercklei
is a shrubby treelet characteristic of high elevations in
Costa Rica. Pyrrhopyge aesculapus spins its nest
lightly and conspicuously among the thin, flimsy leaves
of this plant. Tearing at the nest easily exposes the
boldly patterned caterpillar which does not try to stay
hidden in the nest but actively crawls away, as if to
flaunt its colors.
Too few examples of P. aesculapus have been reared
to draw conclusions about its lack of parasitoids (Table
1). In the habitat, and at the elevations, where P. aes-
culapus occurs, caterpillars of P. cosyra are attacked by
a host-specific tachinid fly.
Pyrrhopyge cosyra H. Druce
Of the many species currently in Pyrrhopyge, this is
one of at least 15 in which the valvae of the male geni-
talia are asymmetric (Godman & Salvin 1893:pl. 74,
fig. 2, Evans 1951:fig. A.1.55) (others include P. creon
and members of the P. maculosa species complex). Su-
perficially, P. cosyra is extremely distinct: both pairs of
wings are iridescent dark blue, but dorsally they sport
a huge, bold, proximal, orange splotch; the forewing
has three distal bands that are white and hyaline
(hence ventral as well as dorsal); and the body, except
for the dorsally orange thorax, is banded black and
white (Fig. 6). In the only other pyrrhopygine that re-
sembles it—a South American species, P. spatiosa
(Hewitson)—the splotch is a redder orange. Pyr-
rhopyge erythrosticta (whose ground color is black in-
stead of iridescent dark blue) presents a color pattern
basically like that of P. cosyra, except that the huge,
central, orange splotch is ventral instead of dorsal (Fig.
23
7). Pyrrhopyge cosyra ranges from Guatemala through
Central America to Colombia and Ecuador,
In Costa Rica P. cosyra occurs from lowland rainfor-
est (sea level on the Osa Peninsula in the southwest) to
ACG rainforest at 400-800 m and extends up into
cloud forest at 1400 m. Neither adults nor caterpillars
have been found in ACG dry forest, even though one
of its foodplants, Clusia rosea Jacq. (Clusiaceae), is
sporadic in wet sites there. In ACG cloud forest and
rainforest, adults have been seen only when they de-
scended from the tree canopy for brief exploration of
potential foodplants. The relatively common caterpil-
lars of P. cosyra are found less than 3 m above the
ground.
The caterpillar of P. cosyra, which is black with or-
ange-yellow, lateral, vertical bars (Fig. 6), looks super-
ficially a lot like the larvae of both Jemadia species
(Figs. 10, 11) and much like those of Pyrrhopyge zen-
odorus (Fig. 2) and several other rainforest skippers
listed in our account of that pyrrhopygine. However,
none of these species feeds on Chrysochlamys
(=Tovomitopsis) or Clusia, the only foodplants of P
cosyra (Table 1).
On small-leaved plants such as Chrysochlamys and
Clusia minor L., the caterpillar of P cosyra spins an
extraordinarily tough and baglike silk nest. It is hard to
tear open; and when one does, the caterpillar retreats
inside in an apparent effort to stay out of sight. When
feeding on the other species of Clusia with their mas-
sive thick leaves, the caterpillar silks a couple of leaves
very tightly together into an equally strong nest.
Pyrrhopyge cosyra is the most heavily parasitized of
all the pyrrhopygines reared, and the only one at-
tacked by Tachinidae, Ichneumonidae, and Bra-
conidae (Table 1). The tachinid Chlorohystricia sp. 1
appears to use it almost exclusively (18 records on P.
cosyra plus one on Venada sp. in the same habitat) and
occurs throughout the caterpillar’s range in the ACG
(400-1400 m). Pyrrhopyge cosyra is the sole known
host for the ichneumonid genus “MCAM” sp. | (n =
10); and it is heavily used by Casinaria sp. 9, an ich-
neumonid wasp that also attacks a few species of
pyrgines. The lone braconid individual reared from
this skipper has yet to be placed to genus.
Pyrrhopyge erythrosticta (Godman & Salvin)
For their original description of Pyrrhopyge ery-
throsticta, Godman and Salvin (1879) used very few
words, no illustrations, and an unspecified number of
specimens (from Chontales, Nicaragua, and Veragua
[part of central Panama]) without designating a type.
Even in the Biologia Centrali-Americana, Godman
and Salvin (1893) did not picture an adult; but they did
—————————
ee Se ae
Fe ee
ee ee
eS ee
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics. 2-4. Last instars, plus adults in dorsal (L) and ventral (R) view, of ACG pyrrhopygines. 2, Pyrrhopyge zenodorus (rainforest) 94-
SRNP-707, L/R 2 94-SRNP-6631. 3, Pyrrhopyge crida (rainforest) 97-Rios-491, L/R ° 97-Cali-188. 4, Pyrrhopyge creon presumably (cloud for-
est) 0O-SRNP-20314.
VOLUME 55, NUMBER |
Fics. 5-7. Last instars, plus adults in dorsal (L) and ventral (R) view, of ACG pyrrhopygines. 5, Pyrrhopyge aesculapus (cloud forest) 97-
SRNP-513, L/R 6 97-SRNP-514. 6, Pyrrhopyge cosyra (rainforest to cloud forest) 98-SRNP-2767, L 2° 97-SRNP-1805, R 2 97-SRNP-6531. 7,
Pyrrhopyge erythrosticta (rainforest) 00-SRNP-15831, L/R 6 00-SRNP-15831.
what was much better in the end by clearly figuring
(vol. 3, pl. 73, fig. 13) and verbally describing (vol. 2, p.
252) the “very peculiar” male genitalia of P. erythro-
sticta. Having found that their type series includes
more than one genitalically divergent species, Burns
hereby chooses as lectotype (BMNH) the male whose
genitalia were shown. This action fixes the taxonomic
concept of the name and ensures its future consistent
interpretation. The lectotype bears a distinctive God-
man-Salvin miniature genitalia slide (numbered 369)
plus three labels: (1) Chontales, / Nicaragua. / T.Belt.
(2) B.C.A. Lep. Rhop. / Pyrrhopyge / erythrosticta, / G.
& S. (3) Godman-Salvin / Coll.1912.-23. Its left wings
have been bleached for study of venation. This is the
species that we have reared in the ACG. At present we
know it only from Nicaragua and Costa Rica. The
more wide-ranging entity that has been confused with
it will be described separately, in the context of the en-
tire P. maculosa complex.
While we were writing the paper you hold, ACG
parataxonomists Petrona Rios and Calixto Moraga re-
ported that twice in 1996 they had seen yellow-ringed,
black pyrrhopygine caterpillars on Marila laxiflora
Rusby (Clusiaceae) at Estacién Bioldgica Pitilla (in
rainforest at 700 m on the northeast side of Volcan
Orosi) but had been unable to rear them. Asked now
to try for more, Rios found a single last instar (Fig. 7)
in its nest on a non-host plant—i.e., a plant that the
caterpillar was not eating in the field and would not eat
in captivity. It fed normally on M. laxiflora until pupa-
tion (Fig. 19) and produced our lone reared adult (Fig.
7). Two wild adults of P. erythrosticta have been
caught in the rainforest at 600-800 m in the pass be-
tween Volcan Cacao and Volcan Rincén de la Vieja.
This species has not been seen anywhere else in the
ACG.
Pyrrhopyge erythrosticta gets its specific name from
the small, dull red spot near the base of the dorsal
forewing between veins 1 and 2 (Fig. 7); but the
forewing also bears nine, more conspicuous, small,
white hyaline spots in fixed positions. It is important
that in true P. erythrosticta, the lowest hyaline spot,
which is in space 1b, is directly under the spot in space
2 (Fig. 7). The spot in space 1b is likewise under the
spot in space 2 in a more northern species, P. hoff-
manni Freeman, to which P. erythrosticta is also geni-
talically related. However, P. hoffmanni is much more
closely related to the still more northern P. mulleri
(Bell): this pair has essentially the same genitalia in
both sexes (for males, cf. Bell 1934:fig. 5 and Freeman
1977:fig. 17), but P. mulleri has lost all hyaline spots.
The members of the P. mulleri-P. hoffmanni / P. ery-
throsticta trio appear to replace each other geographi-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
cally. Their apparent parapatry independently sup-
ports their grouping on morphological grounds. On
the other hand, both P. hoffmanni and P. erythrosticta
coexist with the species that is currently confounded
with the latter. In this unnamed species, the spot in
space lb is not under the spot in space 2 but, instead,
displaced outward so that the spots in spaces 1b, 2, and
the cell form a straight line—as they also do in its more
southern (and closer) relative, P. maculosa Hewitson.
Elbella scylla (Ménétriés)
Elbella scylla amounts to much more than the latest
taxonomic revision implies. Mielke (1995) treats E.
dulcinea (Plétz) as a different species, occurring in
Mexico with E. scylla and extending farther south than
E. scylla into northern South America (see map plot-
ting both in Mielke 1995:fig. 44); but all this is one
species ranging from central Mexico to Colombia,
Venezuela, and Ecuador. Its genitalia (Godman &
Salvin 1893:pl. 73, fig. 6, Bell 1931:fig. 35, Evans
1951:fig. A.2.1, Mielke 1995:figs. 15, 16) look the same
(within normal limits of individual variation) every-
where (41 closely compared KOH-dissections: Mexico
3 6, Guatemala 1 d 1 9, Honduras 1 d 1 2, Costa Rica
14 d 14 9, and Panama 3 d 3 9). Elbella scylla (de-
scribed in 1855 from Nicaragua) and E. dulcinea (de-
scribed in 1879 from the part of Colombia that, in
1903, became Panama) were considered synonymous
in the 1940s. However, Evans (1951) called dulcinea a
subspecies of scylla; and Freeman (1966:227)—who
claimed, without specifics, that the two “differ some-
what genitalically and morphologically” and “occur to-
gether in the same areas of Veracruz,” Mexico—raised
dulcinea back to the level of species (for complete ref-
erences, see Mielke 1995:460).
The main color character that “distinguishes” dul-
cinea from scylla—i.e., inward expansion of white
from the white hindwing fringe into the outer margin
of the black hindwing proper—is variably expressed by
most of the ACG specimens of E. scylla. Since this
“white expansion” is more pronounced and more uni-
formly expressed in specimens from Panama, and
since it is still more pronounced in Venezuela (see
Mielke 1995:fig. 87), it is apparently a clinally varying
trait in southern portions of E. scylla’s range. (The
“white expansion” is better expressed ventrally than
dorsally in every individual showing it; furthermore, it
is sexually dimorphic and better expressed by females
than by coexisting males.) Geographic variation in
fringe-whitening, and in the extent to which white in-
vades the dark hindwing itself, has been well docu-
mented in the pyrgine genus Erynnis in which a white
fringe has evolved eight times independently, always in
VOLUME 55, NUMBER 1
those populations occurring in the southwestern USA
and Mexico plus points south (Burns 1964). In Elbella
scylla a color character of the tegulae—whether their
longitudinal stripe is yellowish, reddish-orange, or
bluish-white (it is occasionally unexpressed)—also
varies geographically, but not in concordance with the
clinal variation noted above. In ACG material the
stripe is reddish-orange.
Adults of E. scylla are occasionally encountered vis-
iting flowers (e.g., Trigonia rugosa Benth. [Trigoni-
aceae], Cordia spp. [Boraginaceae]) in the ACG dry
forest and on the dry side of the intergrades between
dry forest on the one hand and rainforests and cloud
forests on the other in the eastern ACG. Neither
adults nor caterpillars have ever been encountered in
rainforest or cloud forest.
Although the adult of E. scylla does not closely re-
semble adults of other ACG dry-forest hesperiids, it
can—in flight—be confused with two other large and
generally black skippers, Mysoria ambigua (Mabille &
Boullet) and Phocides palemon lilea. The former
species has yellow and magenta along the outer and
costal margins, respectively, of the underside of the
hindwing (Fig. 12), while the latter species has a small
red bar at mid-costa of the forewing both above and
below.
The brightly yellow-ringed caterpillar of E. scylla
(Fig. 8) superficially resembles those of at least 13
other ACG dry-forest skippers (Janzen & Hallwachs
2001): Mysoria ambigua (Fig. 12), Phocides palemon
lilea, Drephalys alemon (Cramer) and D. kidonoi
Burns (see Burns & Janzen 2000:figs. 21-25), Poly-
thrix caunus, Chrysoplectrum pervivax (Hiibner), Ri-
dens mephitis, Astraptes fulgerator azul, Nascus solon
corilla Evans, two species currently going under the
one name Nascus phocus, Pellicia dimidiata Herrich-
Schiffer, and Achlyodes busirus heros, all of which ap-
pear to be involved in an as yet undescribed, multi-fa-
milial, aposematic, caterpillar mimicry complex.
Caterpillars of E. scylla are oligophagous on mature
leaves of a small subset of the species of ACG
Malpighiaceae plus two species of Combretaceae
(Table 1). Most caterpillars eat Byrsonima crassifolia
(L.) Kunth (Malpighiaceae), a shrubby tree that is
common in the old and frequently bummed pastures cut
out of ACG dry forest, but that also persists in many
kinds of old-growth forest ranging from exposed sea
cliffs to deep soils on the volcanic foothills. The other
three malpighiaceous foodplants are woody perennial
vines occurring commonly on forest edges and in un-
derstory of secondary succession. All four of these
foodplants probably will be rarer and more locally dis-
tributed when the ACG dry forest has returned to full
NS)
|
old-growth status. Other common dry-forest Malpighi-
aceae (Heteropterys obovata |Small] Cuatrec. &
Croat, Gaudichaudia hexandra Chodat, Stigmaphyl-
lon ellipticum A. Juss., S. lindenianum A. Juss.,
Mascagnia sinemariensis Griseb., Bunchosia biocellata
Schlecht., Malpighia glabra L., Banisteriopsis cornifo-
lia C. B. Rob.), which occur both on edges and in
heavily-shaded understory of deep forest, have never
yielded E. scylla caterpillars. Neither have any of the
other five or so species of malpighiaceous treelets and
shrubs (e.g., Tetrapterys discolor G. Mey., Hiraea sp.,
Bunchosia sp.) in the wetter portions of the ACG pro-
duced these caterpillars, despite intensive search.
Combretum farinosum H. B. & K. is an occasional
woody vine and scandent shrub that grows along
ravines and watercourses, and E. scylla apparently eats
its semi-evergreen foliage primarily in the dry season.
This is the only native species of Combretaceae in the
ACG dry forest. The others are in the mangroves or in
local riparian wet sites, and we do not know whether
they, too, would be acceptable. Terminalia catappa L.
is an introduced beach-edge and garden evergreen
combretaceous tree that is very rarely used by E.
scylla. Inclusion of Combretaceae in the diet of a
malpig-eater is mirrored by the pyrgine genus
Cephise: caterpillars of the tailless species Cephise
nuspesez Burns as well as the tailed species C. auginu-
lus (Godman & Salvin)—which now is considered a
synonym of C. aelius (Plétz) (Austin & Mielke 2000)—
feed heavily on Malpighiaceae and also use Combre-
taceae (Burns 1996, Janzen & Hallwachs 2001). There
is no hint that E. scylla caterpillars feed on any plants
in the ACG besides the four Malpighiaceae and the
two Combretaceae in Table 1.
For E. dulcinea—which is synonymous with E.
scylla—Mielke (1995) reported that, in Maracay,
Aragua, Venezuela, the caterpillar was reared by F. F.
Yépez on Terminalia catappa. This Asian introduction
was recorded once as a foodplant in the ACG.
In Para, Brazil, Moss (1949) reared larvae of E. in-
tersecta (Herrich-Schiffer) on at least three unspeci-
fied species of Malpighiaceae. Elbella scylla and E. in-
tersecta are very closely related and, since the latter is
primarily South American in distribution, mostly al-
lopatric. For E. intersecta intersecta, Mielke (1995)
stated that larvae are very common on Vismia guia-
nensis (Clusiaceae) in Imperatriz, Para; and for E. i.
peter Evans, he simply listed Vismia guianensis and
Banisteria mossii Morton (Malpighiaceae) as food-
plants.
Elbella scylla appears to be attacked by only one
species of parasitoid, a microgastrine braconid (Cote-
sia) (Table 1). A single individual was found with tachi-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics. 8-10. Last instars, plus adults in dorsal (L) and ventral (R) view, of ACG pyrrhopygines. 8, Elbella scylla (dry forest) 96-SRNP-224,
L/R 2 93-SRNP-6852. 9, Parelbella macleannani (rainforest) 01-SRNP-113, L/R ° 98-SRNP-6468. 10, Jemadia pseudognetus (rainforest) 00-
SRNP-21908, L/R 3 98-SRNP-6196.
VOLUME 55, NUMBER 1
Fics. 11-13. Last instars, plus adults in dorsal (L) and ventral (R) view, of ACG pyrrhopygines. 11, Jemadia sp. X of a J. hewitsonii species
SRNP-272, L 2° 87-SRNP-1162, R d 95-SRNP-
30
nid eggs glued on the front of the head, but neither
flies nor caterpillar survived the rearing process.
Parelbella macleannani (Godman & Salvin)
Evans (1951) grouped five subspecies in his poly-
typic species Elbella polyzona (Latreille). Mielke
(1995) regards this assemblage as a new genus, Parel-
bella; treats four of the subspecies as three distinct
species (one that is polytypic includes not only two of
those four subspecies but also the fifth as a synonym);
and adds a new species. One of Parelbella’s four
species extends north of South America.
Parelbella macleannani ranges from northern
Ecuador (Imbabura) to southern Mexico (Veracruz,
Oaxaca), with intervening records from Colombia,
Panama, Costa Rica (ours), Nicaragua, and Guatemala
(Mielke 1995:fig. 51, 561-562). In the ACG, its cater-
pillars have been found only in the rainforest from
400-800 m—never in the dry forest—and free-flying
adults have not been seen or collected.
Parelbella and Jemadia (the genus we consider next)
are among those distinctive pyrrhopygines in which
adults with a more or less black ground color are
striped and banded pastel blue over the body and on
both pairs of wings and, in addition, are banded hya-
line white on the forewings (Figs. 9-11). (Some
species of the pyrgine genus Phocides have indepen-
dently evolved this striking color pattern; but their
stout antennal club—in sharp contrast to that of
pyrrhopygines—ends in a long, delicate, reflexed
apiculus. Nevertheless, in life these pyrgines fold the
apiculus back so tightly against the body of the club
that even the club suggests that of a pyrrhopygine.)
Though at first glance Parelbella macleannani and
ACG species of Jemadia look much the same, they do
differ—not just in major genitalic ways (cf. Mielke
1995: figs. 32-36 and Evans 1951:figs. A.5.1-A.5.7) but
in wing pattern and shape (cf. Figs. 9-11). In P.
macleannani the slender hyaline spots in spaces 3 and
4 of the forewing (which, at times, are barely or not ex-
pressed) are widely disjunct, whereas in Jemadia these
two spots are in contact and aligned. Parelbella
macleannani has a fine blue line (expressed much bet-
ter ventrally than dorsally) at the distal end of the
forewing cell, but Jemadia does not. In P. macleannani
the inner (basal) blue band on the dorsal hindwing is
single; in Jemadia it is double. The submarginal blue
band on the ventral hindwing of P macleannani
(which is not as well-developed as the three more
proximal blue bands) is lacking in Jemadia. And there
is a small lobe at the hindwing tornus in P. maclean-
nani but not in Jemadia.
Caterpillars of P. macleannani are monophagous to
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
oligophagous on mature leaves of one to three species
(some determinations pending) of Eugenia (Table 1),
These Myrtaceae are forest-understory-to-edge scan-
dent shrubs, on which the caterpillars were all found
within 2 m of the ground. Given that there are more
than 30 species of Myrtaceae in the ACG and that P
macleannani has been found on only one to three con-
generic species, it may eat just it or those. On the
other hand, the caterpillar of this skipper occurs at
such low density that, with more years of search, its list
of myrtaceous foodplants may increase.
In Para, Brazil, Moss (1949) found caterpillars of
what Mielke (1995) treats as Parelbella ahira ahira
(Hewitson) on an undetermined species of Eugenia
(and once also on Virola sebifera Aubl. [Myristi-
caceae]). He described both larvae and pupae as
strongly crimson and indicated that the larvae have
transverse yellow belts.
The caterpillar of Parelbella macleannani is reddish-
black with orange-yellow rings (Fig. 9). This color pat-
tern basically resembles those of various other ACG
rainforest skippers such as Pyrrhopyge zenodorus, as
listed in the account of that species. Except for the
pyrgine Phocides palemon lilea, these other skippers
do not feed on Myrtaceae.
Our only parasitized caterpillar of Parelbella
macleannani (Table 1) was attacked by Hyphan-
trophaga virilis (Aldrich & Webber), which is the most
extreme generalist of all ACG tachinids reared.
Jemadia pseudognetus (Mabille), reinstated status
Evans's (1951:50) polytypic species Jemadia hospita,
ranging from Guatemala to Peru, Bolivia, and Brazil,
includes four subspecies. The two subspecies that are
widely distributed are also the first to have been,de-
scribed (originally as species): Jemadia hospita hospita
(Butler, 1877) and J. hospita pseudognetus (Mabille,
1878). Repeated comparison of the genitalia of 20 3
shows that these two taxa are indeed separate species,
with J. pseudognetus ranging from southern Mexico to
Colombia, at least, and with J. hospita strictly South
American (like the other taxa in this complex). The
characters given by Evans (1951:50-51) for superfi-
cially distinguishing between pseudognetus and hos-
pita are good. Differences in the male genitalia (not
noted by Evans) are subtle and involve a narrow,
pointed, finely dentate projection that sweeps an-
terodorsally from the posterior half of the valva: this
projection is shorter and a little less dentate in J.
pseudognetus (Godman & Salvin 1893:pl. 74, fig. 6)
than it is in J. hospita (Bell 1933:fig. 24, Evans
1951 :fig. A.5.1).
Caterpillars of J. pseudognetus have been found
VOLUME 55, NUMBER 1
only in the rainforest from 400-800 m, and free-flying
adults have never been seen or collected in the ACG.
Since no caterpillars have been found in the dry forest,
J. pseudognetus is clearly a rainforest skipper within
the ACG.
Several differences in external appearance between
adults of ACG species of Jemadia and those of Parel-
bella macleannani are listed under the latter species.
But J. pseudognetus specifically differs from P. mac-
leannani in another respect: the dorsal hindwing has
only two pastel blue bands (inner and outer), instead of
three (inner, central, and outer) as in P. macleannani.
Caterpillars of J. pseudognetus are oligophagous on
mature leaves of Lauraceae (Table 1). Although there
are more than 30 species of Lauraceae in the ACG, J.
pseudognetus has been found on only six and may be
limited to them. But, because the density of cater-
pillars is low, the list of lauraceous foodplants may, in
time, increase. All known foodplants are forest-
understory-to-edge saplings and treelets, and all cater-
pillars have been found within 2 m of the ground.
The caterpillar of J. pseudognetus is black with
short, lateral, vertical bars of orange-yellow (Fig. 10).
More or less similar barred color patterns occur in
Pyrrhopyge aesculapus (Fig. 5), P. cosyra (Fig. 6), and
another species of Jemadia (Fig. 11) to be treated next.
However, each of these ACG wet-forest pyrrhopygines
eats plants in a different family (Table 1).
New member of a Jemadia hewitsonii
species complex
Evans's (1951:53) polytypic species Jemadia hewit-
sonii, which is essentially South American, contains
five subspecies: Jemadia h. pater Evans, 1951, J. h.
ovid Evans, 1951, ]. h. albescens Rober, 1925, ]. h. he-
witsonii (Mabille, 1878), and J. h. brevipennis Schaus,
1902. Consideration of the color-pattern characters
used by Evans to distinguish among these taxa, cou-
pled with comparison of their genitalia (from a total of
30 d—including the holotype of J. brevipennis—in the
USNM), suggests that several closely related species,
rather than subspecies, comprise a J. hewitsonii
species complex. Genitalic differences among these
taxa are modest: for example, the dorsally dentate dis-
tal portion of the valva is relatively long, finely dentate,
and lower distally than proximally in J. hewitsonii (Bell
1933:fig. 27) but relatively short, coarsely dentate, and
fully as high distally as proximally in J. pater, new sta-
tus (Godman & Salvin 1893:pl. 74, fig. 9). Genitalic
study further shows that J. brevipennis is not what
Evans had in mind but, instead, a species well-re-
moved from the rest and closely allied with J. gnetus
(Fabricius).
31
Although Evans recorded J. pater from Venezuela
(3 5 2), Colombia (65 d 1 2), and Panama (1 4 1 9), it
is not the northernmost member of the J]. hewitsonii
species complex. What is, is something deep in the
rainforest of the ACG known from 16 caterpillars
found eating Casearia arborea Urb. (Flacourtiaceae)
(Table 1) and from the three adults (1 ¢ 2 2) they pro-
duced. More adults (especially males) are desired be-
fore formal description of this apparent differentiate.
Here we simply call it Jemadia sp. X. Its caterpillar is
black with short, lateral, vertical bars of bright, clear
yellow (Fig. 11). These short, lateral bars are orange-
yellow in J. pseudognetus (Fig. 10).
At both Obidos and Santarém, Pard, Brazil, Moss
(1949) found caterpillars of J. gnetus feeding on a dif-
ferent flacourtiaceous plant, Laetia corymbulosa
Spruce.
Adults of Jemadia sp. X look like those of its ACG
congener, except that, on the dorsal hindwing, between
the inner and outer pastel blue bands, there is a short,
blue, central band (running only from veins 3 to 7),
which is absent in J. pseudognetus (cf. Figs. 10 and 11).
Mysoria ambigua (Mabille & Boullet)
Although Evans (1951) saw Mysoria ambigua as the
most northern of six subspecies comprising polytypic
species M. barcastus (Sepp), comparative study—par-
ticularly of genitalia (note major valval differences de-
picted in Evans 1951:fig. A.12.1)—shows that M. am-
bigua is a Separate species, occurring from Mexico to
Costa Rica.
In the ACG, M. ambigua frequents dry forest and
the dry side of the intergrades between dry forest and
wetter forests to the east. Neither adults nor caterpil-
lars have ever been encountered in rainforest or cloud
forest itself. When flying in the ACG dry forest, M.
ambigua can be confused with two other large and
generally black skippers, Elbella scylla and Phocides
palemon lilea—but neither of them has a yellow outer
margin and purplish-red costal margin on the ventral
hindwing (Fig. 12).
In the ACG, the caterpillars of M. ambigua (Fig.
12) are oligophagous on mature leaves of a subset of
the resident species of Flacourtiaceae (Table 1).
Casearia corymbosa Kunth, which is unambiguously
the foodplant for the great majority of the caterpillars,
is a common forest-edge treelet in secondary succes-
sion (both anthropogenic and naturally occurring).
Casearia sylvestris Sw., though an equally common
forest-understory treelet, is rarely used by M. ambigua
caterpillars. Casearia arguta Kunth is a somewhat
scarcer secondary successional treelet that likewise is
rarely used. Zuelania guidonia Britton & Millsp.,
————
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
. 14-16. Last instars, plus adults in dorsal (L) and ventral (R) view, and pupa of ACG pyrrhopygines. 14, Myscelus belti (rainforest) 95-
SRNP-8435, L 2 99-SRNP-12882, R 2 99-SRNP-12671. 15, Passova gellias (rainforest to cloud forest) 95-SRNP-8758, L 2 99-SRNP-8542, Rd
99-SRNP-4972. 16, Pyrrhopyge crida (rainforest) 00-Cali-013.
VOLUME 55, NUMBER |
Fics. 17-19. Pupae of ACG pyrrhopygines. 17, Pyrrhopyge aesculapus (cloud forest) 97-SRNP-513. 18, Pyrrhopyge cosyra (rainforest to
cloud forest) 95-SRNP-539. 19, Pyrrhopyge erythrosticta (rainforest) 00-SRNP-15
34
though a common tree, is not commonly used (and
caterpillars occur only on the foliage of saplings, 1-3 m
from the ground). The other four species of ACG dry
forest Flacourtiaceae—Casearia tremula Griseb. and
C. praecox Griseb. (both locally common, but with
very small distributions) and Xylosma horrida Rose
and X. flexuosa (Kunth) Hemsl.—have never yielded
caterpillars of M. ambigua; nor have any of the other
10+ species of flacourtiaceous treelets and shrubs in
the wetter portions of the ACG, despite intensive
search.
At Santarém, Para, Brazil, Moss (1949) found cater-
pillars (described as mauve, belted with lemon-yellow)
of the M. barcastus species complex on small bushes of
Casearia.
The yellow-ringed caterpillar of M. ambigua is su-
perficially similar to at least 13 other ACG dry forest
hesperiid caterpillars: Elbella scylla plus the 12
pyrgine species listed in our account of that pyrrhopy-
gine. The caterpillar of M. ambigua (Fig. 12) has twice
as many rings as does that of E. scylla (Fig. 8), and
they are narrower.
Parasitization frequency of M. ambigua caterpillars
is very low (Table 1). Mysoria ambiguca is used by three
generalist tachinids—Hyphantrophaga virilis (Aldrich
& Webber), Chaetogena scutellaris (Wulp), and Lespe-
sia aletiae (Riley)—as well as by two still unidentified
species of Winthemia, both of which will probably turn
out to be essentially specialists on this pyrrhopygine.
The only two cases of parasitization by Hymenoptera
(microgastrine braconids) appear to be by species that
attack a number of skipper species.
Myscelus amystis hages Godman & Salvin
This taxon ranges from about the Tropic of Cancer
in Mexico to Panama (and probably also Colombia). In
the ACG dry forest, and on the dry side of the inter-
grades between dry forest and wetter forests to the
east, adults of Myscelus amystis hages are very rarely
seen visiting flowers such as Trigonia rugosa (Trigoni-
aceae) and Cordia spp. (Boraginaceae). Neither adults
nor caterpillars have ever been found in ACG rainfor-
est or cloud forest.
Myscelus amystis hages, which looks like no other
skipper in the ACG dry forest, most resembles its rust-
red congeners M. belti Godman & Salvin (Fig. 14) and
M. perissodora Dyar of the ACG rainforest. Owing to
the pronounced sexual dimorphism in color pattern in
M. a. hages, which is not expressed in the other two
species, the females of M. a. hages (Fig. 13) approach
those species in general appearance, while the more
yellow-orange males (Fig. 13) do not.
In the ACG, the caterpillars of M. a. hages are ap-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
parently oligophagous on mature leaves of three
species of Trichilia (Meliaceae) and nothing else
(Table 1). (Trichilia is rarely eaten by other caterpillars
of any kind.) Trichilia americana (Sessé & Moc.) T. D.
Penn., the heavily favored foodplant, is a common for-
est-edge medium-sized tree in all kinds of secondary
succession; the caterpillars (and newly laid eggs) are
most often encountered on branches with fully inso-
lated mature leaves. The other four species of ACG
dry forest Meliaceae—Trichilia martiana C. DC., Swi-
etenia macrophylla King, Cedrela odorata L., and
Guarea excelsa H. B. & K.—have never produced
these caterpillars; and neither have any of the other
five or so species of meliaceous trees in the wetter por-
tions of the ACG, despite intensive search.
The pink/dull green, shaggy caterpillar of M. a.
hages (Fig. 13) does not resemble any other species of
ACG dry-forest hesperiid caterpillar. But it strongly
resembles the dull green, shaggy caterpillars of M. belti
(Fig. 14) and M. perissodora and the pink/dull green,
shaggy caterpillar of Passova gellias (Fig. 15), all found
in the wetter forests on the eastern side of the ACG. It
can easily be distinguished from M. belti and P. gellias
by the reddish long hairs on its head (which, itself, be-
comes red posteriorly instead of remaining black).
Myscelus belti Godman & Salvin
Although Myscelus belti, like M. a. hages, ranges
from Mexico to Panama (and perhaps also Venezuela),
M. belti is a rainforest skipper instead of a dry-forest
one. While the caterpillars of M. belti are found in the
ACG rainforest and in intergrades with dry forest from
400-700 m, no caterpillars have ever been found in
dry forest or cloud forest (and free-flying adults have
not been seen or collected anywhere).
Myscelus belti, with its brick-red dorsal surface and
bright yellow central splotch ventrally (Fig. 14), can be
confused with only one ACG rainforest skipper—M.
perissodora. However, M. belti has a small hyaline spot
in the cell of the hindwing and, ventrally, a small black
spot on the hindwing costa at the angle near its base,
both of which M. perissodora lacks. Moreover, the
three hyaline spots (in spaces 1b, 2, and the cell) that
form the innermost band on the forewing are less well
aligned and less even along their proximal margin in
M. belti than they are in M. perissodora. The dark, dif-
fuse discal band of the dorsal hindwing is a little less
continuous, but at the same time a little darker in
spaces 5 and 6, in the former than in the latter. Finally,
the dorsal ground color of both wings is a bit redder in
M. belti than it is in M. perissodora.
In the ACG, the caterpillars of M. belti are
oligophagous on mature leaves of three species of
VOLUME 55, NUMBER 1
Guarea (Meliaceae)—G. glabra Vahl, G. rhopalocarpa
Radlk., and G. bullata Radlk.—the first of which is also
eaten by both M. perissodora and Passova gellias, and
the second, by P. gellias (Table 1). Guarea glabra,
which is certainly the principal foodplant, is a common
forest understory and edge treelet in all sorts of sec-
ondary succession. Our efforts notwithstanding, M.
belti caterpillars have never been found feeding on any
of some 15 other species of ACG Meliaceae: Trichilia
spp., Swietenia macrophylla, Cedrela odorata, Guarea
spp., and various other rainforest Meliaceae.
The dull green, shaggy caterpillar of M. belti (Fig.
14) does not resemble any species of ACG dry forest
hesperiid caterpillar except the pink/dull green, shaggy
caterpillar of M. a. hages (Fig. 13) which, however, has
reddish hairs on its head. But in the ACG rainforest
and cloud forest, it strongly resembles the pink/dull
green, shaggy caterpillar of Passova gellias (Fig. 15),
with which it is easily confused because they feed on
the same plants in the same place, and because P. gel-
lias varies from pink (usual) to dull green. We know it
must also closely resemble the caterpillar of M. peris-
sodora (of which we have no photo) because our only
reared adult of that species was mistaken, as a cater-
pillar, for M. belti.
In Parad, Brazil, Moss (1949) reared three species
that soon (Evans 1951) came to be called Myscelus
pardalina guarea Evans, Passova gazera (Hewitson),
and Passova passova gortyna (Hewitson), all on
Guarea trichilioides L.; in addition, he found nests of
the Myscelus on a second species of Guarea. Moss
(1949:34) noted not only that the larvae of these three
skipper species “are scarcely distinguishable” from one
another but also that they are very distinct from those
of other pyrrhopygines he had reared because “they do
not possess belts or patches of lighter colour. The gen-
eral tone is olive brown to greenish, ventrally pink with
pinkish extremities and the head is warm brown... .”
Although Evans (1951) viewed Myscelus and his new
genus Passova as separate genera, they are closely re-
lated. For those two genera, Evans (1951:72, 79) ex-
plicitly stated that “All males have dense brushes on
the inside of the femora and tibiae of the hind legs”;
and he had already pointed out (Evans 1951:5) that
this is “the only secondary sexual character in the sub-
family” Pyrrhopyginae. Close similarities in the male
genitalia of Myscelus and Passova are evident in plates
8 and 9 of Evans (1951).
Myscelus perissodora Dyar, reinstated status
Evans (1951)—whose material (except from the
Colombian department of Cauca) was both limited
and variable—treated Myscelus pegasus as a polytypic
species comprising a pair of subspecies: Myscelus p.
perissodora ranging from Mexico to Colombia and
Myscelus p. pegasus Mabille occurring in Venezuela,
French Guiana, and Ecuador. Although we lacked
males of the latter, we had enough females of both taxa
to allow critical comparison, which led to the conclu-
sion that they represent different species. We studied
5 2 of M. perissodora (1 from Mexico [holotype,
USNM], 2 Costa Rica, 2 Panama) and 4 2 of M. pega-
sus (Venezuela). Evans (1951) saw even fewer females:
4 2 of the former (1 Costa Rica, 1 Panama, 2 Colom-
bia) and 2 2 of the latter (1 Venezuela, 1 Ecuador).
Comparison of KOH-dissected genitalia of the holo-
type, 3 other ° of M. perissodora, and 2 ° of M. pega-
sus showed nothing but slight individual variation on a
single theme. However, absence of genitalic differen-
tiation in a group whose genitalia are generally conser-
vative does not argue strongly for conspecificity.
What is more important here is that species of
Myscelus are also conservative in superficial appear-
ance and that a number of independent superficial fea-
tures distinguish our females of the taxa in question.
The three large, white hyaline spots (in spaces Ib, 2,
and the cell) that form the innermost band on the
forewing are much smaller in M. perissodora than they
are in M. pegasus and, moreover, they are broadly,
rather than narrowly, edged in black. With respect to
the four contiguous, subapical hyaline spots on the
forewing, which run from the upper part of space 5
through space 8, the spot in space 6 is half again to
twice as high as those in spaces 7 and 8 in M. peris-
sodora, but only slightly higher than those in spaces 7
and 8 in M. pegasus; the subapical spots are not as
elongate in M. perissodora as they are in M. pegasus;
and the spot in the top of space 5 is only a point in M.
perissodora but a short dash in M. pegasus. The dark,
diffuse discal band of the dorsal hindwing is offset dis-
tad at vein 4 more conspicuously in M. pegasus than it
is in M. perissodora. The lobe in the outer margin of
the hindwing at the end of vein 4 is a little more pro-
nounced in M. pegasus than it is in M. perissodora.
The rusty-red ground color of the wings dorsally looks
slightly paler in M. pegasus than it does in M. peris-
sodora. The dark central band in the proximal yellow
area of the ventral hindwing extends to vein 7 in M. pe-
gasus but limits itself to a dash in space Ic in M. peris-
sodora. The dark discal band (which extends at least to
vein 6 or, more often, to vein 7, with sometimes a de-
tached spot in space 7) is separated from the dark dis-
tal half of the ventral hindwing by a strip of yellow in
M. perissodora (except in the type where just traces of
JOURNAL OF THE LEPID ERISTS’ SOCIETY
13509.
VOLUME 55, NUMBER 1
ines. 23, Myscelus amystis hages (dry forest) 93-SRNP 7, 24, 25, Passova gellias (rainforest to
Fics. 23-25. Pupae of ACG pyrrhor
cloud forest) 94-SRNP-7815.
fo}
38
yellow remain) but is fused with the dark distal half in
M. pegasus.
Because the caterpillar that produced our lone adult
of M. perissodora was identified in the field by its
finder as M. belti, the larvae of these two skippers must
look about the same. The look-alike M. perissodora
caterpillar was likewise feeding on Guarea glabra, the
usual foodplant of M. belti. However, the caterpillar
came from the upper Rio Mena (700 m) in the inter-
grade between dry forest and rainforest on the north
slope of Volcén Orosi. No examples of M. belti have
been reared from there, although the habitat looks
similar to that on the southeast side of the Volcan
Orosi-Volcén Cacao complex where caterpillars of M.
belti have commonly been found.
Passova gellias (Godman & Salvin)
Passova gellias (Fig. 15) is definitely known from
Honduras, Costa Rica, and Panama. While its caterpil-
lars are found in the ACG rainforest, cloud forest, and
intergrades with dry forest from 400-700 m, they have
not been found in the dry forest proper; and no free-
flying adults have been seen or collected in the ACG.
In the wetter forests of the ACG, the caterpillars of
P. gellias (Fig. 15) are oligophagous on mature leaves
of two of the three species of Guarea (Meliaceae) that
Myscelus belti eats (one of which is also eaten by M.
perissodora); and G. glabra is almost always the food-
plant of choice (Table 1). See the account of M. belti
for (1) a list of some 15 other ACG rainforest species
of Meliaceae that, despite repeated search, have never
yielded caterpillars of M. belti, M. perissodora, or P.
gellias and (2) comments on basic similarities not only
in the foodplants and the caterpillars but also in cer-
tain critical morphological features of the adults of the
genera Myscelus and Passova.
THE ONLY PYRRHOPYGINE THAT ENTERS THE USA
Pyrrhopyge arizonae Godman & Salvin,
reinstated status
Because this taxon has been considered a northern
subspecies of the Mexican Pyrrhopyge araxes (Hewit-
son), it has generally been called either Pyrrhopyge
araxes arizonae (also, for a time, Apyrrothrix araxes
arizonae) or—by those who have chosen to downplay
subspecies—simply P. araxes. Godman and Salvin’s
(1893:253) original description of P. arizonae was both
minimal (because this Arizonan skipper was outside
the fauna that they were treating) and incidental to
their account of P. araxes (which belonged to that
fauna):
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Hewitson’s description of this species [P. araxes] was based upon
a Mexican specimen now in the British Museum. Our Mexican se-
ries of examples agrees generally with the type, though there is a
considerable amount of variation as regards the definition of the
markings of the underside of the secondaries. In Arizona specimens
these marks are evanescent, and the dark outer border is not clearly
defined on its inner edge. The difference is sufficient to constitute a
distinct race. Plétz also recognized two forms, but unfortunately
gave a new name, cyrillus, to the true araxes, the Arizona form be-
ing his araxes. To rectify this we propose to call the northern insect
Pyrrhopyge arizonae, specimens of both sexes being in our collec-
tion, sent us by Morrison from the neighbourhood of Fort Grant
[which almost certainly means from the Pinalefio Mountains].
It is worth noting that, even though they wrote
“Pyrrhopyge arizonae,’ Godman and Salvin used the
term “race” rather than “species” in referring to it. Ap-
parently, they themselves saw this taxon as equivalent
to a subspecies.
But P. arizonae and P. araxes are separate (albeit ex-
tremely close) species. Superficially—besides the pair
of differences noted by Godman and Salvin—the
brown ground color of P. arizonae is not nearly so dark
as that of P. araxes; the white wing fringes of P. ari-
zonae are more prominent and more heavily check-
ered than those of its sister; and, ventrally, the proxi-
mal yellow of the (mainly hind-) wings is not as
intense, not as orangy, in P. arizonae as in P. araxes. On
an average, P. arizonae is a little smaller than P. araxes.
Genitalically, in males, there are several subtle dif-
ferences which are not evident when comparing pub-
lished genitalia figures of P. araxes (Godman & Salvin
1893:pl. 73, fig. 16, Bell 1931:fig. 51, Evans 1951:fig.
A.1.58) and P. arizonae (Skinner & Williams 1922:fig.
1, reprinted in Lindsey et al. 1931:pl. 2, fig. 1). These
differences are described here in strict lateral view.
Immediately anterior to the uncus, at the posterodor-
solateral corners of the tegumen (where in many
pyrrhopygines paired, long, caudally-projecting prongs
originate), there are very short, dorsally-projecting
lobes in P. araxes but not in P. arizonae. The proximal
(massive) portion of the valva is longer in P araxes
than it is in P arizonae; and its dorsal edge, as it
changes (anterior to posterior) from roughly horizontal
to nearly vertical, is somewhat angled in P. araxes but
more evenly curved in P. arizonae. Although the den-
tate, distal projections of the valvae are variable (and
slightly asymmetric) in both species, the ventral ones
are wider in P. araxes than they are in P. arizonae. In
the female genitalia of both species, the posteriormost
end of the ductus bursae is closely surrounded by
heavy sclerotization that forms a long, caudally-ex-
tending tube which brings the ostium bursae far back
beneath the proximal ends of the ovipositor lobes.
Originating from the posterior end of the sclerotized
VOLUME 55, NUMBER 1
39
Fics. 26-28. Last instar and pupa of Pyrrhopyge arizonae (oak eater) in summer 1959; collected as larvae along the Swift Trail, 1615 m,
Pinalefio Mountains, Graham County, Arizona, USA, on 5 June 1959. 26, Larva in left lateral view. 27, Prothoracic segment (enlarged) in dor-
sal view. 28, Pupa in left lateral view.
tube (i.e., at the ostium bursae), a midventral, long,
narrow, delicate, terminally-pointed prong and a mid-
dorsal, equally long, but wide (and progressively
widening) projection, which, in its distal half, is mid-
dorsally divided so as to suggest a pair of terminally-
blunt paddles, all extend back to the level of the distal
ends of the ovipositor lobes. Compared genitalic dis-
sections include 7 ¢ 4 ° of P. arizonae from the Chiric-
ahua and Huachuca mountains, Cochise Co., plus
Pena Blanca Lake, Santa Cruz Co., Arizona, USA, and
30 mi (48 km) N of Mante, Tamaulipas, Mexico; and 4
3 3 2 of P. araxes from Mexico City, Cuernavaca, and
Tizapan, Mexico.
Pyrrhopyge arizonae occurs in the southwestern
USA (in the montane oak belt of southeastern Arizona,
adjacent southwestern New Mexico, and also the Big
Bend region of Texas) and in northern Mexico (from
Sonora, Chihuahua, and Tamaulipas southward to un-
certain limits). Pyrrhopyge araxes occurs in central
and southern Mexico (at least from Jalisco through the
Distrito Federal and Morelos to Veracruz, south to at
least Guerrero and Oaxaca).
In southeastern Arizona (Cochise and Graham
counties), Comstock (1956) collected a single caterpil-
lar of P. arizonae on oak at 1860 m in Miller Canyon,
Huachuca Mountains, on 27 July 1955; and Burns
(1964:109, unpubl.) collected three caterpillars on
Quercus arizonica Sarg. and another on Quercus sp. at
1615 m in the South Fork of Cave Creek Canyon,
Chiricahua Mountains, on 22, 23, 24 April 1959, plus
two, on Q. arizonica and Q. emoryi Torr., respectively,
at 1615 m along the Swift Trail, Pinalefio Mountains,
on 5 June 1959 (Quercus is in the Fagaceae). After fur-
ther growth, these last two specimens posed for Figs.
26 to 28. This skipper has also been reported on Q. ob-
longifolia Torr. (Bailowitz & Brock 1991).
Although the adult of P. arizonae is a classic “skip-
per-brown’—with the “usual” hyaline spots on the
forewing—and is in no way a flashy pyrrhopygine, the
caterpillar (Fig. 26) ranks with the best of them. Its
head is black (clypeus tan), densely clothed with (for
the most part, relatively short) white hairs—paired
ventrolateral patches of which are rusty-orange; and its
body is a rich, deep maroon crossed by brilliant yellow
bands (one per segment), with long white hairs spring-
ing mainly from the yellow bands, but also from a pale,
creamy-yellow prothoracic shield. This pale shield
bears a very thin, black band dorsolaterally (but not
meme aaaaaaacaaaaeaamaaaamaacaaaaaaaaaaaaa
40
medially) about two-thirds of the way from its anterior
to its posterior margin (Fig. 25).
The pupa (Fig. 26) j is dark and heavily clothed with
hairs, which are mostly white, but brilliant orange-
yellow on the prothorax. The wing covers, the thorax
dorsally, plus the proboscis, legs, antennae, the greater
part of the eyes, and a few other head plates are all a
dark (purplish) black with a whitish blue-gray bloom.
The thoracic spiracles, part of the eyes, the anterior
head region, and the paired plates anterior to the pro-
thorax are all a rich chestnut-brown. The abdomen
looks dirty yellowish-brown, with alternating yellower
and browner zones, the latter coated with blue-gray
bloom. The terminal segment is mostly brown and
blackish-brown, and the cremaster is brown. Although
descriptions of the last instar and pupa provided by
Comstock (1956) variously supplement, support, and
depart from those above, both his and our pupal de-
scriptions clearly fit the subfamily mold (Figs. 16-25).
DISCUSSION
Ecology. Twenty-two years of rearing 2192
pyrrhopygine caterpillars and collecting far fewer
adults show that the ACG supports at least 15 species
of pyrrhopygine hesperiids: 3 in (lowest elevation) dry
forest (Elbella scylla, Mysoria ambigua, and Myscelus
amystis hages); 4 in (highest elevation) cloud forest
(Pyrrhopyge creon, P. aesculapus, P. cosyra, and
Passova gellias); and 10 in (middle elevation) rainfor-
est (Pyrrhopyge zenodorus, P. crida, P. cosyra, P. ery-
throsticta, Parelbella macleannani, Jemadia pseudo-
gnetus, Jemadia sp. X (an unnamed Jemadia belonging
to a J. hewitsonii species complex), Myscelus belti, M.
perissodora, and Passova gellias). Although caterpillars
of some of these species are occasionally found in the
intergrades between their “usual” ecosystem and an
adjacent one, no pyrrhopygine species spans all three
ecosystems; and none occurs in both dry forest and
rainforest. Only two range from rainforest to cloud for-
est: Pyrrhopyge cosyra, which occurs from hot low-
land rainforest to high, cold cloud forest; and Passova
gellias, which generally goes less far up into the cloud
forest.
ACG pyrrhopygines seem not to migrate seasonally
between major ecosystems. Even in ne dry season,
sexually dormant adults of the three dry-forest species
appear in local moist areas of the dry forest rather than
in the adjacent rainforest. (Note that these adults do
not break reproductive dormancy and oviposit on their
respective foodplants when those plants—responding
to, say, the clearing of right-of-ways and firebreaks—
produce new, leafy, sucker shoots from fresh stumps in
the middle of the dry season.)
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Ecosystem-level specificity of ACG pyrrhopygines
does not stem entirely from the ecology of their food-
plants. On the one hand, dry-forest pyrrhopygine
foodplants do not occur in rainforest and cloud forest
(although congeners do). On the other hand, some
wet-forest pyrrhopygine foodplants do occur in dry
forest as well (along with congeners), while the
pyrrhopygines that eat them do not. For example, Vis-
mia baccifera and Clusia rosea occur in both ACG dry
forest and rainforest; but Pyrrhopyge zenodorus and P.
cosyra, their respective grazers, stick to the wetter for-
est. However, many other ACG skipper species do oc-
cur in both dry forest and rainforest. The plant families
Lauraceae, Flacourtiaceae, Myrtaceae, Combretaceae,
and Malpighiaceae—which are used by ecosystem-
specialist pyrrhopygines—all contain species that oc-
cur in both dry forest and rainforest where they sup-
port larval pyrgines that likewise range across both
ecosystems.
Costa Rica has about 25 species of pyrrhopygines, or
about one-eighth of our estimated total of all
pyrrhopygine species. This estimate of 200 may still be
too low, but it far exceeds the number (“almost 150”)
offered by Ackery et al. (1999). Since we now have
reared 15 species (and anticipate only one to three
more), the ACG is home to at least 60% of the Costa
Rican pyrrhopygine fauna. Though an ecological is-
land, the ACG seems to offer enough habitat, with
abundant foodplants, to sustain these skippers. How-
ever, substantial global warming, with a concomitant
rise in the elevation of the cloud layer (e.g., Pounds et
al. 1999), could seriously reduce the area of cloud-for-
est habitat for Pyrrhopyge creon and P. aesculapus.
Moreover, the gradual return of large areas of ACG
secondary successional forest to old-growth forest over
the coming centuries will substantially reduce the sizes
and ubiquity of populations of important rainforest
foodplants such as Vismia baccifera and dry-forest
foodplants such as Byrsonima crassifolia and Casearia
corymbosa.
Pyrrhopygine caterpillars, like those of other ACG
skippers that feed on broadleaf plants, are concen-
trated primarily in the zone between 10 cm and 3 m
above the ground, even when individuals of their food-
plants rise as much as 20-40 m. Although further in-
ventory exploration of the ACG canopy may turn up
exceptions, they are not yet evident. While adult skip-
pers can be seen visiting flowers in the crowns of the
tallest trees, they do not seem to be ovipositing at
these heights.
Foodplants. ACG pyrrhopygine larvae have exclu-
sive tastes. The caterpillars of Myscelus belti, M. peris-
sodora, and Passova gellias are the only ones that share
VOLUME 55, NUMBER 1
their foodplants with other pyrrhopygines—in these
cases, each other (Table 1). The caterpillars of Elbella
scylla, Parelbella macleannani, and Jemadia pseudo-
gnetus are the only ones that share their foodplants
with non-pyrrhopygine hesperiid larvae.
The strong host specificity shown by pyrrhopygine
caterpillars is frequent among ACG butterflies and
certain families of moths (e.g., host records in Janzen
& Hallwachs 2001), and especially among dicot-eating
skippers. Even though the foodplant records for our
pyrrhopygines (Table 1) are dotted across the botanical
taxonomic landscape (i.e., in the families Lauraceae,
Clusiaceae, Flacourtiaceae, Cunoniaceae, Myrtaceae,
Combretaceae, Meliaceae, Malpighiaceae, and Arali-
aceae), some patterns emerge. M yscelus amystis
hages, M. belti, M. perissodora, and Passova gellias,
which (aside from their adult color patterns) are simi-
lar, all focus on Meliaceae—mainly Guarea glabra.
Several species currently in the large genus Pyrrho-
pyge (P. zenodorus, P. Gide P. cosyra, and P. erythro-
sticta) are monophagous to oligophagous on members
of the Clusiaceae. A female of another Costa Rican
species of Pyrrhopyge, P. haemon Godman & Salvin,
was found and reared (in 1995 at 1600 m in Tapanti
National Park by R. Delgado) on a clusiaceous plant in
the genus Vismia, species of which are the foodplants
of P. zenodorus and P. crida.
In this connection, it is also clear that ACG
pyrrhopygine caterpillars are specialists at levels far
below that of the plant family. For each species of
caterpillar, there are many ACG species of plants
within the plant family fed upon that are not used. It is
particularly striking, for example, that Pyrrhopyge
zenodorus eats the common Vismia baccifera (occur-
ring from dry forest to rainforest) and the rare V. fer-
ruginea, while P. crida eats neither, restricting its diet
to the low density and local rainforest V. billbergiana.
Within an oligophagous species, percentages of
caterpillar records among its small set of foodplants
(Table 1) do not reflect the relative abundances of
those plants. For example, Mysoria ambigua uses
Casearia corymbosa most of the time, even though its
other three foodplants are common in the same dry-
forest habitats.
More or less close relatives of half of the pyrrhopy-
gine species that we have reared in the ACG were
reared previously in Para, Brazil, by Moss (1949). The
plants he found them eating are usually in the same
genera as the foodplants of their Costa Rican counter-
parts, and are always in the same families. Not surpris-
ingly, the strong host specificity shown by ACG
pyrrhopygine caterpillars seems to be geographically
conservative.
4]
In also rearing a few pyrrhopygines most of which
were well removed from the species we have reared,
Moss (1949) recorded foodplants in some other fami-
lies: Annonaceae, Myristicaceae, Sterculiaceae, and
Sapindaceae (although he questioned the accuracy of
the identification of the last). In revising both Elbella
and its near relatives (most of whose diets remain un-
known), Mielke (1995) added one foodplant genus
(Quiina and Miconia) from each of the flees Qui-
inaceae and Melastomataceae. While the total taxo-
nomic spread of pyrrhopygine foodplant families be-
comes a little wider, there is also a little clustering. The
Annonaceae may relate to the Myristicaceae; the Qui-
inaceae, to the Clusiaceae; and the Melastomataceae,
to the Combretaceae or Myrtaceae (Heywood 1993).
The only pyrrhopygine to enter the USA, Pyrrho-
pyge arizonae, puts another isolated genus and family
(Quercus, Fagaceae) on the list of disparate food-
plants.
Biogeography. To judge from current distribu-
tions, the Pyrrhopyginae (a strictly New World group)
probably arose in tropical South America. Most of the
existing species are there. Nearly half of the ACG
pyrrhopygines we have reared (Pyrrhopyge zenodorus,
Elbella_ scylla, Parelbella macleannani, Jemadia
pseudognetus, Jemadia sp. X, Mysoria ambigua, and
Myscelus amystis hages) appear to be northern exten-
sions of species complexes that are better represented
and more widely distributed in South America. Four
more (Pyrrhopyge crida, P. aesculapus, P. cosyra, and
Myscelus perissodora) are species with significant South
American ranges (usually Colombia and Ecuador) as
well as Middle American ones. Only four (Pyrrhopyge
creon, P. erythrosticta, Myscelus belti, and Passova gel-
lias) seem to be confined to Middle America. There are
no pyrrhopygines in the West Indies.
Parasitoids. ACG pyrrhopygines are notably free of
parasitoids. Despite their slow larval growth rates
(among the longest within the ACG _hesperiids),
pyrrhopygines are at the low end of the spread of lar-
val parasitization frequencies—and not just among
skippers but among ACG macrolepidoptera gener:
Only 7.1% of the total wild-caught pyrrhopygine cater-
pillars were parasitized. Ten oF the 15 species reared
from wild-caught caterpillars produced no parasitoids
whatsoever, even when samples were large (Table 1).
It is especially noteworthy that none a our many
caterpillars (n = 295) belonging to the closely related
genera Myscelus and Passova has yielded parasitoids
(Table 1).
Ichneumonids have been reared only from
Pyrrhopyge cosyra caterpillars. However, Casinaria
sp. 9, which frequently attacks P. cosyra (31 records),
also attacks the pyrgines Phocides nigrescens (2
records) as well as Achlyodes thraso (Hiibner) (1
record), A. selva Evans (2 records), and A. busirus
heros (6 records) in the same habitat (but has not yet
been found to attack tens of other hesperiid species in
this habitat). To date, the ichneumonid genus
“MCAM"” sp. 1 (10 records) has been found parasitiz-
ing only Pyrrhopyge cosyra. Only Elbella scylla is at-
tacked by a braconid with enough frequency to suggest
that the relationship might be host-specific. Parasitiza-
tion by tachinid flies is light, and only four of the 15
species reared from wild-caught caterpillars had any at
all. Whether the slow growth rate of pyrrhopygine
caterpillars is evolutionarily “permitted” by the low
parasitization frequency, or the low parasitization fre-
quency stems from traits of the caterpillars, will be the
subject of family-wide analysis at a later time.
Taxonomy. It is hardly surprising that most of the
parasitoids attacking ACG pyrrhopygines are unde-
scribed whereas their large and ostentatious hosts are
not. Nevertheless, when it came to identifying these
Costa Rican skipper butterflies (which led to examin-
ing them in a much broader geographic and taxonomic
context), fully half of them posed biological questions.
The recurring one involved the status of an ACG en-
tity vis-a-vis its nearest relative: Are those perceptible
(but closely similar) differentiates species or sub-
species? In general, taxa that Evans (1951) treated as
subspecies are here granted species rank. Other bio-
logical problems bearing on accurate application of
names involved (1) something currently passing as a
single species that actually comprises a pair of sym-
patric species, (2) the reverse of this—a supposed pair
of sympatric species that actually are one and the same
thing, and (3) an entity that belongs to an essentially
South American species complex not previously
recorded from Costa Rica. Pyrrhopygines are a flashy
tip of a taxonomic iceberg. If some of their identities
were this uncertain, what about those of all the rest of
the ACG skipper butterflies (which reflect the world’s
uniquely rich neotropical hesperiid fauna)? And then,
by extension, what about the placement of all the rest
of ACG invertebrate biodiversity? Taxonomists and
time are of the essence.
ACKNOWLEDGMENTS
We thank people and institutions for help as follows: Loaning
specimens, P. R. Ackery, The Natural History Museum (BMNH),
London, England; and Jacqueline Miller, Allyn Museum of Ento-
mology/Florida State Museum, Sarasota, Florida, USA. KOH-
dissecting many skipper genitalia, Patricia Gentili, Donald Harvey,
and Elizabeth Klafter. Translating a few paragraphs, Astrid Caldas
and Gentili. Formatting tables, Gentili. Drawing Figs. 26 to 28, E. P.
Catts. Digitally composing them, James DiLoreto and John Clarke.
Discussing problems, John Brown, Jason Hall, Harvey, Gerardo
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Lamas, Robert Robbins, and Chris Thompson. Reviewing the man-
uscript, Deane Bowers, Brown, and an anonymous reviewer. Read-
ing proofs, Sarah Burns. Finding and rearing caterpillars, Winnie
Hallwachs, Guillermo Pereira, Lucia Rios, Manuel Pereira, Elieth
Cantillano, Osbaldo Espinosa, Ruth Franco, Harry Ramirez, Gloria
Sihezar, Roberto Espinosa, Roster Moraga, Felipe Chavarrfa, Ale-
jandro Masis, Adrién Guadamuz, Mariano Pereira, Jon Sullivan,
Eric Olson, Jessica DiMauro, Roger Blanco, Daniel Perez, Benigno
Guadamuz, Carolina Cano, Freddy Quesada, Dunia Garcia, Calixto
Moraga, and Petrona Rios. Identifying plants, Nelson Zamora,
Roberto Espinosa, Barry Hammel, and Maria Marta Chavarria.
Identifying parasitoids, lan Gauld, Monty Wood, Jim Whitfield, and
Mike Sharkey. Supporting this research, the National Science Foun-
dation ( (DHJ: BSR 80-11558, BSR 83-07887, BSR 86-10149, BSR
90-24770, DEB 93-06296, DEB 94-00829, DEB 97-05072, DEB
00-72730), Costa Rica’s INBio, Costa Rica’s Ministerio del Ambiente
y Energia (MINAE), the Area de Conservacion Guanacaste, Earth-
watch, US-AID, US-NIH, and Asociaci6n BioGuanacaste.
LITERATURE CITED
ACKERY, P. R., R. DE JONG & R. I. VANE-WricHT. 1999. The but-
terflies: Hedyloidea, Hesperioidea and Papilionoidea, pp.
263-300. In Kristensen, N. P. (ed.), Handbook of zoology. Vol.
IV. Arthropoda: Insecta, part 35, Lepidoptera, moths and but-
terflies. Vol. 1. Evolution, systematics, and biogeography. Wal-
ter de Gruyter, Berlin & New York.
AusTIN, G. T. & O. H. H. MIELKE. 2000. Hesperiidae of Rondénia,
Brazil: Cephise Evans (Pyrginae), with descriptions of new
species from Mexico and Brazil. Revta bras. Zool. 17:757-788.
BaiLowiTz, R. A. & J. P. BRocK. 1991. Butterflies of southeastern
Arizona. Sonoran Arthropod Studies, Inc., Tucson, Arizona. ix +
342 pp.
BELL, E. L. 1931. Studies in the Pyrrhopyginae, with descriptions
of several new species (Lepidoptera, Rhopalocera, Hesperi-
idae). J. New York Entomol. Soc. 39:417-491.
. 1933. Studies in the Pyrrhopyginae, with descriptions of
new species (Lepidoptera, Rhopalocera, Hesperiidae). J. New
York Entomol. Soc. 41:265-295, 481-529.
. 1934. New American Hesperiidae (Lepidoptera, Rhopalo-
cera). Bull. Brooklyn Entomol. Soc. 29:89-96.
Burns, J. M. 1964. Evolution in skipper butterflies of the genus
Erynnis. Univ. Calif. Publ. Entomol. 37:1-217.
. 1996. Genitalia and the proper genus: Codatractus gets
mysie and uvydixa— a compact cyda group—as well as a hys-
terectomy, while Cephise gets part of Polythrix (Hesperiidae:
Pyrginae). J. Lepid. Soc. 50:173-216.
Burns, J. M. & D. H. JANZEN. [2000] “1999.” Drephalys: Division
of this showy neotropical genus, plus a new species and the im-
matures and food plants of two species from Costa Rican dry
forest (Hesperiidae: Pyrginae). J. Lepid. Soc. 53:77-89.
Comstock, J. A. 1956. Notes on the life histories of two southern
Arizona butterflies. Bull. South. Calif. Acad. Sci. 55:171-179.
Evans, W. H. 1949. A catalogue of the Hesperiidae from Europe,
Asia and Australia in the British Museum (Natural History).
British Museum, London. xix + 502 pp., 53 pls.
. 1951. A catalogue of the American Hesperiidae indicating
the classification and nomenclature adopted in the British Mu-
seum (Natural History). Part I. Introduction and Group A,
Pyrrhopyginae. British Museum, London. 92 pp., pls. 1-9.
FREEMAN, H. A. 1966. New Hesperiidae records for Mexico. J.
Lepid. Soc. 20:226-228.
. 1977. Six new species of Hesperiidae from Mexico. J.
Lepid. Soc. 31:89-99.
Gopman, F. D. & O. SALvIN. 1879. Descriptions of new species of
Rhopalocera from Central and South America. Proc. Zool. Soc.
London 1879(1):150-155, pl. 14.
. 1879-1901. Biologia Centrali-Americana; Insecta; Lepi-
doptera-Rhopalocera. Vol. 2, 782 pp. Vol. 3, 113 pls.
HAYWARD, K. J. 1948. Genera et species animalium argentinorum.
Vol. 1. Insecta., Lepidoptera, Hesperiidae, Pyrrhopyginae and
VOLUME 55, NUMBER 1
Pyrginae. Guillermo Kraft Ltda., Buenos Aries. 389 pp., 27 pls.
Heywoop, V. H. 1993. Flowering plants of the world. Oxford Univ.
Press, New York. 336 pp.
HOFFMANN, C. C. 1941. Catalogo sistematico y zoogeografico de
los Lepidopteros Mexicanos. Segunda parte. Hesperioidea. An.
Inst. Biol. Mex. 12:237-294.
JANZEN, D. H. 1988a. Guanacaste National Park: Tropical ecologi-
cal and biocultural restoration, pp. 143-192. In Cairns, J. J.
(ed.), Rehabilitating damaged ecosystems. Vol. II. CRC Press,
Boca Raton, Florida.
. 1988b. Ecological characterization of a Costa Rican dry
forest caterpillar fauna. Biotropica 20:120-135.
. 1993. Caterpillar seasonality in a Costa Rican dry forest,
pp. 448-477. In Stamp, N. E. & T. M. Casey (eds.), Caterpillars.
Ecological and evolutionary constraints on foraging. Chapman
& Hall, New York, New York.
. Inpress. Ecology of dry forest wildland insects in the Area
de Conservaci6n Guanacaste, northwestern Costa Rica. In
Frankie, G. W., A. Mata & S. B. Vinson (eds.), Biodiversity con-
servation in Costa Rica: learning the lessons in seasonal dry for-
est. Univ. Calif. Press, Berkeley.
JANZEN, D. H. & W. Haiwacus. 2001. Event-based database of
caterpillars, their host plants, and their parasitoids in the Area
de Conservacién Guanacaste, northwestern Costa Rica.
(http://janzen.sas.upenn.edu).
43
LinpsEy, A. W., E. L. BELL & R. C. WILLIAMS JR. 1931. The Hes-
perioidea of North America. Denison Univ. Bull., J. Sci. Lab.
26:1-142.
Mayr, E. 1942. Systematics and the origin of species. Columbia
Univ. Press, New York, New York. xiv + 334 pp:
MIELKE, O. H. H. 1995. Revisao de Elbella Evans e géneros afins
(Lepidoptera, Hesperiidae, Pyrrhopyginae). Revta. bras. Zool.
11:395-586.
Moss, A. M. 1949. Biological notes on some “Hesperiidae” of Para and
the Amazon (Lep. Rhop.). Acta Zool. Lilloana 7:27-79, pls. I-V.
Nico.ay, S. S. & G. B. SMALL Jr. 1969. A new subspecies of
Pyrrhopyge creon (Hesperiidae) from Panama. J. Lepid. Soc.
23:127-130.
. 1981. Illustrations and descriptions of some species of
Pyrrhopyginae from Costa Rica, Panama and Colombia (Hes-
periidae). J. Res. Lepid. 19:230-239.
Pounps, J. A., M. P. L. FOGDEN & J. H. CAMPBELL. 1999. Biologi-
cal response to climate change on a tropical mountain. Nature
398:611-615.
SKINNER, H. & R. C. WILLIAMS JR. 1922. On the male genitalia of
the larger Hesperiidae of North America. Trans. Am. Entomol.
Soc. 48:109-127.
Received for publication 27 November 2000; revised and accepted
12 July 2001.
BOOK REVIEWS
Journal of the Lepidopterists’ Society
55(1), 2001, 44-45
A CONTRIBUTION TO RIODINID SYSTEMATICS (LEPIDOPTERA: RIO-
DINIDAE), by Jason P. W. Hall with Keith R. Willmott & Donald J.
Harvey, Contributors. 1998. Tropical Lepidoptera. Volume 9, Sup-
plement 11. 42 pp., 7 color plates. Published by The Association for
Tropical Lepidoptera (www.troplep.org), P.O. Box 141210,
Gainesville, Florida, USA. ISSN 1048-8138, available from the pub-
lisher.
A REVISION OF THE GENUS THEOPE, ITS SYSTEMATICS AND BIOLOGY
(LEPIDOPTERA: RIODINIDAE: NYMPHIDIINI), by Jason P. W. Hall.
1999. Published by Flora & Fauna Books (www.ffbooks.org),
Gainesville. 127 pp., 9 color plates. Paperback, 8.5 x 11 in., glossy
paper, ISBN 0-945417-95-0, available from the publisher.
As an amateur rather than a professional entomologist, it is easy
to wax enthusiastic about these two publications. Those of us inter-
ested in riodinid butterflies have seen a steady increase in the quan-
tity and quality of publications regarding this group. Supplement 1
to Volume 9 of Tropical Lepidoptera consists of four papers au-
thored or co-authored by Jason P. W. Hall. The co-authored papers
describe a total of 13 new riodinid species from Ecuador. The longer
paper is a systematic revision of Sarota, including 5 species and 2
subspecies previously not described. A particularly helpful key is
provided which will prove invaluable for the worker trying to differ-
entiate these rather small butterflies.
The second publication, “A Revision of the Genus Theope,” car-
ries the process to an even higher level. For this 125 page tome, the
author has personally examined 4253 Theope specimens in 23 major
collections on three continents, including all but three of the extant
primary types. Most Theope are rare in collections (with a few no-
table exceptions), and half the described species are known from 20
or fewer specimens. Nevertheless the paper contains dorsal and ven-
tral photographs of all the recognized species with genitalia illustra-
tions of 70% of the females and all the males save Theope villai. Ter-
minal abdominal tergite drawings are also included, which show a
remarkable diversity of sizes and shapes. To avoid confusion by fu-
ture researchers, the author has extensively documented the mate-
rial examined and figured, including the location for each figured
specimen and genitalia dissection. Many lectotypes are also desig-
nated.
This revision identifies 68 Theope taxa. A binomial key for identi-
fication of males is provided, but not females, since many are un-
known, especially in the “foliorum group.” The author has included
corrected identifications for specimens figured by Seitz, Godman
and Salvin, Barcant, D’Abrera, DeVries, and others, updating all
these works to conform with this revision.
Most interesting is the discussion of Theope as an indicator of
species diversity in Upper Amazonian areas. The extensive range
maps are most helpful and the tables listing perching times for 34
species of Theope will provide a basis for correlation of further field
observations. I believe this publication could serve as a template for
any doctoral thesis in this field, much less any taxonomic revision. To
have this much valuable information, so concisely compiled, is an
unexpected pleasure.
DaviD AHRENHOLZ, 9255 County Road 6, Maple Plain, Min-
nesota 55359.
THE WESTERN PALEARCTIC ZYGAENIDAE (LEPIDOPTERA), by C. M.
Naumann, G. M. Tarmann and W. G. Tremewan, 1999. Published
by Apollo Books, Kirkeby Sand 19, DK-5771 Stenstrup, Denmark.
304 pp., 178 line drawings and black and white photographs, 12
color plates. Hardback, 24 x 17 cm, ISBN: 87-88757-15-3. Available
from Apollo Books, Price DK 6000.00 (about $80.00 US) excluding
postage, available from the publisher.
Three eminent specialists have combined their expertise to cre-
ate a book on one of the most striking groups of day-flying moths,
the Zygaenidae. In addition, they have compiled the results of many
studies by other workers, amateur or professional, over the last 100
years. The foreword by Miriam Rothschild, written in her typical en-
tertaining style, introduces the reader to this book in the context of her
own interests such as their extensive variation, complex diapause pat-
terns, behavior, conservation, and toxicity. The purpose of the book,
as explained in the authors’ foreword, is to provide an up-to-date
summary of past and current biochemical, physiological, behavioral,
and ecological studies, and to present an overview of the distribu-
tion, ecology, and systematics of the zygaenid moths of the western
Palaearctic. They have certainly succeeded in achieving these aims.
The book is divided in two parts. In the “General Part” the sys-
tematics and phylogeny of the Zygaenidae are briefly discussed, fol-
lowed by a description of life cycles and their peculiarities. The mor-
phology of the immature and imaginal stages, with frequent
references to their phylogenetic implications, is briefly, but compre-
hensively, described and illustrated in the section “Structures and
functions” that includes a discussion on “Senses and orientation” and
“Nutrition.” “Genetics and individual variation” deals with wing pat-
tems and coloration, including color mutants and other aberrations,
of some Zygaena species. In “Zoogeography” the geographical vari-
ation in Zygaena moths, hybrid belts, littoral and montane
melanism, and geographical variation in color and size of larvae is
discussed. The distribution patterns in the western Palaearctic are
assigned to a Mediterranean, Syro-Anatolian, Iranian, Transcau-
casian or Euro-Siberian type, which but for the latter, reflect and in-
clude Pleistocene refugia and survivors of the last glaciation. Fossil
zygaenids are briefly mentioned. “Ecology and behavior” deals ex-
tensively with their diverse habitat preferences. A discussion on the
food plants leads to a highly informative section on cyanogenensis,
cuticular secretion of defensive compounds by larvae and imagoes as
a defensive strategy. The strong attraction of adults to certain flow-
ers is ascribed to compounds related to sex attractants of some Zy-
gaena species. Two independent mate-locating systems, the early
morning (optical cue) strategy and the pheromone induced matings,
are described. The toxicity and aposematic coloration of zygaenids is
related to mimicry, behavior, and predation. Diapause in these in-
sects reveal complex patterns which regulate phases of growth and
dormancy. Parasites of larvae and pupae include wasps and tachinid
flies. In “Zygaenids as indicator species’ their importance as such is
emphasized. Conservation should concentrate on the protection of
habitats. “Breeding” and “Collecting techniques” is followed by an
“History of research on the Zygaenidae” highlighting the contribu-
tions made in taxonomy, collecting expeditions, biology, genetics,
biochemistry, ecology and behavior, reproductive biology, and bibli-
ography by past and present researchers and naturalists. The first
part ends with a list of vernacular names as used in the western
Palaearctic. “Selected references” introduces the reader to more de-
tails of zygaenid biology and systematics.
The “Systematic Part” starts with a check list of the 116 species
found in the western Palaearctic. This list is divided into the three
VOLUME 55, NUMBER 1
subfamilies, the Procridinae, Chalcosiinae and Zygaeninae, each
with their constituent taxa and currently accepted names. The diag-
nosis of the family is followed by a dichotomous key to the three
subfamilies of the western palaearctic Zygaenidae. Under Procridi-
nae there is a list of the constituent genera and characters that sep-
arate the procridines from other subfamilies; these general charac-
ters and their variation are then described in more detail. A
dichotomous key to the genera of the Procridinae is presented. Each
genus and subgenus, where applicable, is described in terms of di-
agnostic characters, followed by a detailed standardized description
of the constituent species. The pertinent details of each species in-
clude a reference to the imago by referring to plate number, length
of fore wing, brief description of both sexes, including descriptions
and drawings of their genitalia, similar species, individual geograph-
ical and ecological variation, and its distribution, including a range
map for every species. Where known, details of the immature stages,
including food plants, are also given. Every species is illustrated in
color. Dichotomous keys to species, sometimes between sexes, are
presented. The treatment of Chalcosiinae is as for Procridinae. The
Zygaeninae are introduced by a diagnostic overview of the subfam-
ily and the genus Zygaena, followed by a dichotomous key to the
species of the western Palaearctic. Descriptions of the various
species are then presented. An “Appendix” provides information on
the recently described Jordanita (J.) fazekasi. A comprehensive “In-
dex” concludes the text.
The overall design of the book is excellent. The book is well
bound with a quality decorative cover. The text, text figures, and pho-
tographs are on glossy paper and provide excellent detail. The text is
Journal of the Lepidopterists’ Society
55(1), 2001, 45-46
FORESTER Motus, by K. A. Efetov and G. M. Tarmann. 1999. Apollo
Books, 192 pp., 12 color plates with 241 illustrations, 174 line draw-
ings, 24 x 17 cm. Available from Apollo Books, price: 460 Danish Kro-
ner, excl. postage (hardback).
As a worker on the other half of the superfamily Zygaenoidea (i.e.,
the limacodid group), the publication of two books on Zygaenidae at
roughly the same time is like meeting your lost relatives. For those of
us that have large quantities of unsorted palaearctic zygaenids, this is a
welcome event indeed. It is also a major benefit for those interested in
the biologies and life histories of these beautiful diumal creatures and
makes literature more accessible to the English-speaking lepidopterist.
The scope of the two books on western palaearctic Zygaenidae
overlap in part, which at first glance is a bit confusing. “Forester
Moths” by Efetov & Tarmann is solely about the subfamily Procridinae
(also known as foresters), and its four genera and 63 species from the
western palaearctic. “The Westen Palearctic Zygaenidae” (1999, C. M.
Naumann, G. M. Tarmann and W. G. Tremewan, Apollo Books) covers
the same 4 genera, but only 44 species of Procridinae, in addition to the
Zygaeninae and Chalcosiinae. In fact, one might question why the two
books could not have been combined because of the overlap of the
Procridinae and in genitalia drawings, checklist, and keys.
“Forester Moths” by Efetov & Tarmann, perhaps the last book of
the two to be published in 1999, mentions the other as a
complementary text in its introduction. Indeed the focus on the
Procridinae by Efetov & Tarmann provides more detailed information
about this subfamily, including citations of misspelling and all known
synonymies and homonymies. It should be noted that there is a lack of
agreement between the checklists of the Procridinae regarding format
45
in an easy, well-organized reading style. The 178 text figures consist
of an array of diagrams, drawings and photographs; for every species
a distribution map is also given. The color reproduction of the plates
is of the highest quality, accurate, and shadow-free. Plates 1-6 illus-
trate 318 set specimens, often more than one exemplar of a species,
at life size with facing pages giving the species or subspecies, sex, lo-
cality and reference page number. Plates 7-8 show imagoes resting,
in copula or feeding; various aspects of behavior are shown on plate
9. Plate 10 is devoted to larvae. Plates 11-12, depicting various habi-
tats, will put shame to any tourist brochure! The book has been well
proofed and I could spot only smaller errors (misspelling of
Somabrachyidae (p. 14), alkaloids (p. 15) and food plants (p. 16)).
The only distraction is that the entry for the last species ends in mid-
sentence (p. 262), interrupted by the plates, to continue on p. 288;
the plates could have been bound following p. 290.
This book is beautifully produced and reasonably priced. Its com-
prehensive treatment resulting from the combination of authorita-
tive authorship and editorial care guarantee that this work will be
the standard reference on the zygaenid fauna of the western
Palaearctic for many decades to come. This book should not only be
on the bookshelf of lepidopterists, but it will also be of interest to
students of biogeography and evolutionary history of Lepidoptera
and other insects from a highly interesting faunistic region. I recom-
mend this book enthusiastically.
HENK GEERTSEMA, Department of Entomology and Nematology,
University of Stellenbosch, Private Bag X1, 7602, Matieland, South
Africa
and inclusion of taxa. Efetov & Tarmann separate the subgenera into
species groups and have subspecies, whereas Naumann et al. does not.
The discrepancy between the procridine species included (63 in
Efetov & Tarmann and 44 in Naumann et al.) between the two books
is in part due to the covered region and primarily concerns the genus
Adscita Retzius. Naumann et al. only contains species from the
Iberian Peninsula and North Africa to the Urals, including Turkey,
Transcaucasia and Caucasus, but Efetov & Tarmann, in addition,
include species from northern Iran and some countries east of the
Caspian Sea of “the western and central parts of Asia (i.e., the western
and central Palaearctis).”
“Forester Moths” begins with a concise introduction and definition
of the Procridinae. This is followed by an intriguing chapter (III) on
the characters of the subfamily including a chart and associated
illustrations that review the broad variation of dorsal and subdorsal
setae on the first abdominal segment in first instar larvae. This is
reminiscent of the broad variation found within the entire limacodid
group (Epstein, M. E. 1996. Revision and phylogeny of the limacodid-
group families, with evolutionary studies on slug caterpillars
(Lepidoptera: Zygaenoidea), Smithsonian Contributions to Zooology,
582). Next is a review of the chromosomal numbers in the subgenera.
The final part Chapter III includes a novel character combination
diagram and table, which divides the forester genera or subgenera
into groups based on shared characters of adult and immature stages.
These include biological (e.g:, host plants, diapause stage) and
chromosomal information, in addition to morphological data. There is
also a similar table for genera and host plant families. These tables
serve as a concise way to organize information on these genera,
although do not necessarily reflect the relationships between the
46
forester moths. Chapter IV on phylogenetic relationships, while
centering on the palaearctic genera of Procridinae, provides a useful
review for future workers on the worldwide fauna.
Chapter V is a checklist where each species is numbered and
corresponds to the species numbers in the next chapter, the systematic
catalogue. The text is very telegraphic in the catalogue, but there is
detailed information on the types (localities and place of deposition)
and host plants. I applaud the authors who took care of nomenclatural
business by designating 17 lectotypes. Distributional information is
provided, but according to the authors, maps are forthcoming in an
atlas which will complement the text. Two new subgenera, Tremewa-
nia and Procrita, are also described.
In the next chapter (VII), the authors have chosen to replace the
usual descriptions of taxa with keys to genera and species, and provide
large numbers of illustrations of key taxonomic characters, especially
of the genitalia in Chapter VII. These two chapters might have been
more logically placed before the systematic catalogue, but this is only
a minor inconvenience. Sadly, these excellent illustrations will be the
last by V. V. Kislovsky who died during a heart operation at age 24,
shortly before the book was published.
The penultimate chapter is on the immature stages and life
histories of six species in the genera Jordanita Verity and Adscita.
These include observations of the adults and biotype where the
species occur, along with detailed life history information on the
immature stages, with color photographs of each in the back of the
book. The final chapter presents a table of new parasitic records based
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
on host identification by the authors and parasitic information by
specialists for each group.
The book is made up of rag type paper, which has the disadvantage
of having somewhat dull images for the color plates and taking up
more space on your shelf than glossy paper for the same number of
pages. In the back of the book are seven plates. The first five plates
contain 130 paintings by N. V. Dyadenko of all included forester
species shown at 2x natural size. This is followed by a plate with
photographs of 16 of the 17 lectotypes designated in this book along
with early stages and adults in natural settings of Theresiminia Strand,
Rhagades, and Jordanita. The figure of the lectotype of Ino paupera
Christoph would have benefited from a lighter background. The next
two plates correspond to the life history information of Jordanita and
Adscita, mentioned above. These are followed by a plate of beautiful
zygaenid host plant photographs and three plates of scenic
photographs illustrating the diverse biotypes of foresters of the
western palaearctic.
I recommend both books on western palaearctic Zygaenidae for
serious study, because it is clear that you would be missing a great deal
of useful information on this group by owning just one of them.
Marc E. EPSTEIN, Department of Entomology, National Museum
of Natural History, Smithsonian Institution, Washington, D.C. 20560-
0105
Journal of the Lepidopterists’ Society
55(1), 2001, 47-50
EDITORIAL STAFF OF THE JOURNAL
CARLA M. PENZ, EDITOR
Department of Invertebrate Zoology,
Milwaukee Public Museum,
Milwaukee, Wisconsin 53233 USA
flea@mpm.edu
PHIL DEVRIES, BOOK REVIEW EDITOR
Center for Biodiversity Studies,
Milwaukee Public Museum,
Milwaukee, Wisconsin 53233 USA
pjd@mpm.edu
ASSOCIATE E’DITORS:
GERARDO LAMAS (Peru), KENELM W. PHILIP (USA), ROBERT K. ROBBINS (USA),
FELIX SPERLING (Canada), DaviD L. WAGNER (USA), CHRISTER WIKLUND (Sweden)
INSTRUCTIONS TO AUTHORS
The Journal of the Lepidopterists’ Society is a quar-
terly publication by the Lepidopterists’ Society. Con-
tributions to the Journal may address any aspect of
Lepidoptera study, including systematics, natural his-
tory, behavior, ecology, distribution, biogeography, and
evolution. Categories are Articles, Profiles, General
Notes, Technical Comments, Book Reviews, Obituar-
ies, Feature Photographs, and Cover Illustrations.
Obituaries must be authorized by the president of the
Society. Requirements for Feature Photographs and
Cover Illustrations are in volume 42(2):111.
Send Journal submissions to the editor at the above
address (or electronically to: flea@mpm.edu). Contrib-
utors should feel free to recommend one or two re-
viewers upon submission of their manuscript. Send re-
quests to review books for the Journal and book review
manuscripts to Phil DeVries at the above address (or
electronically to: pj}d@mpm.edu). Short manuscripts
concerning new state records, current events, and no-
tices should be sent to the News, Phil Schappert, Dept.
Zoology, University of Texas, Austin, TX 78712, USA
(or electronically to: philjs@mail.utexas.edu). Specific
instructions for electronic submissions appear below.
GENERAL MANUSCRIPT GUIDELINES
Submit manuscripts in triplicate, typewritten,
entirely double-spaced, with wide margins, on one
side only of white letter-sized paper. Use a legible
typeface (preferably a serif font) to allow for easy
identification of letters and numbers (e.g., number 1,
lower case L, and upper case I). Do not break words
with hyphenation at the right margin of the page. Do
not use right justification. Number pages in the upper
right corner.
When submitting hardcopies of the manu-
script via regular mail, submit them flat, not folded.
Submit original illustrations only after the manuscript
has been revised and accepted. Authors should pro-
vide a diskette or electronic copy of the final ac-
cepted manuscript in a standard PC or Macintosh-
based word processing format. Submit original
illustrations either in mounted or electronic form.
When submitting electronically, attach a stan-
dard PC or Macintosh-based word processing file, or a
PDF file, to your letter of submission via email. If your
word processing files include symbols such as ¢ ° p a
+ ®, or special table formats, please submit one hard-
copy via regular mail. Contributors can also send
scanned figures electronically. Upon receipt, the editor
will check your files for problems of incompatibility
between computer applications or systems. Electronic
submission expedites revision and publication, and it is
strongly recommended.
ARTICLES, PROFILES, AND TECHNICAL COMMENTS
Organize articles, profiles, and technical comments
as follows: title page (page 1), abstract and key words
page (page 2), text (pages 3 +), acknowledgments page
(numbered), literature cited pages (numbered), tables
(numbering optional), explanation of figures (number-
ing optional), and copies of figures (page numbers not
recommended).
Title page. The title page should include the man-
uscript title, author's name, affiliation, and full address,
RS ag
Te SS Eo Sr
oe
=H
including email address when available. Unless indi-
cated otherwise, it is assumed that the senior author of
multiple authored contributions will revise both the
manuscript and page proofs.
Make title explicit, descriptive, and as short as pos-
sible. It is unnecessary to include the word Lepi-
doptera in the title. List family names and other higher
level taxonomic categories in parenthesis.
Abstract and key words page. The indented
word “ABSTRACT.” (in large capitals, boldface, and
followed by a period) precedes a meaningful digest of
the manuscript. An abstract in Spanish can follow the
English abstract if desired by the author.
Up to five key words or terms not in the title should
accompany Articles (it is unnecessary to include the
word Lepidoptera), entitled Additional key words.
Text. Write with precision, clarity, and economy.
Use the active voice and the first person whenever ap-
propriate. Mark major sections of the manuscript
with centered headings set in small caps, but do not
use a heading for the introduction. Headings for taxo-
nomic, natural history, and experimental contributions
will differ and should follow standard scientific format.
Use articles published in the Journal as models. Sub-
headings within a section are followed by a period
and set in boldface. The example below shows a head-
ing and subheading:
RESULTS
Field studies. From 1959 through 1988, 18,255
Biston betularia .. .
In taxonomic manuscripts, taxon names can be
used both as headings and subheadings. They should
be centered, italicized, and followed by the author's
name in roman type. Year of publication is optional; if
provided, it should be separated from author's name
by a comma. Indicate new taxa and change in status of
a taxon in English, spelled fully, and in boldface (e.g.,
new species, new combination, new synonym). New
taxa with author and status appear entirely in boldface.
When needed, authors’ initials may be added to eluci-
date their identity. In the examples below, the genus
name appears as a section heading, and species names
are subheadings; new status is assigned to one species,
and one is being described as new:
Mylon Godman & Salvin, 1894
Mylon orsa Evans, 1953
Mylon exstincta Mabille & Boullet, 1917, new status
Mylon simplex Austin, new species
Descriptions should be clear and concise, employ-
ing standard terminology (e.g., head scoli, not head
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
horns) and traditional plural derivations (e.g., larva,
larvae; tarsus, tarsi; valva, valvae), and include mea-
surements (mean, range) and number of examined
specimens when applicable. Abbreviations such as FW
(forewing) and HW (hindwing) can be used to abridge
text. Depending on the circumstances, terminology for
descriptions can be used either in the singular or
plural forms, but text should be internally consistent.
When appropriate, manuscripts must name a public
repository where voucher specimens documenting
the identity of organisms can be found. Kinds of re-
ports that require vouchering include descriptions of
new taxa, life histories, host associations, immature
morphology, and some experimental studies.
In both descriptive and experimental studies, make
reference to figures (abbreviated Fig. or Figs.) and ta-
bles (Table, not abbreviated) whenever appropriate,
and these should be numbered and listed sequentially
in the text (i.e., Fig. 1 should be listed in the text be-
fore Fig. 2).
Examples of how to list references in text are:
Remington (1963), (Fruhstorfer 1913), Vansconcellos-
Neto (1986, 1991), DeVries (1991a, b), Vane-Wright
and Ackery (1989), (Clarke & Sheppard 1960), Roth-
schild et al. (1979). Unpublished data should be
cited as (unpublished), or (HFG unpublished) to
single out one of multiple authors when appropriate.
Personal communications should be cited as (J. W.
Brown pers. com.). Manuscripts in press should be
cited as such both in the text and Literature Cited sec-
tion, e.g., Epstein (in press). Only those manuscripts
accepted for publication should be cited as “in press;”
do not cite submitted papers or papers in review as “in
press.”
Use the following general guidelines for notations,
measurements, symbols, and other items. The first
mention of a plant or animal in the text should in-
clude the full scientific name with author and family.
For measurements, use metric units and abbreviate
them correctly (e.g., 15 km, 20 pg). For time, use a 24
h clock (0930 h, not 9:30 AM). For date, use “day
month year” format. Spell months fully; use full nota-
tion for year. As a recommendation of manuscript
style, numerals can be used when indicating day of
the month, measurements, statistics, anatomical
counts (e.g., 4 setae), standard entomological termi-
nology (e.g., forewing vein M3), and numbers of spec-
imens examined (e.g., 2 specimens), but should be
otherwise expressed in their word equivalent between
one (1) and nine (9). It is desirable to use male and
female symbols (¢ and 2) to condense accounts of ex-
amined material. Use italics for scientific names only
(genus and below). Underline only where italics are
VOLUME 55, NUMBER 1
intended. Do not use italics for emphasis (e.g., this
species occurs only at elevations above 800 m). Use ro-
man type for Latin abbreviations and expressions
(i.e., e.g., ca., et. al., sensu strictu, in situ, ad libitum, a
priori).
Acknowledgments page. Under the centered
heading ACKNOWLEDGMENTS, in one single paragraph
list persons that contributed to the study using either
their full name or initials, but remaining consistent
throughout the text. Fully spell out names of institu-
tions. List permit granting institutions when applica-
ble. Acknowledge financial support at the end of the
paragraph, followed by grant number when applicable.
Literature Cited pages. List references alphabet-
ically under the centered heading LITERATURE CITED.
Write authors’ names in small caps, and use roman
type for both reference and journal titles, except for
scientific names and special notations. Abbreviate
journal titles should as listed in the international Seri-
als Catalogue: Part I: Catalogue (International Council
of Scientific Unions Abstracting Board, 1978), except
when they consist of a single word (e.g., Biotropica).
Examples are listed below.
Books:
SHEPPARD, P.M. 1959. Natural selection and heredity.
2¢ ed. Hutchinson, London. 209 pp.
Book chapters:
JANZEN, D. H. 1988a. Guanacaste National Park:
Tropical ecological and biocultural restoration, pp.
143-192. In Cairns, J. J. (ed.), Rehabilitating dam-
aged ecosystems. Vol. II. CRC Press, Boca Raton,
Florida.
Journal articles:
POLLARD, E. 1977. A method for assessing changes in
the abundance of butterflies. Biol. Conserv.
12:115-124.
Multiple authors:
Nico.ay, S. S. & G. B. SMALL JR. 1969. A new sub-
species of Pyrrhopyge creon (Hesperiidae) from
Panama. J. Lepid. Soc. 23:127—130.
FAIRCHILD, W. L., D. C. Eipt, & C. A. A. WEAVER.
1987. Effects of fenitrothion insecticide on inhabi-
tants of leaves of the pitcher plant, Sarracenia pur-
purea L. Canad. Entomol. 119:647-652.
Multiple citations of the same author:
BELL, E. L. 1931. Studies in the Pyrrhopyginae, with
descriptions of several new species (Lepidoptera,
Rhopalocera, Hesperiidae). J. New York Entomol.
Soc. 39:417—491.
. 1933. Studies in the Pyrrhopyginae, with de-
scriptions of new species (Lepidoptera, Rhopalo-
cera, Hesperiidae). J. New York Entomol. Soc.
4]:265-295, 481-529.
49
Manuscripts in press:
JANZEN, D. H. In press. Ecology of dry forest wild-
land insects in the Area de Conservacifn Gua-
nacaste, northwestern Costa Rica. In Frankie, G.
W., A. Mata & S. B. Vinson (eds.), Biodiversity con-
servation in Costa Rica: learning the lessons in sea-
sonal dry forest. Univ. Calif. Press, Berkeley.
Proceedings of meetings:
PHILBRICK, R. N. (ed.) 1967. Proceedings of the Sym-
posium on the biology of the California islands. Santa
Barbara Botanic Garden, Santa Barbara, California.
Theses and dissertations:
PENZ, C. M. 1996. The higher-level phylogeny of the
passion-vine butterflies (Nymphalidae, Heliconi-
inae). Ph.D. Dissertation. University of Texas,
Austin, Texas.
Computer programs:
Maddison, W. P. & D. R. Maddison. 2000. MacClade:
version 4.0 PPC. Sinauer, Sunderland.
Anonymous institutional or organizational publi-
cations:
International Code of Zoological Nomenclature. 1985.
3" ed. International Trust for Zoological Nomen-
clature (BM[NH]). University of California Press,
Berkeley, California.
Tables. Number tables consecutively in Arabic nu-
merals. Label them in small caps (e.g., TABLE 1. ) and
use a concise and informative heading. Type each table
on a separate sheet and place after the Literature Cited
section, with the approximate desired position indicated
in the text. Avoid vertical lines and vertical writing.
Explanation of Figures. Type figure legends dou-
ble-spaced, on a separate sheet (not attached to the il-
lustrations), headed EXPLANATION OF FIGURES. Use a
separate paragraph for each legend. Color illustrations
are encouraged; contact editor for submission require-
ments and cost.
Figures. Illustrate only half of symmetrical objects
such as adults with wings spread, unless whole illustra-
tion is crucial. Mount photographs and drawings on
stiff, white backing, arranged in the desired format.
Bear in mind that your illustrations will be reduced to
fit a Journal page (plan to make lettering sufficiently
large), and that the figure legend will appear at the
bottom of the page (Journal page:16.5 cm width, 22.5
cm height). Illustrations larger than letter size are not
acceptable and should be reduced photographically to
that size or smaller. On the back of each illustration,
print the author's name and figure numbers as cited in
the text. Figures, both line drawings and photographs,
should be numbered consecutively in Arabic numer-
als; do not use “plate.”
eee
PRS SE AEE
FE
SSPE LO RS
EER aE
amas
50
GENERAL NOTES
Organize notes without page breaks as follows: title
(all capitals), additional key words, text, literature
cited, author name and full address, including email
address when available. Do not divide text into sec-
tions, but use boldface, indented headings when nec-
essary (for an example, see Cordero 1999, J. Lepid.
Soc. 53(4):169-170). Keep figures and tables to a min-
imum, but use when necessary. Acknowledgments are
given in the last paragraph of the text, without a sec-
tion heading. Title, Additional key words, and Litera-
ture Cited follow the same format as articles. General
guidelines for notations, measurements, symbols, and
other items are also the same as for articles.
Book REVIEWS
Provide full reference and supplementary informa-
tion as in the example below, followed by text, author's
name, and full address, including email address when
desired. Text should include a detailed overview of the
book reviewed and the author's critical appreciation of
contents, scientific merit, illustrations, and other perti-
nent issues.
THE MOTHS OF BORNEO. Part 9, Family Geometridae
(incl. Orthostixini), subfamilies Oenochrominae,
Desmobathrinae, Geometrinae, and Ennominae ad-
denda, by Jeremy Daniel Holloway. 1996. Malayan
Nature Journal 49: 147-326. Published in the Malayan
Nature Journal and also produced in paper covers by
Southdene Sdn. Bhd., Kuala Lumpur, Malaysia. 427
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
figures, 12 color plates. Soft cover, sewn binding, 17.9
by 25.3 em, ISBN: 983-99915-3-1. Available from
Southdene Sdn. Bhd., P.O. Box 10139, 50704 Kuala
Lumpur, Malaysia; Phone: 603-4022-2643; FAX: 603-
4022-226; e-mail: hsbar@pc.jaring.my; website:
www.edi.co.uk/barlow; Price $26.00, £18.00 (including
surface mail overseas).
PAGE CHARGES
For authors affiliated with institutions, page charges
are $50 per Journal page. For authors without institu-
tional support, page charges are $25 per Journal page.
For authors who are not members of the Society, page
charges are $75 per Journal page. Authors will be
charged for a full page for any partially filled pages.
Authors unable to pay page charges for any reason
should apply to the editor at the time of submission for
a reduced rate or free publication. Authors of Book Re-
views and Obituaries are exempt from page charges.
PAGE PROOFS
The edited manuscript and galley proofs will be
mailed to the author for correction of printer’s errors.
Excessive author's changes will be charged to authors
at the rate of $3.00 per line. A purchase order for
reprints will accompany proofs.
CORRESPONDENCE
Address all matters relating to the Journal to the ed-
itor. Address book reviews directly to the book review
editor.
Journal of the Lepidopterists’ Society
55(1), 2001, 51
MANUSCRIPT REVIEWERS, 2000
The merit of a scientific journal depends on the quality of its reviewers as well as its authors, but the former are usu-
ally unknown to the readers of the published articles. The Journal relied on the expertise of 49 reviewers last year
to provide 66 evaluations of manuscripts. It is with much gratitude that the Journal acknowledges the services of
the people listed below, who reviewed manuscripts for the 1999 volume of the Jowrnal. Those who reviewed two or
more manuscripts are denoted by asterisks. Our thanks to all of you.
Ackery, Phil, London, England
Aiello, Annette, Smithsonian Tropical
Research Institute
Bettman, David, Boulder, CO
Boggs, Carol, Stanford, CA
*Bowers, Deane, Boulder, CO
Brower, Lincoln, Sweet Briar, VA
Brown, John, Washington, DC
Burns, John, Washington, DC
Chew, Francis, Medford, MA
Common, Ian, Toowomba, Queensland, Australia
*Debinski, Diane, Ames, IO
*DeVries, Phil, Milwaukee, WI
Drummond, Boyce, Florissant, CO
Fitzgerald, Terrence, Cortland, NY
*Gall, Larry, New Haven, CT
Jenkins, Dale, Sarasota, FL
Landry, Bernard, Aylmer, Quebec, Canada
McCorkle, David, Monmouth, OR
*Metzler, Eric, Columbus, OH
Miller, Jackie, Sarasota, FL
Miller, Scott, Washington, DC
*Miller, William, St. Paul, MN
Nitao, James, Beltsville, MD
Nice, Chris, Madison, WI
Passoa, Steven, Powell, OH
*Penz, Carla, Milwaukee, WI
Pogue, Michael, Washington, DC
Porter, Adam, Amherst, MA
Powell, Jerry, Berkeley, CA
Prudic, Kathleen, Boulder, CO
Ritland, David, Due West, SC
*Robbins, Robert, Washington, DC
Rubinoff, Dan, Berkeley, CA
Rutowski, Ron, Tempe, AZ
Sargent, Theodore, Amherst, MA
Scholtens, Brian, Charleston, SC
Shapiro, Art, Davis, CA
Shappert, Phil, Austin, TX
Shuey, John, Indianapolis, IN
Simms, Steve, Maryland Heights, MO
*Solis, Alma, Brownsville, TX
Stamp, Nancy, Binghamton, NY
Tuskes, Paul, San Diego, CA
Tuttle, James, Tucson, AZ
Vane-Wright, Richard, London, England
*Wagner, David, Storrs, CT
Webster, Reginald, New Brunswick, Canada
Weiss, Martha, Washington, DC
Williams, Ernest, Clinton, NY
Date of Issue (Vol. 55, No. 1): 21 December 2001
ee
LEPIDOPTERISTS’ SOCIETY PUBLICATIONS
Memoir No. 5: Basic Techniques Manual, by William D. Winter, 2000. Members $29.00, non-members $44.00;
Canada, add $6.00; other countries, add $10.00.
Memoir No. 4: Foodplants of World Saturniidae, by Stephen E. Stone, 1991. Paper. ISBN 0-930282-05-1.
Members $9.00, non-members $14.00; outside US, add $5.00.
Memoir No. 3: Supplement to A Catalogue / checklist of Butterflies of America North of Mexico, edited by
Clifford D. Ferris, 1989. Paper. ISBN 0-930282-04-3. Members $8.00, non-members $12.00; outside
US, add $4.00.
Commemorative Volume 1947-1972, 1977. Includes cumulative Journal index, history of the Society, biographical
sketches. ISBN 0-930282-01-9. Cloth bound. Members $5.00: outside US, add $6.00.
All of the above prices include postage. Remittance must be in U.S. dollars, drawn on a U.S. bank, payable to The
Lepidopterists’ Society.
For Memoirs 3, 4, and 5 mail to:
The Lepidopterists’ Society
Kenneth Bliss, Publications Manager
P.O. Box 1366
Edison, NJ 08818-1366 USA
For all other publications, mail to:
Ron Leuschner
1900 John Street
Manhattan Beach, CA 90266-2608, USA
For more information on publications and membership, visit the Society's web site at:
http://www.furman.edu/~snyder/snyder/lep/
EDITORIAL STAFF OF THE JOURNAL
Carua M, Penz, Editor
Department of Invertebrate Zoology
Milwaukee Public Museum”
Milwaukee, Wisconsin 53233 USA
flea@mpm,.edu
Pit DeVries, Book Review Editor
Center for Biodiversity Studies
Milwaukee Public Museum
Milwaukee, Wisconsin 53233 USA
pjd@mpm.edu ;
Associate Editors:
“Grrarpo Lamas (Peru), Kenetm W. Puiu (USA), Ropert K. Rospins (USA),
Fe.ix Srertinc (Canada), Davin L. Wacner (USA), Cnrister Wikiunp (Sweden)
NOTICE TO ) CONTRIBUTORS
me or two reviewers upon obiasion of their rious Send rentees to review Baeks for the fi ral and na re-
ripts to Phil DeVries at the above address (or electronically to: pj\d@mpm.edu). Short manuscripts concerning new state
rrent events, and notices should be sent to the News, Phil Schappert, Dept. Zoology, University of Texas, Austin, TX 78712,
ee to: philjs@mail. utexas. od ). Before submitting manuscripts or electronic files, consult instructions in vol-
juscripts in triplicate, typewritten, Butieely double-spaced, with wide margins, on one side only of white letter-sized pa-
anuscript according to the following instructions, and submit them flat, not folded. Submit original illustrations only af-
script has been revised and accepted. Authors should provide a diskette cODy of the final bereptes manuscript in a stan-
cintosh-based word processing format.
Include an informative abstract for Articles, Profiles, and Technical Comments.
ditio ane words: Upto five. oe words or terms not in the title should accompany Articles, Profiles, General Notes, and
M. 1959. ‘Natural selection and feeds: 2nd ed. Hutchinson, evar 209 pp.
1961a. Some contributions to population genetics resulting from the study of the Lepidoptera. Adv. Genet. 10:165-216.
3: Illustrate only half of symmetrical objects such as adults with wings spread, unless whole illustration is crucial.
P Aerio and drawings on stiff, white backing, arranged in the desired format. Bear in mind that your illustrations will be
) fit a Journal page (plan to make lettering sufficiently large) and that the figure legend will appear at the bottom of the
rnal page: 16.5 em width, 22.5 cm height). Illustrations larger than letter-size are not acceptable and should be reduced
phically to that size or smaller, On the back of each illustration, print the author's name and figure numbers as cited in the
: roth line en and aaa sould be numbered aan in. sale numerals; cee not use eden Type
al enewl oe “Type each es ona separate sheet and Fae A the Literature Cited euuok, stath this spore tate desired po-
e 1 indicated i in the text, Avoid vertical lines and vertical writing,
oucher cece, - When ppprepnats aeecuns must name a public repository where ia ee hae se
ge i For ees affiliated with institutions: page charges are $50 per hee page. For sige swithout institutional
page charges are $25 per Journal page. For authors who are not members of the Society, page charges are $75 per Journal
uthors unable to pay page charges for any reason should apply to the editor at the time of submission for a reduced rate or
lication. Authors of Book Reviews and Obituaries are exempt from page charges.
_ Correspondence: Address all matters relating to the Journal to the editor. Address book reviews diosbsly to the book review
c ee by :
i Pantien BY THE ALLEN PRESS, INC.. LAWRENCE, KANSAS 66044 U.S.A.
xa: Riot i
oa ae He its SisteMarics. “AND. 5 BioLoer Le IDO TE
David Ahrenholz -
: ; Volume 55 2001 Number 2
ISSN 0024-0966
JOURNAL
: of the
_ _Lepmpoprerists’ SOCIETY
Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN
Publicado por LA SOCIEDAD DE LOS LEPIDOPTEROLOGOS
i AE ‘ " AN x Ri CN
fut ; = DAI renee
Mita
Ua
15 February 2002
_ mote the science of lepidopterology in all its branches, . . . to issue a periodical and other publications on Lepidopter te
gether with their full name, address, and special lepidopterological interests. Tn alternate years a list.of members of aut !
-sued, with addresses and special interests. ‘ cat
THE LEPIDOPTERISTS’ SOCIETY
EXECUTIVE Cisunce.
Joun W. Brown, President en SUSAN $i Borkin, Vice President ‘iia ater ee z
Micuaet J. Smiru, Immediate Past President Mirna M. CAsaGRANDE, Vice President
.Manuen A. BatcazAr-Lara, Vice President ~ Davin C. Irtner, Treasurer
Ernest H. Wiiuiams, Secretary Tes
ay ; Members at large: avn tera
Ronald L. Rutowski ~ M. Deane Bowers - George L. Balogh
Felix A. H. Sperling Ron Leuschner _ Andrew V. Z. Brower
Andrew D. Warren Michael Toliver _. Brian Scholtens:
EprroriaL BoarD
Rosert K, Rossins (Chairman), Joun W. Brown (Member at large)
Caria M. Penz (Journal),
WituaM EB. Mitrer (Memoirs)
Purr J. SCHAPPERT (News)
HONORARY Lire MEMBERS OF THE SOCIETY
Onsnins L, Remincton (1966), E. G. Munror (1973), Ian F. B. Common (1987),
Joun G. Francuemont (1988), Lincoun P. Brower (1990), Doucras C. Fercuson “ey
- Hon. Miriam Roruscurp (1991), Cuaupe Lemaire (1992), Freperick H, ee? (i997) hes
The object of The Lepidoptedsts Sanee, which was formed in May 1947 and formally constituted in Dedenben 19 a;
the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cOOne on in, o
sures” directed towards these aims. :
Membership im the Society is open to all persons interested in the Study) of Lepidoptera, All tambet receive the
News of The Lepidopterists’ Society. Prospective members should send to the Assistant Treasurer full dues for the «
Active members—annual dues $45.00 within the U. Se $50. 00 ated the U.S.
Affiliated members—annual dues $10.00 within the U.S., $15.00 outside the US,
Student members—annual dues $20.00 within the U.S., $95, 00 outside the U.S.
Sustaining members—annual dues $60.00 within the U. S., $65. 00 outside the U. se
Life members—single sum $1,800.00 ‘dae ue
Institutional subscriptions—annual $60.00 rer eS a SN A) Si
Airmail postage for the News $15.00 cao
Send remittances, payable to The Lepidopterists’ Society, to: Kelly M. Richers, Asst. Tredsines, 9417 Camsalhe Coles B field,
CA 93311; and address changes to: Julian P. Donahue, Natural History Museum, 900 Exposition Blyd., Los Angeles, CA 90007-4
For information about the Society, contact: Ernest H. Williams, Department of Biology, Hamilton College, Clinton, NY 1332
order back issues of the Memoirs, write for availability and prices to Kelly M. Richers, Asst. Treasure er, 9417 Carvalho Court, !
field, CA 93311. we
The additional cost for members outside the U.S. is to cover mailing costs.
ai
ng
and at additional crtlials offices POSTMASTER: “send mace changes to The ieee soa Ni stra I History Musev
900 Exposition Blvd., Los Angeles, CA 90007-4057. ;
Cover illustration: Rotschildia zacateca male. Drawn by artist John Cizeski (johnc@coredes.com) from a photo by K, Wolfe. Ey
JOURNAL OF
tne ILPPIDOPTERISTS’ SOCIETY
Volume 55
Journal of the Lepidopterists’ Society
55(2), 2001, 53-58
2001 Number 2
SPECIES’ COMPOSITION OF MOTHS CAPTURED IN TRAPS BAITED WITH ACETIC ACID AND
3-METHYL-1-BUTANOL, IN YAKIMA COUNTY, WASHINGTON
PETER J. LANDOLT
USDA-ARS, Yakima Agricultural Research Laboratory,
5230 Konnowac Pass Road, Wapato, Washington 98951, USA
AND
PAUL C. HAMMOND
Department Entomology, Oregon State University, Corvallis, Oregon 97331, USA
ABSTRACT. Moths were captured in traps baited with acetic acid and 3-methyl-1-butanol at four sites in Yakima County, Washington,
from March to October, 1999. Nine hundred and ninety-one moths captured were identified to 60 species. Three families of Lepidoptera were
represented: Noctuidae, Thyatiridae, and Pyralidae. The majority of species (90%) and individuals (90%) were noctuids. These included many
non-pest species and numbers of several pest species: the forage looper Caenurgina erechtea Cramer, glassy cutworm Apamea devastator Brace,
bertha armyworm Mamestra configurata Walker, true armyworm Pseudaletia unipuncta Haworth, and spotted cutworm Xestia c-nigrum (L.).
Noctuids collected included representatives of the subfamilies Catocolinae, Cuculliinae, Hadeninae, Amphipyrinae, and Noctuinae. The ma-
jority of moths trapped were captured during August and September.
Additional key words: trapping, attractant, survey, biodiversity, sampling.
Many insect species, including Lepidoptera, are at-
tracted to baits containing fermenting sugar solutions
or other sweet materials (Norris 1935). Such baits have
long been used by moth and butterfly collectors (e.g.,
Holland 1903, Sargent 1976) and have been employed
or investigated by applied entomologists attempting to
monitor or control pest species of moths (Ditman &
Cory 1933, Eyer 1931, Frost 1926, Landolt 1995, Lan-
dolt & Mitchell 1996). Recently, Yamazaki (1998) used
sweet baits in traps as a means of sampling moths for
survey and ecological work.
The identification of attractive chemical odorants
from fermented sugar baits provides opportunities to
develop useful insect lures. Three pest species of Noc-
tuidae, Lacanobia subjuncta (Grote & Robinson), the
bertha armyworm Mamestra configurata Walker, and
the spotted cutworm Xestia c-nigrum (L.), are at-
tracted to a combination of acetic acid and 3-methyl-1-
butanol (Landolt 2000). Acetic acid and 3-methyl-1-
butanol are found in fermented sweet baits and some
microbial cultures (DeMilo et al. 1996, Utrio & Eriks-
son 1977). These compounds are thought to be feed-
ing attractants for those moths (Landolt 2000) and are
being developed as lures for monitoring and control-
ling pest species of Noctuidae (Landolt & Alfaro 2001,
Landolt & Higbee 2002).
The purpose of this study was to determine what
types of moths are attracted to acetic acid and 3-methyl-
1-butanol more thoroughly than in previous surveys.
In a study that documented attraction of L. subjuncta,
M. configurata, and X. c-nigrum to these chemicals
(Landolt 2000), other moths were captured but were
not identified. We sought to determine if other pest
species of Lepidoptera are attracted to this lure. A sec-
ondary objective was to document responses of non-
target species in order to assess the complexity of us-
ing the lure to monitor pests. Knowledge of the types
of insects responding to this lure may suggest addi-
tional uses of the attractant in survey and sampling
studies, such as that of Yamazaki (1998).
MATERIALS AND METHODS
Moths were trapped with Butterfly Bait Traps (Bio-
Quip Products, Santa Monica, California). These cage
e
|
|
|
54 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
TABLE 1. Species of moths captured in traps in Yakima County, 1999.
Total
Species Orchard Parker Moxee Ahtanum 2 c)
Thyatiridae
Euthyatira semicircularis (Grote) 0 0 0 1 0
Habrosyne scripta (Gosse) 0 1 0 0 1 0
Pseudothyatira cymatopheroides (Guenée) 0 1 0 Il
Noctuidae
Catocolinae
Caenurgina erechtea (Cramer) 0 IL 8 I 8 2)
Catocala stretchi Behr 0 0 2 0 2 0
Catocala briseis Edwards 0 0 0 2 2 0
Zale lunata (Drury) 0 3 1 1 5 0
Plusiinae
Autographa californica (Speyer) 0 0 2 0) 2 0
Amphipyrinae
Protagrotis obscura (Barnes &
MacDunnough) 0 0 0 1 0 1
Apamea (Agroperina) dubitans (Walker) 0 1 il 2) 1 3
Apamea amputatrix (Fitch) 0) 1 0 1 2 0)
Apamea (C rymodes) devastator (Brace) 0) 5 a 58 54 89
Apamea occidens (Grote) 0 0) 1 0) Il 0
Aseptis characta (Grote) 0 0 18 0) 9 9
Caradrina meralis (Morrison) 0) 1 9 0 4 6
Caradrina morpheus (Hufnagel) 0 0 1 0 0 il
Chytonix divesta (Grote) 0 1 0 0 0 1
Oligia indirecta (Grote) 0 0 0 3 1 2
Oligia tonsa (Grote) 0 i 9 0 5 5
Athetis mindara (Barnes & MacDunnough) 0 0 1 0 1 0
Spodoptera praefica (Grote) 0 0 2 0 0 2
Cuculliinae
Epidemas cinerea Smith 0) 0 8 0 4 4
Hadeninae
Aletia oxygala (Grote) 0 7 18 91 63 53
Leucania farcta (Grote) 0 1 0) 1 il 1
Discestra oregonica (Grote) 0 0 0 1 1 0
Discestra mutata (Dod) 0) 0 0 il Il 0
Dargida procinta (Grote) 1 3 130 4 67 71
Lacanobia subjuncta (Grote & Robinson) 1 0 0 0 1 0
Lacinipolia stricta (Walker) 0 3 0 14 10 7
Lacinipolia vicina (Grote) 0) 0 8 ) 4 4
Lygephila victoria (Grote) 0 0 1 0 IL 0
Mamestra configurata (Walker) 0) 0 Dil 3 12 12
Protorthodes curtica (Smith) 0 4 17 24 27 18
Anhimella contrahens (Walker) 0 0 2 0 0 2
Pseudaletia unipuncta (Haworth) 0 26 15 3 20 24
Noctuinae
Abagrotis negascia (Smith) 0 2 1 1 4 0
Agrotis ipsilon (Hufnagel) 1 0 0 0 il 0
Agrotis venerabilis (Walker) 0 0 2 0 2 0
Agrotis vetusta (Walker) 0 PD} 2 4 5 3
Diarsia rosaria (Grote) I It 0 3 4 I
Euxoa albipennis (Grote) 0 0 1 1 2 0
Euxoa atomaris (Smith) 1 0) 1 0 1 1
Euxoa auxiliaris (Grote) 0) 0 il 0 1 0
Euxoa choris (Harvey) 0 0 1 () 1 0
Euxoa declarata (Walker) 0) 0 1 0 0 1
Euxoa idahoensis (Grote) ] 0 1 il 3 0
Euxoa infausta (Walker) 1 0 0 0 1 0
Euxoa septentrionalis (Walker) I 8 135 ) 101 52
Feltia jaculifera (Guenée) 0 0 33 1 17 17
VOLUME 55, NUMBER 2 55
TABLE 1. Continued
Total
Species Orchard Parker Moxee Ahtanum )
Peridroma saucia (Hiibner) 0 0 1 0 0 ]
Rhynchagrotis formalis (Grote) 0 0 6 0 6 0
Spaelotis bicava LaFontaine 0 0 1 0 1 0
Spaelotis clandestina (Harris) 0 0 ) 0 2 0
Xestia c-nigrum (L.) 1 5 12 8 15 11
Xestia (Anomogyna) infimatis (Grote) 0 0 2 1 3 0
Xestia plebeia (Smith) 0 0 1 4 5 0
Xestia xanthographa (Denis &
Schiffermiiller) 0 0 1 0 1 0
Pyralidae
Hypsopygia costalis (Fabr.) 0 0 0 93 38 55
Udea profundalis (Packard) 0) 0 1 0 0 1
Pyralis farinalis L. 2 2 3 1 i 1
traps are fiberglass screen cylinders (90 cm high and 38
cm wide) hung vertically about 3 cm above a 40 cm wide
wooden platform (figured in Covell 1984:16). The trap
entrance is a screen cone at the bottom of the screen
cylinder and above the wooden platform. The lure was a
30 ml polypropylene vial containing 15 ml of a 1:1 mix-
ture of glacial acetic acid (Baker Chemical, Pittsburgh,
Pennsylvania) and 3-methyl-1-butanol (Aldrich Chemi-
cal, Milwaukee, Wisconsin) on cotton. Release of the
chemicals from the vial was through a 6.4 mm diameter
hole in the vial lid. The vial was positioned on the
wooden platform directly beneath the center of the trap
opening, held in place by three corks glued to the plat-
form. Our objective was to obtain relatively undamaged
specimens that are identifiable; large numbers of insects
captured in the butterfly bait trap can perch relatively
undisturbed on the screening of the trap after capture.
Four cage traps were set up on 12 March 1999, each
at a different site with somewhat different habitats in
Yakima County, Washington. All traps were hung from
tree branches. The first trap was placed within a com-
mercial apple orchard near the town of Donald, 11 km
southeast of Union Gap. This site received multiple
applications of conventional pesticides for control of
insect pests. The second trap was placed along the
edge of a forested riparian habitat along the Yakima
River, 7 km southeast of Union Gap, and adjacent to
an apple orchard. The third trap was placed in a wind-
break of Douglas Fir trees, at the Moxee Experimen-
tal Farm, 20 km east of Moxee. This trap was near
mixed irrigated agriculture and native dryland or
steppe plant community. The fourth trap was placed
along a riparian habitat, along Ahtanum Creek, 10 km
west of Union Gap, and adjacent to small farms and
pasture. Traps were checked once or twice per week,
depending on the amount of moth activity, and lures
were replaced every 2 weeks. Traps were maintained
until 1 October 1999.
To assess the relative effectiveness of the butterfly
bait trap in comparison to methods used in previous
studies (Landolt & Alfaro 2001) we compared four
butterfly bait traps to four Universal moth traps (Uni-
Trap). Universal moth traps have been used in previ-
ous experiments for trapping L. subjuncta (Landolt &
Alfaro 2001). All traps were baited with 30 ml
polypropylene vials with a 6 mm hole in the lid. Vials
were loaded with 15 ml of a 1:1 mixture of acetic acid
and 3-methyl-1-butanol. The eight traps were placed
in one row of an apple orchard, with butterfly bait
traps and UniTraps alternated in the row. All traps
were positioned so that the bait was at a 2 m height
and traps were 20 m apart. Treatment (trap design)
positions were switched each time that traps were
checked. Traps were checked twice per week from 4 to
17 May 2000. Treatment means for pooled male and
female trap catch data were compared using Student's
t-test, with separate analyses for L. subjuncta, M. con-
figurata, and X. c-nigrum trap catch data.
Voucher specimens are deposited in the collection
of the Department of Entomology, Washington State
University, Pullman, Washington.
RESULTS
A total of 991 moths were collected in the 4 butter-
fly bait traps maintained from 12 March to 1 October
1999. Sixty species were recovered (Table 1). Some
specimens (<10%) in several late summer samples
were not identifiable because of severe loss of wing
scales. Noctuids constituted 90% of specimens and
91% of species (56 of 62) of moths captured in traps.
The remaining species were in the families Thyatiridae
(3 species) and Pyralidae (3 species) (Table 1). A rela-
56
400
”
<
Lu
Lu
Ss 300
N
Qa
Lu
a 200
ou
<=
oe
-
”
+ 100
-E
2)
=
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
3/19 4/2 4/16 4/30 5/14 5/28 6/11 6/25 7/9 7/23 8/6 8/20 9/3 9/17 10/1
DATE (MONTH/DAY)
Fic. 1. Numbers of Noctuidae (black bars) and Pyralidae (white bars) captured in 4 cage traps baited with acetic acid and 3-methyl-1-bu-
tanol lures, at 2 week intervals through the season. Yakima County, Washington State, 1999.
tively small number of species constituted a large per-
centage of the moths captured. The four noctuid
species Apamea devastator Brace, Dargida procincta
Grote, Aletia oxygala (Grote), and Euxoa septentrion-
alis Walker and the pyralid Hypsopygia costalis (Fab.)
together constituted 65% of the moths trapped and
identified. For most species of noctuids trapped, both
sexes were caught (Table 1).
During the seasonal study there were relatively few
types of other insects (non-Lepidoptera) captured in
the traps. Of note, no butterflies and no bees
(Apoidea) were caught. However, eight Sceliphron
caementarium (Drury) mud dauber wasps (Sphecidae)
were captured at the Ahtanum Creek site and the yel-
low jackets Vespula germanica (Fabr.) and Vespula
pensylvanica (Saussure) (Vespidae) were captured
fairly consistently in late summer at all sites. Very few
moths were captured in traps before August, with
most moths captured from late August through late
September (Fig. 1).
TABLE 2. Means (+SE) numbers of 3 pest species of Noctuidae
captured in Universal moth traps (Unitrap) and in butterfly bait traps
(cage) baited with acetic acid and 3-methyl-1-butanol, in apple or-
chards in Yakima County, Washington. May 2000.
Moth species Unitrap Cage trap t value p
Lacanobia subjuncta 3.7+0.9 2.5 + 0.6 1.40 0.09
Mamestra configurata 11+04 06+0.2 2.24 0.02
Xestia c-nigrum 3.6 + 1.1 3.2 + 0.9 0.49 0.31
In the comparison of butterfly bait traps and Uni-
Traps, significantly more M. configurata moths were
captured in UniTraps (Table 2). However, numbers of
the other 2 species of pest moths targeted in this study,
L. subjuncta and X. c-nigrum, were not significantly
different between trap types (Table 2).
DISCUSSION
Landolt (2000) found that the combination of acetic
acid and 3-methyl-1-butanol is attractive to both sexes
of 3 noctuid pests that are common in Washington
state apple orchards; L. subjuncta, M. configurata, and
X. c-nigrum. The results of this study confirm that this
lure attracts a variety of moths, predominantly of the
family Noctuidae (Table 1). The capture of several
other pest species, including the glassy cutworm A.
devastator, the true armyworm Pseudaletia unipuncta
(Haworth), and the forage looper Caenurgina erechtea
(Cramer) (Table 1) indicates that this chemical attrac-
tant may have broad application for use in monitoring
and control of noctuid pests of agricultural crops.
These and other species captured (Table 1) indicate a
taxonomically diverse response within the family Noc-
tuidae, with 5 of the noctuid subfamilies represented
in the sampling. This lure in a suitable trap may prove
useful also as a means of sampling moth biodiversity,
as was attempted by Yamazaki (1998) using fermented
sugar solutions.
It is interesting that numbers of these 3 moth
species were captured consistently in apple orchards
VOLUME 55, NUMBER 2
during the comparison of the two trap designs and
were only infrequently trapped at the 3 non-agricul-
tural sites used for the season-long study (Table 1),
while only one female L. swhjuncta was captured in
the season-long study in a different apple orchard.
Populations of these moths are expected to vary with
the season (Hitchcox 2000) and appear to vary greatly
between orchards, depending on prior history and
management practices (unpublished data).
The reasons for the pronounced seasonal pattern in
captures of moths are not known but may include a
relative abundance of some species of moths at trap-
ping sites late in the season compared to early sum-
mer. For example, the cutworm Euxoa septentrionalis
(Walker) (135 captured) is univoltine and flies from
late August into October (Lafontaine 1987). There are
other species of moths known to be attracted to acetic
acid and 3-methyl-1-butanol which are bivoltine and
are present earlier in the season, such as M. configu-
rata and L. subjuncta, (Landolt 2000). Perhaps they
were simply not present at the trapping sites used in
this study but have been common in other apple or-
chards.
There was a near absence of species in the noctuid
subfamily Cuculliinae in traps (Table 1). In contrast to
our study, Yamazaki (1998) captured 15 species of cu-
culliine moths during spring with fermented sugar so-
lutions. Their general absence in our traps (excepting
Epidemas cinerea Smith) may be due to an absence of
host plants at the sites sampled. Moreover, Yamazaki
(1998) worked in a secondary forest, which was not a
habitat type included in this study. It is also possible
that some moths attracted to fermented sweet baits
may be attracted by different blends of chemicals, and
not to the combination of acetic acid and 3-methyl-1-
butanol.
It is not known if the lack of captures for other
groups of insects, such as butterflies, relates to low or
nonexistent populations or to low or non-existent
responsiveness of those insects to acetic acid and
3-methyl-1-butanol. It is suspected that yellow jacket
wasps may have been visually attracted into traps by
the presence of prey items (captured insects). How-
ever, both V. germanica and V. pensylvanica are
weakly attracted to some degree by the combination of
acetic acid and 3-methyl-1-butanol (Landolt et al.
2000).
Based on the results of the trap comparison test, we
do not think that many species of moths were missed
due to the use of the cage trap, rather than the Uni-
Trap, which has been used extensively in studies to de-
velop the lure for pest species (Landolt & Alfaro
2001). However, some species of moths attracted to
|
Ol
acetic acid and 3-methyl-1-butanol likely are better
captured with different designs of traps.
In summary, this study indicates broad attractive-
ness of acetic acid with 3-methyl-1-butanol to a diver-
sity of moth species, predominantly Noctuidae, with a
number of agricultural species included. This lure, in
an appropriate trap, may then be useful as a means of
sampling moth populations in ecological or environ-
mental studies, for general collecting BE moths, and for
monitoring of pest “yqpullaitons in agricultural crops.
The general attractiveness of the lure to noctuids is of
some concern for lure use in monitoring specific agri-
cultural pests, because the monitoring of targeted
species will be complicated by the trapping of non-tar-
get species, requiring time spent sorting specimens.
ACKNOWLEDGMENTS
Technical assistance was provided by D. L. Larson, P. S. Chap-
man, J. R. Beauchene, and L. L. Biddick. Helpful comments to im-
prove the manuscript were made by H. C. Reed and R. S. Zack. This
work was supported in part by a grant from the Washington Tree
Fruit Research Commission.
LITERATURE CITED
CovELL, C. V. JR. 1984. A field guide to the moths of eastern
North America. Houghton-Mifflin Co., Boston, 496 pp.
DeEmILO, A. B., C. J. LEE, D. S. MORENO & A. J. MARTINEZ. 1996.
Identification of the volatiles derived from Citrobacter freundii
fermentation of a trypticase soy broth. J. Agric. Food Chem.
44:607-612.
Dirman, L. P. & E. N. Cory. 1933. The response of com earworm
moths to various sugar solutions. J. Econ. Entomol. 26:
109-115.
EYER, J. R. 1931. A four year study of codling moth baits in New
Mexico. J. Econ. Entomol. 24:998—1001.
Frost, S$. W. 1926. Bait pails as a possible control for the oriental
fruit moth. J. Econ. Entomol. 19:441—450.
Hircucox, M. 2000. Seasonal phenology and monitoring of La-
canobia subjuncta (Noctuidae: Lepidoptera) in apple orchards
of Washington State. Masters Thesis, Washington State Univer-
sity, Pullman, Washington. 75 pp.
HOLLAND, W. J. 1903. The moth book. A guide to the moths of
North America. Dover Publications Inc., New York. 479 pp.
LAFONTAINE, J. D. 1987. Noctuidae. Noctuinae (Part Euxoa). In
Hodges, R. W. (ed.), Moths of America north of Mexico. Fasci-
cle 27.2. 237 pp., Wedge Entomological Research Foundation,
Washington, D.C.
LANDOLT, P. J. 1995. Attraction of Mocis latipes (Lepidoptera: Noc-
tuidae) to sweet baits in traps. Florida Entomol. 78:523—550.
. 1998. Chemical attractants for trapping yellowjackets
Vespula germanica and Vespula pensylvanica (Hymenoptera:
Vespidae). Environ. Entomol. 27:1229-1234
2000. New chemical attractants for trapping Lacanobia
subjuncta, bertha armyworm, and spotted cutworm (Lepi-
doptera: Noctuidae). J. Econ. Entomol. 93:101—106.
LANDOLT, P. J. & J. F. ALFARO. 2001. Trapping Lacanobia sub-
juncta, Xestia c-nigrum, and Mamestra configurata (Lepi-
doptera: Noctuidae) with acetic acid and 3 -methyl-1-butanol
in controlled release dispensers. Environ. Entomol. 30:
656-672.
LANDOLT, P. J. & B. S. HicBEE. In press. Both sexes of the true
armyworm (Lepidoptera: Noctuidae) trapped with the feeding
attractant composed of acetic acid and 3-methyl-1- inieraall
Florida Entomol.
PR a
ee
58
LANDOLT, P. J. & E. R. MITCHELL. 1996. Attraction of tobacco bud-
worm moths (Lepidoptera: Noctuidae) to jaggery, a palm sugar
extract. Florida Entomol. 80:403—407.
LANDOLT, P. J., C. S. SMITHHISLER, H. C. REED & L. M. McDo-
NouGH. 2000. Trapping social wasps (Hymenoptera: Vespidae)
with acetic acid and saturated short chain alcohols. Environ.
Entomol. 93:1613-1618.
Norris, M. J. 1935.The feeding habits of the adult Lepidoptera:
Heteroneura. Trans. Roy. Entomol. Soc. London 85:61—90.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
SARGENT, T. D. 1976. Legion of night. Univ. Massachusetts Press,
Amherst, Massachusetts. 222 pp
Urrio, P. & K. ERIKSSON. 1977. Volatile fermentation products as
attractants for macrolepidoptera. Ann. Zool. Fennici 14:98-104.
YAMAZAKI, K. 1998. Communities of early spring noctuid and thy-
atirid moths (Lepidoptera) molasses-trapped in secondary
forests. Entomol. Science 1:171-178.
Received 3 November 2000; revised and accepted 21 August 2001.
Journal of the Lepidopterists’ Society
55(2), 2001, 59-62
A NEW AGONOXENINE MOTH FROM THE GALAPAGOS ISLANDS
(GELECHIOIDEA: ELACHISTIDAE)
BERNARD LANDRY
Muséum dhistoire naturelle, C.P. 6434, CH-1211, Genéve 6, Switzerland
ABSTRACT. The only member of the Elachistidae, Agonoxeninae so far encountered in the Galapagos Islands is described as a new
species: Haplochrois galapagosalis, new species.
RESUMEN. Se describe la unica especie de Elachistidae, Agonoxeninae de las islas Galapagos Haplochrois galapagosalis, sp. n.
Additional key words: Ecuador, South America, Croton sp., Euphorbiaceae.
The Agonoxeninae moths of the world are poorly
known. Hodges (1997) reviewed the classification of
their Neotropical members, and Sinev (1999) syn-
onymized Tetanocentria Rebel (1902) with Hap-
lochrois Meyrick (1897), the genus to which the only
Galapagos agonoxenine species belongs. This is the
first record of Agonoxeninae for the famous Ecuado-
rian archipelago.
The original description of Haplochrois (Meyrick
1897; type-species H. chromatella Meyrick 1897) is
brief, but Bottimer (1926), Clarke (1965), and Riedl
(1969) provide more information on adult and larval
morphology of congeneric species. Based on Hodges
(1983, 1997), Sinev (1999), and the new species
recorded here, Haplochrois includes 30 described
species distributed in tropical and subtropical regions
of the world. The genus is diagnosed by characters of
the male genitalia; ie., the elongate and narrow tegu-
men and saccus, paired setose gnathos, and soft-lobed
valvae (Sinev 1999). The available information on host
plant comes from Kusnezov (1916), who described H.
theae from larvae reared from the tea bush (Camellia
sinensis (L.) O. Kuntze, Theaceae) in Transcaucasia,
and from Bottimer (1926), who reported H.
bipunctella (Chambers) larvae from leaf petiole galls
and seeds of Croton engelmannii Ferguson (=C. capi-
tatus Michaux, Euphorbiaceae) in Texas.
According to Hodges (1997), Tetanocentria (now
Haplochrois) includes seven Neotropical species; six
described by E. Meyrick and one by A. Busck. I exam-
ined the holotype (or a paratype in one case) of each of
these species, as well as specimens of the North
American species, H. bipunctella, to conclude that the
Galapagos taxon is new.
Haplochrois galapagosalis B. Landry, new species
(Figs. 1-9)
Diagnosis. The long and narrow forewing and hindwing, the up-
turned labial palpus, the scaled haustellum, and the simple wing pat-
tern will separate this species from all other Galapagos moths. Two
species of Gracillariidae may appear somewhat similar in color and
size, but their forewing patterns are different and they don’t have
the haustellum scaled. Pyroderces rileyi (Walsingham), a Cosmopte-
rigidae, also appears superficially similar to H. galapagosalis, but its
forewing is mostly dark brown with an oblique paler line subterminally,
and its antennae are annulate black and white (see Landry 2001).
Description. Male (n = 17). Head beige along median line with
brown to dark-brown scales laterally. Haustellum beige. Labial pal-
pus beige medially on segment II and at base and apex of segment
II, brown laterally on segment II, dark brown on most of segment
III. Antenna beige to dark brown, slightly paler on scape and first
few flagellomeres; pecten made of about 25 long and thin scales; fla-
gellomeres simple, with short setae and two sets of scales, the sec-
ond erect. Thorax beige with three longitudinal dark-brown lines
apically, or dark brown, but “greasy” in all specimens and conse-
quently hard to characterize. Foreleg coxa dark brown with beige
scales at base; femur dark brown, some specimens with scales bicol-
ored beige basally and dark brown distally; tibia mostly dark brown
with beige scaling at 1/3, 2/3, and terminally; tarsomeres I and II
mostly dark brown with scales bicolored beige basally and dark
brown distally, with beige spot subterminally; tarsomere III mostly
beige, with dark-brown scales at base and apex; tarsomere IV dark
brown; tarsomere V beige with dark brown at base. Midleg coxa and
femur dark brown; tibia dark brown with scales bicolored beige
basally and dark brown distally, darker brown toward apex, with
short and thin projecting scales dorsally at base, middle, and apex;
tarsomeres dark brown with scales bicolored beige basally and dark
brown distally, with darker brown scales apically on tarsomeres I-IV.
Hindleg coxa with a mixture of beige and brown to dark-brown
scales; femur dark brown; tibia dark brown with scales bicolored
beige basally and dark brown distally, with darker brown spots at
base, middle, and apex, and with projecting long and thin pale-
brown scales on entire dorsal margin and on ventral margin as a
patch posterior to anterior pair of spurs; tarsomeres I-III with beige
to dark-brown scales bicolored beige basally and dark brown distally,
with darker brown scales apically; tarsomeres IV-V uniformly pale
grayish brown; tarsomere I also with darker brown spot at middle
and with patch of thin and long projecting scales dorsoapically; tar-
somere II also with a few long and thin pale-brown scales projecting
at apex dorsally. Forewing (Figs. 1, 2) length: 4-6 mm; ground
color beige to dark grayish brown; pale specimens sometimes with
slightly paler dorsal half and with most conspicuous dark-brown
markings consisting of a small dot above middle subbasally, a short
streak subbasally on cubital fold, a spot on cubital fold submedially
(usually the most conspicuous, and sometimes only, marking), a
smaller spot above end of cubital fold, some specimens with dark-
brown scales dispersed between latter spot and costa, and three or
four small subapical and apical spots on dorsal margin; fringe mostly
dark brown at apex, pale brown to beige on dorsal margin. Hind-
wing uniformly grayish brown; fringe pale brown to beige. Frenu-
lum simple. Retinaculum with frenulum hook. Abdomen usually
appearing uniformly beige to beige brown, some specimens with
brown scales laterally and ventrally, available specimens greasy and
consequently difficult to characterize in color. Male genitalia (n = 5)
ToS
ee
SST
SEL
ae oR
BESET CEE SPS Ln a ES
60
1
Fics. 1-2. Adults of Haplochrois galapagosalis. 1. Female from Isabela (Volcan Darwin, 1000 m eley.). 2
Puerto Villamil).
(Figs. 3-6). Uncus rounded, flattened dorsoventrally but bulged and
shortly setose along apical and lateral margins, about 1/4th length of
tegumen. Gnathos with thin arms broadly curved, constricted before
apical elongate club bearing series of long, narrow, and curved spi-
nules in five transverse rows forming incomplete rings, except for
the most apical row. Tegumen rectangular, with parallel margins,
elongate, fused dorsally and ventrally for most of length (basal arms
very short), dorsal surface scaled, ventral surface spiculate, connect-
ing with vinculum by an elongate and subtriangular projection orig-
inating dorsomedially on each short arm. Transtilla narrow, well scle-
rotized, straight, with lateral partially articulated setose knobs
projecting posteriorly to apex of valva. Anellus forming a median,
crescent-shaped, and crested structure projected posteriorly. Juxta
narrow and poorly sclerotized, U-shaped to accommodate anellus,
with a few setae ventrally, with two short and sparsely setose arms
projecting posteroventrally. Valva short, flat, roughly semicircular,
somewhat angular, curved backward, with minute setae on most of
surface, dorsal and lateral margins with longer setae and long thin
scales forming a lateral fan on undissected specimens. Vinculum
very narrow, ventrally with long and thin saccus reaching middle of
segment VIL. Aedeagus of medium length, broadly curved in lateral
view, apex directed dorsally, without coecum penis, distal half dor-
sally with longitudinal separation between two lateral halves, apically
asymmetrical with only the lateral margins well sclerotized into nar-
row laterally flattened and apically rounded projections; the left pro-
jection longer, broader and coming out on the left side of the anel-
lus; manica surrounding aedeagus at median third, with short
sclerotized section apicodorsally; vesica without cornuti.
Female (n = 30). Forewing length: 5-7 mm. Antennal flagello-
meres simple, without noticeable setae, with erect scales on last
third of flagellum only. Frenulum with two acanthae. Retinaculum
with evenly spaced thin scales on Sc+R, and Radial stem. Female
genitalia (n = 4) (Figs. 7-9). Papillae anales laterally flattened, nar-
rowing slightly toward rounded apex, with setae of medium length
on lateral surface and very long setae around base; dorsal margin
straight; ventral margin convex. Posterior apophyses narrow, straight,
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
. Female from Isabela (+15 km N
reaching anterior margin of segment VIII. Segment VIII about as
wide as preceding segment, unsclerotized medioventrally. Anterior
apophyses short and narrow, directed medially at half right angle.
Ostium bursae in fold in middle of ventral membranous section of
segment VIII, without sclerotization. Middle of intersegmental
membrane VII-VIII with two small subtriangular plates variable in
width and degree of sclerotization between them. Ductus bursae
long, narrow, slightly variable in width and length, coiled twice, with
pair of very short sclerotized plates laterally at about 1/5th of length,
posterior to inception of ductus seminalis. Corpus bursae elongate,
about 1/3rd length of ductus bursae, without scobination, reaching
apical margin of abdominal segment II; single small signum a very
short inwardly directed spine on a small irregularly shaped plate.
Holotype 3d (CNC # 22679): 1—“ECU[ADOR]., GALAPAGOS/
Santiago, Central/ 700 m elev[ation]., 9.iv.1992/ M|ercury] Vapor]
Llamp], leg. B. Landry” [white, rectangular, printed in black ink].
2—“HOLOTYPE/ Haplochrois/ galapagosalis B. Landry/ CNC
#22679” [red, rectangular, hand written in black ink].
Paratypes: 16 3, 30 2 ECUADOR, Galapagos, MVL, leg. B
Landry. Floreana: Las Cuevas, 23.iv.1992 (1 4, slide BL 1254).
Punta Cormoran, 21.iv.1992 (1 2). Genovesa: Bahia Darwin,
10.iii. 1992 (3 2, slide BL 1256): 26.iii.1992 (1 5, 1 °). Isabela: Puerto
Villamil (8.5 let N), 11.iii.1989 (1 2); (ca. 15 km N), 25.v.1992 (2 2,
slide BL 1228). Tagus Cove, 13.v.1992 (1 4, slide BL 1253). Volcan
Darwin, 300 m elev., 15.v.1992 (1 2): 630 m elev., 16.v.1992 (1 2);
1000 m elev., 18.v.1992 (2 2); 1240 m elev., 19.v.1992 (3 9).
Marchena: 12.iii.1992 (1 4, 3 2 slide BL 1258): 23.iii.1992 (1 2).
Pinta: arid zone, 15.iii.1992 (1 2, slide BL 1255); 200 m elev.,
16.iii.1992 (1 2): 400 m elev., 18.iii.1992 (1 d, 2 2). San Cristébal: 4
km SE Puerto Baquerizo, 12.ii.1989 (1 d, CNC slide MIC 4677).
Base of Cerro Pelado, 22.ii.1989 (1 °). Santa Cruz: Charles Darwin
Research Station, arid zone, 19.1.1989 (1 4, slide JFL 604). 4 km N
Puerto Ayora, 20.i.1989 (1 3). Los Gemelos, 31.i.1989 (1 ?); 27 May
1992 (1.4). Tortuga Reserve (W Santa Rosa), 6.ii.1989 (4 6, 1 2),
Finca S. Devine, 17.iii.1989 (1 2); Finca Vilema (2 km W Bella
Vista), 1.iv.1992 (1 2). Santiago: Bahi Espumilla, 4.iv.1992 (1 d slide
VOLUME 55, NUMBER 2 61
Fics. 3-9. Genitalia of Haplochrois galapagosalis. 3-6. Males (scales = 0.5 mm). 3. Apical view, valvae flattened, aedeagus removed [slide '
BL 1257]. 4. Lateral view of whole genitalia except aedeagus [slide BL 1254]. 5. Aedeagus in lateral view [slide BL 1257]. 6. Aedeagus in dor-
sal view [slide 1254]. 7-9. Females. 7. Ventral view [from slides BL 1228, 1256, 1258] (scale = 0.5 mm). 8. Bursa in ventral view [slide BL 1228].
9. Plates on membrane below ostium bursae [slide BL 1258] (scale = 0.0625 mm).
62
BL 1257, 1 2). 200 m elev., 5.iv.1992 (1 2). Aguacate, 520 m elev.,
12.iv.1992 (1 3, 1 2).
The material collected in 1989 belongs to the Canadian National
Collection of Insects, Ottawa, Ontario, Canada (CNC). The material
collected in, 1992 will be distributed among the Muséum dhistoire
naturelle, Geneva, Switzerland (MHNG), The Natural History Mu-
seum, London, England (BMNH), the CNC (holotype), the Charles
Darwin Research Station, Santa Cruz Island, Galapagos, Ecuador
(ECCD), and the National Museum of Natural History, Washington,
D.C., U.S.A. (USNM).
Remarks. The type locality, Central, is a campsite, and so is Agua-
cate on Santiago. The nomenclature used for the description of the
male genitalia structures surrounding the aedeagus (the transtilla,
anellus and juxta) is tentative. No recent or older species descrip-
tions in this genus treat these structures in detail. Sinev (1999) men-
tions that the lobes of the gnathos bear setae, but I was not able to
see any evidence of sockets and so I have called these projections
spinules, in concordance with Hodges (1999). The two original spec-
imens of one of Meyrick’s species, H. catholica, are without ab-
domen and superficially similar to the Galapagos species. Resem-
blance in color pattern is common in Haplochrois and it is possible
that the two taxa are conspecific. However, the type locality of H.
catholica is Mallali, Guyana, where non endemic Galapagos species
are not typically encountered, unless they have a broad Neotropical
distribution.
Etymology. This species is named for the Ecuadorian archipel-
ago where it was collected.
Distribution and biology. The species was found on the Gala-
pagos Islands of Floreana, Genovesa, Isabela, Marchena, Pinta, San
Cristobal, Santa Cruz, and Santiago. I suspect that it may be distrib-
uted also on the other larger islands of the archipelago. Moths were
collected between January 19 and May 25. They were attracted to
light and found from sea level to 1240 m elevation on Volcan Dar-
win. The host plant is unknown.
ACKNOWLEDGMENTS
I express my sincere gratitude to the authorities of the Galapagos
National Park and those of the Charles Darwin Research Station for
allowing fieldwork and for logistical support in 1989 and 1992. The
fieldwork and other costs were partially supported by an operating
grant to Stewart B. Peck, Carleton University, Ottawa, from the Nat-
ural Sciences and Engineering Research Council of Canada, for
field research on arthropod evolution. Without the following com-
panions, some of my work in the Galapagos would have been im-
possible, or at least much less pleasant, and I thank them sincerely:
Joyce Cook, Moraima Inca, Ricardo Palma, S. B. Peck, Bradley J.
Sinclair, and Eduardo Vilema. I thank curators Kevin Tuck (BMNH),
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
David Adamski (USNM), and Richard Hoebeke, Comell University
Insect Collection, Ithaca, New York, U.S.A., for their help in the ex-
amination of critical specimens. I am grateful to Ronald W. Hodges
(formerly at USNM) for a preliminary determination of the insect
described herein. David Adamski and Ronald Hodges also provided
useful comments on the manuscript. Finally, I am greatly indebted
to Jean-Francois Landry, Agriculture and Agrifood Canada, Ottawa,
for facilitating lab work, for his help in processing the images, and
for mentioning critical literature.
LITERATURE CITED
BOTTIMER, L. J. 1926. Notes on some Lepidoptera from Eastern
Texas. Jour. Agric. Res. 33:797-819.
CLARKE, J. F. G. 1965. Catalogue of the type specimens of mi-
crolepidoptera in the British Museum (Natural History) de-
scribed by Edward Meyrick. Vol. V. London, Trustees of the
British Museum (Natural History). 581 pp.
Hopces, R. W. 1983. Agonoxenidae, p. 17. In Hodges, R. W. et al.
(eds.), Check list of the Lepidoptera of America north of Mex-
ico. E. W. Classey Ltd. and the Wedge Entomol. Res. Found.,
London. xxiv + 284 pp.
. 1997. A new Agonoxenine moth damaging Araucaria
araucana needles in Western Argentina and notes on the
Neotropical Agonoxenine fauna (Lepidoptera: Gelechioidea:
Elachistidae). Proc. Entomol. Soc. Wash. 99:267-278.
. 1999. The Gelechioidea, pp. 131-158. In Kristensen, N. P.
(ed.), Handbook of zoology, Lepidoptera, moths and butterflies.
Vol. 1. Evolution, systematics, and biogeography. Walter de
Gruyter, Berlin & New York. x + 491 pp.
KusNEZOy, N. J. 1915. Description of Parametriotes theae, gen. n.,
sp. n. (Lepidoptera, Tineidae), a new enemy of the tea bush in
Transcaucasia. Ent. Obozr. 15:627-652.
Lanpry, B. 2001. The Cosmopterigidae (Lepidoptera) of the Gala-
pagos Islands, Ecuador. Rev. suisse zool. 108:513-539.
MEyRICK, E. 1897. Descriptions of Australian microlepidoptera.
Proc. Linnean Soc. N.S. Wales 22:297—-435.
REBEL, H. 1902. Lepidopteren aus Morea gesammelt von Herm
Martin Holtz im Jahre 1901. Berl. ent. Z. 47:83-110.
RIEDL, T. 1969. Matériaux pour la connaissance des Momphidae
paléarctiques (Lepidoptera). Partie IX. Revue des Momphidae
européennes, y compris quelques espéces d'Afrique du Nord et
du Proche-Orient. Polskie pismo ent. 39:635-919.
SINEY, S. Y. 1999. Notes on synonymy of narrow-winged moths
(Lepidoptera: Agonoxenidae, Cosmopterigidae, Momphidae) of
the Palaearctic. Ent. Rev. 79:30-39.
Received 20 September 2000; revised and accepted 6 August 2001.
Journal of the Lepidopterists’ Society
55(2), 2001, 63-68
MORTALITY OF LEPIDOPTERA ALONG ROADWAYS IN CENTRAL ILLINOIS
DUANE D. MCKENNA
KATHERINE M. MCKENNA
Department of Organismic and Evolutionary Biology,
Harvard University, 26 Oxford Street,
Cambridge, Massachusetts 02138, USA
STEPHEN B. MALCOM
Department of Biological Sciences,
Western Michigan University, 3151 Wood Hall,
1903 W. Michigan Avenue, Kalamazoo, Michigan 49008, USA
May R. BERENBAUM
Department of Entomology, University of Illinois,
320 Morrill Hall, 505 S. Goodwin,
Urbana, Illinois 61801, USA
ABSTRACT. We conducted this study to investigate the magnitude of roadway mortality of Lepidoptera in central Ilinois. To quantify the
number and kinds of Lepidoptera killed along roadways, dead adult Lepidoptera were collected, identified, and counted from along 13 roadside
transects in the vicinity of Champaign/Urbana, Illinois, with collections occurring weekly on each transect for six weeks. During the six weeks
of this study, 1824 presumably road-killed Lepidoptera were collected. At traffic rates of 1000, 13,500, and 19,700 vehicles per day, more Lep-
idoptera were collected per 100 m than at other traffic rates. A peak in monarch butterfly mortality may coincide with the timing of their annual
migration through the area. Based on these data, the number of Lepidoptera killed along roadways for the entire state of Illinois during one
week was estimated at more than 20,000,000 individuals. The number of monarch butterflies killed may have exceeded 500,000 individuals. Our
results suggest that increases in traffic rate and speed limit may to a certain extent increase mortality.
Additional key words: Butterflies, Danaus plexippus, traffic.
Although roadway traffic is known to affect popula-
tion densities of amphibians (Fahrig et al. 1995),
snakes (Bernardino & Dalrymple 1992), koalas (Can-
field 1991), wolves (Mech 1989), turkeys (Holbrook &
Vaughan 1985), badgers (Davies et al. 1987), and other
vertebrates (Lalo 1987, Putman 1997), practically
nothing is known about the impact that roadways have
on invertebrates (Seibert & Conover 1991). In fact, a
recent book on butterfly conservation (New 1997)
makes no mention of the subject, and roads are men-
tioned only briefly in two other recent books on insect
conservation, as barriers to butterfly movement
(Samways 1994, Pullin 1995). Samways (1994) states
“Roads are line corridors that can cause high mortality
where traffic volume is high. In 1989, 100 m of Ten-
nessee roadside was a graveyard for over 120 traffic-
killed butterflies” (Samways 1994:117). Other than
these sources, the best information on road mortality
of butterflies is by Munguira and Thomas (1992) in
England. They found that roads were not a serious
barrier to butterfly movement, but that vehicles killed
up to 7% of adult butterflies from some populations.
Illinois has 2050 miles of interstate, 276 miles of toll
road, 14,892 miles of highway, and 120,782 miles of
county, municipal, and other roads (Illinois Depart-
ment of Transportation pers. com. 1998). Although this
comprises the third largest state highway system in the
United States, nothing is known about the magnitude
of lepidopteran mortality along Illinois roadways.
This study was conducted to investigate the magni-
tude of roadway mortality of Lepidoptera in central
Illinois. To quantify the number and kinds of Lepi-
doptera killed, dead adult Lepidoptera were collected,
counted, and identified from along 13 roadside tran-
sects in the vicinity of Champaign/Urbana 40°2’N,
88°17’'W, Champaign Co., Illinois, with collections oc-
curring weekly on each transect for six weeks. This
study is the first to document systematically the mag-
nitude of roadway mortality of Lepidoptera anywhere
in the United States.
MATERIALS AND METHODS
Eleven roadside transects were chosen at the outset
of this study to represent different rates of traffic and
different roadway types (Table 1). Traffic rates ranged
from 0-26,000 vehicles per 24 hour period (Illinois
Department of Transportation (IDOT) pers. com.). To
facilitate comparisons, traffic rates above 500 vehicles
per day were rounded to the nearest 100. Each road-
side transect fit into one of the following general plant
community types: remnant prairie, agriculture, or
woodland. Roadways were classified into four types:
64.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
TABLE 1. Transect Characteristics.
Roadside Speed Vehicles/24
Transect Transect Roadway community Transect limit hours **
number name type type* length (m) (mph) (#Vehicles)
1 Highway 150 #1 Highway Prairie/Ag 150 Do 1,000
2 Highway 150 #2 Highway Prairie/Ag 250. 55 1,000
3 Highway 150 #3 Highway Prairie/Ag 180 55 1,000
4 Cunningham #1 Divided Highway Ag 180 55 13,500
5 Cunningham #2 Divided Highway Ag 250 55 13,500
6 Cunningham #3 Divided Highway Ag 180 55 13,500
i I-74 Interstate Ag 200 65 26,000
8 M-57 Interstate Ag 200 65 19,700
9 Trelease Woods Country Road Old Field/Woodland 160 35 150
10 Brownfield Woods Country Road Woodland 200 0-35*** 1000
11 Airport Grass Control | N/A (Mowed Airfield) | Mowed Grass 100 N/A 0
12 Country Road Control Country Road Ag 200 O0-45*** 150
13 Highway 150 Paved Country Road Prairie/Ag 600 45 50
*Ag = Agriculture.
** The number of vehicles per day (24 hour period) was obtained through the Illinois Division of Highways.
*%* The range of speeds given are those typically observed for vehicles accelerating from the stop signs at the beginning of these transects.
paved country road, highway, divided highway, and in-
terstate. One additional transect was chosen to control
for the effects of roadside mowing. A second addi-
tional transect was chosen to quantify the baseline
mortality in mowed grass, a habitat resembling the
mowed roadsides of all transects.
Two transects were abandoned one week into the
study due to road construction. These transects were
replaced with two new transects, and two interstate
transects were also added, for a total of 13 transects
(Table 1). The nine original transects were first sam-
pled on 25 August 1998. The two replacement tran-
sects were first collected on 2 September 1998, and
the two interstate transects were first collected on 9
September 1998. We collected along transects weekly
until 19 October 1998. Over the two weeks after that
date, no dead Lepidoptera were found and the sam-
pling was terminated.
Once every seven days each transect was walked and
road-killed Lepidoptera were collected into a plastic
bag. All 13 transects were collected over a two-day pe-
riod each week. Transects with a median were searched
on all sides of the road, including both sides of the me-
dian. As a safety precaution, the medians of interstate
transects were not sampled. Both sides of the road
were walked against the flow of traffic. Lepidoptera
were collected that lay dead within two meters of the
edge of the road. This usually included the shoulder
and about one meter of “ditch”. Lepidoptera lying on the
roadway itself were not collected, but fewer than 10 such
individuals were noticed throughout the entire study.
Transect length varied from 100 to 610 m, with a
mean of 219 m and standard deviation of 121 m (Table
1). A total of 2850 m of transect was sampled during
weeks when all 13 transects were sampled. The
Thomasboro, Illinois office of the Illinois Division of
Highways reported that transect locations received sim-
ilar roadside maintenance, but on different days. Typical
maintenance included mowing and trash removal.
After collection, the Lepidoptera were sorted by
species or species-group and were counted. They were
sorted into six taxonomic groups:
1. Hesperiidae, mostly Epargyreus clarus (silver-
spotted skipper).
2. Lycaenidae, which were not identified below the
family level.
3. Moths, mostly Arctiidae and Noctuidae, grouped to-
gether because too few individuals from most fami-
lies were collected to warrant separate treatment.
4. Nymphalidae, including Danaus plexippus (mon-
arch), Euptoieta claudia (variegated fritillary),
Libytheana carineta (American snout butterfly), Li-
menitis archippus (viceroy), Limenitis arthemis
astyanax (red-spotted purple), Phyciodes tharos
(pearl crescent), Polygonia interrogationis (question
mark), Junonia coenia (common buckeye), and
Vanessa cardui (painted lady).
5. Papilionidae, represented in the roadside transects
only by Papilio polyxenes (black swallowtail).
6. Pieridae, including Colias eurytheme (yellow sulfur)
and Colias philodice (clouded sulfur), which due to
hybridization and difficulty in separation into
species were grouped together and called the C. eu-
rytheme/C. philodice hybrid complex. Pieridae also
included Eurema lisa (little yellow) and Pieris rapae
(cabbage white).
VOLUME 55, NUMBER 2
65
| LSIO
> 8
[o) |
ro
Sess
°
fe
ems 4
o |
a 4
hae
=
Q 2
o. 214
a 1
a 69
; bu | fesacea SASL te 52S I el lal
Pieridae Nymphalidae Moths Hesperiidae Papillionidae Lycaenidae
Group
Fic. 1. Proportion of total mortality for each group of Lepidoptera studied. The total number of individuals collected from each group is
given above each column. N = 1824.
Data were analyzed using analysis of variance
(ANOVA) techniques. Weeks one and two were ex-
cluded from the analyses because they included data
for only a subset of the 13 transects. Results were con-
sidered significant at p $ .05.
Calculations of theoretical statewide mortality were
made by multiplying the number of meters of roadway
in Illinois of each of the roadway “types” (see above)
by the number of butterflies killed per meter of road-
side transect for each roadway type during the week of
9 September 1998 (IDOT pers. com.).
RESULTS
During the six weeks of this study, 1824 Lepi-
doptera were collected from along the 13 roadside
Number of Lepidoptera Killed per 100 m
transects, including 1510 Pieridae, 214 Nymphalidae,
69 moths, 15 Hesperiidae, 13 Papilionidae, and 3 Ly-
caenidae (Fig. 1). Insects belonging to orders other
than Lepidoptera were only infrequently encountered
and are not reported here. The greatest number of
Lepidoptera killed per 100 m of transect occurred
during week 1 (Fig. 2). Mortality decreased there-
after, except for a slight increase at week 4. Not in-
cluded in the figures are the last two weeks of transect
samples during which no dead Lepidoptera were
found.
The first week of collection resulted in the two high-
est mortality rates per 100 m recorded for any of the
traffic rates during the study (Table 2). At 1000, and
13,500 vehicles per day, 51.28 and 49.34 individuals
Week
Fic. 2. Number of Lepidoptera sampled per 100 m during the six-week study.
66
TABLE 2. Lepidoptera sampled (number/100 m) over six weeks
at each of nine traffic rates (number of vehicles/day). An “X” indi-
cates that no data were collected.
Vehicles/Day
Week 0 50 150 1000 13,500 19,700 26,000
1 0.17 O 51.30 49.30 xX xX
9) 0 0 6.40 25.0 19.50 xe xX
3 0 0 3.60 4.50 27.70 25.50 1.50
4 0 0 2.80 14.70 27.90 23.0 1.50
5 0 0 0.280 1.90 10.50 13.50 50
6 0 0) 0 1.30 3.30 8.50 2.50
respectively, were collected per 100 m of transect. At a
traffic rate of 13,500 vehicles per day, more Lepi-
doptera were routinely sampled each week per 100 m
than at any other traffic rate (Table 2). The lowest
numbers were consistently recorded from transects
with traffic rates of 150 vehicles per day and at the
highest traffic rate, 26,000 vehicles per day. The num-
ber of Lepidoptera collected per 100 m of roadway in-
creased from 0.03 at 50 vehicles per day to 23.03 at
13,500 vehicles per day (Fig. 3). At 19,700 vehicles per
day 17.63 individuals were sampled per 100 m of tran-
sect, and at 26,000 vehicles per day only 1.5 individu-
als were sampled per 100 m of transect. Thus, at a rate
of 13,500 vehicles per day, observed mortality was
greatest, and at the highest traffic rate mortality was
much lower. The mean number of butterflies collected
per 100 m along transects with a traffic rate of 26,000
vehicles per day was not significantly different from
that along transects with traffic rates of 19,700 (p =
0.13), 1,000 (p = 0.67), and 150 (p = 0.19) vehicles per
day. At a traffic rate of 13,500 vehicles per day, signifi-
cantly more Lepidoptera were collected per 100 m
than at 26,000 vehicles per day (p = 0.01).
2] 0 50 150
Number of Lepidoptera Killed Per 100 m
c
2
oo
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
The C. eurytheme/C. philodice hybrid complex was
most frequently collected. Over the course of the
study, 1492 individuals were found dead along tran-
sects. Monarch butterflies, D. plexippus, were the next
most abundant species found dead on transects; 99
were collected, including 55 males, 31 females, and 13
that were not identified to sex due to damaged or de-
tached wings and abdomens. The number of mon-
archs collected varied greatly from transect to transect,
and less so from week to week (Table 3 and Fig. 4).
From 0.013 to 0.119 monarch butterflies were killed
per 100 m of transect each week. The most monarch
butterflies collected per 100 m from a transect was 6.5
at a traffic rate of 19,700 vehicles per day during week
four. The second and third greatest mortality, 2.13 and
1.64 individuals per 100 m respectively, were observed
at a traffic rate of 13,500 vehicles per day, also during
week four. The mean number of monarch butterflies
collected per 100 m during week four was significantly
greater than during weeks three, five, and six (p =
0.02). Relatively few individuals of other species of
Lepidoptera were collected.
The red-spotted purple, L. arthemis, and viceroy, L.
archippus, were found only along transects with wood-
land or prairie roadsides. The little yellow E. lisa was
found only along transects with prairie roadsides.
Here, larvae were observed feeding on Cassia (a
legume). We collected both day-flying and night-flying
moths. No rare Lepidoptera were collected.
Using the roadway types and mortality statistics of
the study transects to estimate rates of mortality for
different roadway types in Illinois, we estimated the
potential number of Lepidoptera killed for the entire
state during the seven days prior to transect collection
on week three; 9 September 1999. According to our
1,000
13,500 19,700 26,000
Vehicles Per Day
Number of Lepidoptera sampled per 100 m at each traffic rate. Bars indicate standard deviations.
VOLUME 55, NUMBER 2
TABLE 3. Monarch butterflies killed (number/100 m) over six
weeks at each of seven traffic rates (number of vehicles/day). An “X”
indicates that no data was collected.
Traffic rate
Mowed a ee ee ee
Week _ grass 50 150 1000 13,500 19,700 26,000
1 0 0 0.56 0.90 1.64 Xx xX
2 0 0 0.83 1.28 0.82 Xx xX
3 0 0 0 0.38 1.15 1.00 1.00
4 0 0 0 0.90 213 650 0.50
5 0 0 0 0.77 0.33 150 O
6 0 0 0 0.77 0 0.50 ‘1.00
estimates, the number of Lepidoptera killed along in-
terstates in Illinois during this week could have been
more than 500,000, the number killed along highways
could have been more than 5,000,000, and the number
of Lepidoptera killed along other roads could have
been more than 15,000,000. In total, the number of
Lepidoptera killed by automobiles was estimated at
more than 20,000,000 individuals in this seven—day pe-
riod.
Using similar methods, we estimated the potential
number of monarch butterflies killed during one week.
According to our estimates, the number of monarchs
killed along interstates in Illinois in one week during
this study may have been more than 500,000.
DISCUSSION
Quantifying definitively the impact of automobile
traffic on Lepidoptera is operationally challenging for
many reasons. The small number of Lycaenidae in the
samples for example, may result from the timing of our
study. The collection methods used may have also re-
sulted in a significant undercount of dark-colored and
0.3
0.25 |
0.2 4
0.15 5
0.1
Number of Monarchs Killed per 100 m
small moths, as they were difficult to see lying on the
ground. Moreover, small Lepidoptera of all kinds may
stay attached to the automobile that hits them (DDM
pers. obs.). As well, ants and other insects, birds, and
rodents were observed removing the remains of small
dead insects from the transect roadsides. Weathering
and mowing were observed to disintegrate a few Lep-
idoptera specimens before they could be counted.
Thus, the numbers we report here should be consid-
ered a minimum estimate of mortality for the area
studied; more precise measurements of mortality
await further studies.
The numbers of individuals killed per 100 m on the
first sample date in Table 2, Fig. 2, and Fig. 4, are not
likely to reflect accurately actual mortality along the
transects during the week prior to collection because
these samples include all dead Lepidoptera that had
accumulated and persisted along the transects prior to
initiation of the study.
Somewhat surprising is the result that, at highest
traffic rates, mortality declines. There are several pos-
sible explanations for this finding. At speeds around 55
mph or greater, Lepidoptera were seen to be caught in
a “wind current” going over the roof of the car, with
the result that they were “catapulted” over the car in-
stead of colliding into it. It is not clear if this catapult
effect may have resulted in fewer dead butterflies at
sites with the highest traffic rates and speed limits
compared to sites with lower traffic rates and speed
limits. A further complication is the lack of collections
from expressway medians. The length of expressway
transects and the mortality data do not incorporate this
sampling anomaly. The number of lanes of traffic may
also be an important factor. An alternative explanation
is that the observed decrease in numbers of dead Lep-
Week
Fic. 4. Mean number of monarch butterflies sampled per 100 m during the six-week study. Bars indicate standard deviations. Week one col-
lections were started on September 25.
i
68
idoptera when traffic volume and speed limit were
relatively high (Table 2 and Fig. 3) may be due to a
relatively high per capita morality rate at these sites re-
sulting in decreased population density and the obser-
vation of fewer dead Lepidoptera. While we did not
test this hypothesis, it is an interesting possibility be-
cause it suggests population-level effects of roadway
mortality.
The peak in mortality of monarch butterflies ob-
served on or about week 4 (September 16) may have
been due to their southward migration (Fig. 2). Mi-
grating monarchs usually fly high enough to avoid col-
lision with vehicles, but during mid-morning, they
generally fly lower to the ground (Orley Taylor pers.
comm. ). They also fly low to the ground during windy
weather such as that which prevailed during the study
period (DDM pers. obs.).
Many more male monarchs were found in the sam-
ples than females. Along Highway 150, where four
transects were located, male monarchs were observed
chasing other butterflies for distances of up to 100 m,
often across the roadway. This behavior may account
for the apparent overrepresentation of males in the
samples.
More black swallowtails than monarchs were ob-
served flying at highway sites, but more monarchs
were found dead. Along most highway and expressway
transects, considerable numbers of stems of whorled
milkweed (Asclepias verticillata, Asclepiadaceae) were
observed within two m of the roadway edge. These
plants were observed to be an important larval host
plant and source of nectar for adult monarch butter-
flies at these sites. Black swallowtails were observed to
use whorled milkweed as a nectar plant, but were
chased away from the extensive clonal growths of
whorled milkweed along roadsides by monarch butter-
flies. Black swallowtails generally frequented areas fur-
ther removed from the roadside than monarch butter-
flies where nectar plants for adults such as thistles and
clovers (Cirsium spp., Asteraceae and Trifolium spp.,
Fabaceae) and larval host plants such as wild parsnip
(Pastinaca sativa, Apiaceae) and Queen Anne’s lace
(Daucus carota, Apiaceae) were most abundant (DDM
pers. obs.). These behavioral differences may account
for the observation of fewer dead black swallowtails.
Regardless of the practicality of making estimates of
statewide mortality from this small data set, it is appar-
ent from this study that roadways kill significant num-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
bers of adult Lepidoptera in central Illinois. Unfortu-
nately, estimates of adult lepidopteran mortality
caused by other factors in this region are generally
lacking. The implications of roadways, roadsides, and
traffic rates for lepidopteran mortality and populations
are unclear, but evidence suggests that increases in
traffic rate and speed limit may increase mortality to a
certain extent and may have a detrimental effect on
some populations. Future studies should explicitly ad-
dress these questions. Studies are also needed from
other regions and throughout the season in order to
determine overall impacts of traffic on Lepidoptera
and as well to provide baseline information that may
be helpful in designing programs for reducing this
mortality as it affects threatened or endangered
species.
ACKNOWLEDGMENTS
We would like to thank the undergraduates who assisted the au-
thors with the task of sorting and counting dead butterflies. Thanks
to Dr. Arthur Zangerl for reading through this manuscript. We
would also like to thank the Illinois Department of Transportation
for access to several of the study sites, and two anonymous reviewers
and the editor of this journal for their comments on an earlier ver-
sion of this paper. This work was supported in part by NSF DEB99-
03867.
LITERATURE CITED
BERNARDINO, F. S. & G. H. DALRYMPLE. 1992. Seasonal activity
and road mortality of the snakes of the Pahay-okee wetlands of
Everglades National Park, USA. Biol. Conserv. 62:71-75.
CANFIELD, P. J. 1991. A survey of koala road kills in New South
Wales. J. Wildl. Dis. 27:657-660.
Davies, J. M. 1987. Seasonal distribution of road kills in the
European badger (Meles meles). J. Zool. 211:525-529.
Fannic, L., J. H. PEDLAR & S. Pope. 1995. Effect of road traffic on
amphibian density. Biol. Conserv. 73:177-182.
Howsrook, H. T. & M. R. VAUGHAN. 1985. Influence of roads on
turkey mortality. J. Wildl. Manage. 49:611-614.
Lato, J. 1987. The problem of road kill. Am. Forests 93:50-52.
Mecu, D. 1989. Wolf population survival in an area of high road
density. Am. Mid]. Nat. 121:387—-389.
Muncuira, M. L. & J. A. THomas. 1992. Use of road verges by
butterfly and burmet populations, and the effects of roads on
adult dispersal and mortality. J. Appl. Ecol. 29:316—329.
New, T. R. 1997. Butterfly conservation. Oxford University Press,
Oxford. 248 pp.
Putin, A. §. 1995. Ecology and conservation of butterflies.
Chapman & Hall, London. 363 pp.
PurmaNn, R. J. 1997. Deer and road traffic accidents: options for
management. J. Environ. Manage. 51:43—57.
Samways, M. J. 1994. Insect conservation biology. Chapman &
Hall, London. 358 pp.
SEIBERT, H. C. & J. H. Conover. 1991. Mortality of vertebrates
and invertebrates on an Athens County, Ohio, highway. Ohio J.
Sci. 91:163-166.
Received 11 October 2000; revised and accepted 28 August 2001.
Journal of the Lepidopterists’ Society
55(2), 2001, 69-73
ONTOGENY OF DEFENSE AND ADAPTIVE COLORATION IN LARVAE OF THE COMMA
BUTTERFLY, POLYGONIA C-ALBUM (NYMPHALIDAE)
SOREN Ny LIn!, GABRIELLA GAMBERALE-STILLE, AND BIRGITTA S. TULLBERG
Department of Zoology,
Stockholm University,
SE-106 91 Stockholm, Sweden
ABSTRACT. In many butterfly species early and late larval instars differ in coloration. The first three larval instars in the comma butter-
fly, Polygonia c-album, have coloration that, to the human eye, appears to be disruptively cryptic. The last two instars, which are defended by
strong branching spines, are instead strikingly colored in white, black, and orange. It has been suggested that this coloration is also cryptic, mim-
icking bird droppings. We test the idea that this coloration is instead (or simultaneously) aposematic, using young chickens as models of bird
predators. Some chicks readily ate third-instar larvae, both initially and after having tasted them, suggesting that no chemical defense is present
in comma larvae. The chicks also ate dead fifth-instar larvae from which the spines had been removed. Chicks initially attacked fifth-instar lar-
vae with intact spines, but learned to avoid them already after the first attack. This suggests an aposematic function of the coloration, which can
have evolved by individual selection in this and many other nymphalid species with spiny larvae, because larvae were not harmed during the
learning process. The evolutionary causes of the ontogenetic shift in defense tactics are discussed.
Additional key words: aposematism, mimicry, life history, predation, adaptive function.
Adaptive coloration in animals has been a very active
research field in evolutionary biology over the years
(e.g., Poulton 1890, Cott 1940, Kettlewell 1973, Sillén-
Tullberg 1988, Malcolm 1990), and one in which the
Lepidoptera have always featured prominently as
model species. Adaptive coloration includes crypsis
(helping to avoid detection by other animals), sig-
nalling, and thermoregulation. Crypsis can be achieved
by having colors similar to the background, or by hav-
ing mosaic patterns of spots or stripes that break up
the contour and surface area of the animal (called “dis-
ruptive crypsis”; Cott 1940, Edmunds 1990). Signals can
be of several types, including “aposematism,” defined as
conspicuous colors advertising unprofitability to preda-
tors (Guilford 1990a, b). A phenomenon that may in-
clude aspects of both signalling and crypsis is “mim-
icry,’ a term used for organisms that adaptively
resemble another species, as in the cases of Miillerian
and Batesian mimicry (Malcolm 1990). In the former
case, unpalatable aposematic organisms mimic each
other. In the latter case, a palatable organism mimics
an aposematic one, capitalizing on its defenses. Some-
times the term mimicry is extended to cases when or-
ganisms mimic the shape and colors of objects or im-
mobile organisms for the purpose of crypsis, for
example as in stick insects (Edmunds 1990). Finally,
variation in coloration can be due to different require-
ments for thermoregulation rather than predator
avoidance (e.g., Shapiro 1976, Kingsolver & Watt
1983). For all of these types of (presumably) adaptive
coloration, it is probably fair to say that the function is
often assumed, but seldom tested.
"Email: Soren.Nylin@zoologi.su.se Tel: +46-8-164044 Fax:
+46-8-167715.
The coloration of the first three larval instars in the
comma butterfly, Polygonia c-album L. (Nymphali-
dae), appears to the human eye to be disruptively cryp-
tic (Fig. 1A), at least when seen against a naturally var-
iegated background of light and shadow. The last two
instars, which are defended by strong branching spines,
are much more strikingly colored in white, black and
orange (Fig. 1B). The same type of coloration is present
in the probable sister species to P. c-album, the Nearc-
tic P. faunus (Scudder 1889, Scott 1986). It has been
suggested that this type of coloration is also cryptic, the
large continuous white area supposedly mimicking bird
droppings (Thomas 1986). Resemblance to bird drop-
pings is seen in many butterfly larvae, e.g., in Limenitis
and Papilio (Scott 1986), although to our knowledge
the function has never been tested.
Late instar larvae of other Polygonia species (e.g., P.
interrogationis, P. comma, P. satyrus, and P. c-aureum)
and the related Kaniska canace are similarly colored in
conspicuous white, black and orange, but with no con-
tinuous white areas (Scott 1986, Teshirogi 1990). This
suggests the possibility that the coloration is apose-
matic in these species, and then perhaps in P. c-album
and P. faunus as well. Here, we study the function of
the coloration of late instar larvae of P. c-album using
young chickens as models of bird predators. For com-
parison and as a control for effects of novel food, we
also tested the chicks with third-instar larvae.
MATERIALS AND METHODS
Females of P. c-album were captured near Akers-
berga, north of Stockholm, Sweden. Eggs were ob-
tained in flight cages where the host plant Urtica dioica
(Urticaceae) was present. Larvae were reared on this
plant in the laboratory. Several asynchronous rearings
70
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fic. 1. Larvae of Polygonia c-album on leaves of Urtica dioica. A. Third-instar larva. B. Fifth-instar larva.
were made, so that third- and fifth-instar larvae were
available for trial simultaneously.
Third-instar larvae. In this instar, the larvae lack
large areas of any color except for the background
color, black. Small spines are present, and towards the
end of this instar the spines have a yellowish color on
the foremost part of the body and a whitish color to-
wards the rear (as seen in Fig. 1A). The impression to
the human eye is not that of a striking coloration but
rather a pattern that could function disruptively
against a variegated background.
Fifth-instar larvae. In this instar, the foremost
part of the body is colored orange, and the rear part
is continuously colored white. The rest of the body
is black with orange markings (Fig. 1B). Large,
chitinous, branching spines are present on the back
and sides of the body, and they are colored orange,
white or black, according to position. Larvae are sev-
eral centimeters in length (Figs. 1A and B are approx-
imately to scale).
Chicks. We used domestic chicks (Gallus gallus do-
mesticus) as predators. The chicks had not eaten when
they arrived from the hatchery at an age of less than 20
h. Batches of 30-40 chicks were housed in cages (100
cm X 55 cm X 20 cm) with wooden sides, steel-net
floor, and a roof made partly of wood and partly of
chicken wire. The cages were heated with 60 W car-
bon light bulbs and the floors were covered with saw-
dust. Chicks were fed chick starter crumbs and water,
and at least from their second day on they were also
fed live mealworms (Tenebrio molitor).
Experiments. The experiments took place in an
arena of the same kind of cage in which chicks were
housed. Part of the cage was screened off, leaving a
testing floor of 30 em x 55 cm. Experiments took place
when the chicks were about three days old. The chicks
VOLUME 55, NUMBER 2
7
|
~ — |
— -3d instar ||
|| 5th instar |
Attacks (%)
3
1 2 3
Trial #
Fic. 2. Results from trials with domestic chicks presented with
third-instar (dashed line) or fifth-instar (solid line) larvae of Polygo-
nia c-album. The same pair of chicks was tested on several occasions
during one experimental day (subsequent trials from left to right).
Both young and old larvae were attacked in initial trials, but fifth-in-
star larvae were avoided in the next trial. N = 10 pairs of chicks in
trial #1 and 2 for both larval instars, N = 7 pairs of chicks in trial #3
with young larvae (see text).
were tested in pairs, because single chicks become dis-
tressed and do not feed normally (Gamberale & Tull-
berg 1996, and references therein). One of the chicks
was fed as many mealworms as it would eat before a
trial, which made it satiated and uninterested in feed-
ing during the trial. This chick was used as a compan-
ion to the experimental chick. We used the same com-
panion chick in all trials.
Prey was presented in a petri dish with transparent
bottom, i.e., chicks saw the prey against a background
of sawdust. Before each trial, the chicks were pre-
sented two mealworms in the petri dish. This was done
to show the place where prey was displayed, and as a
check before each trial that the chicks were in fact in-
terested in insect prey. The chicks were then pre-
sented one mealworm and one larva of P. c-album. We
collected data concerning chick attack behavior and
the mortality of the attacked insects. Chicks were ex-
posed to the prey throughout the trial and allowed to
make as many attacks as they wished. A trial was ended
when the P. c-album larva had been eaten, or after 60
seconds if the chick did not peck on the P. c-album
larva at all. If it did, we waited another 60 seconds to
see if the larva was eaten.
Twenty chicks (other than the companion chick)
were used in total, originating from two separate
batches. Half of them were tested with third-instar lar-
vae and the other half with fifth-instar larvae, using
chicks from both batches in both types of experiments.
All chicks were tested twice with the same type of
larva. In addition, to test for the presence of any aver-
sion learning against third-instar larvae, seven chicks
(of the same batch) were also tested with such larvae a
third time. Finally, four of these chicks, that had eaten
or attacked third-instar larvae on all three occasions,
were tested a fourth time. All trial runs were per-
formed consecutively on the same day, within five
hours. We used Fisher's exact test for statistical com-
parisons.
A follow-up experiment was performed next season,
with new larval stock and a new batch of chicks, this
time they were one week old. Five pairs of chicks were
presented with a spineless fifth-instar larva together
with a mealworm in a petri dish. The larvae had been
killed by freezing and their spines had been cut off.
RESULTS
General observations. The two mealworms pre-
sented at the start of each trial were always eaten rapidly,
demonstrating that the chicks were very interested in
insect prey. In all trials the chicks were initially more
interested in attacking the mealworms than the
comma larvae when subsequently given a choice, not
surprisingly since this was a food type with which they
had previous experience. The third-instar P. c-album
larvae might also have been somewhat hard to see
against the variegated background of sawdust. How-
ever, the experimental chicks were curious and in 19
out of 20 initial trials (both age classes combined)
eventually pecked at the comma larvae.
Third-instar larvae. In the initial trials all 10 ex-
perimental chicks attacked the third-instar comma lar-
vae (Fig. 2). In five of these cases the larva was also
eaten (after a time span of, respectively, 19, 30, 35, 45,
and 45 seconds). In the next trial the same chicks at-
tacked in nine out of ten cases (decrease in attack fre-
quency not statistically significant: Fisher's exact test),
and ate the larva in two cases (both after 45 seconds).
Both of these cases involved chicks which had also
eaten the comma larva in the previous experiment.
As explained in Materials and Methods one group of
chicks was tested a third time, with attacks by all seven
experimental chicks (Fig. 2) and the larvae being eaten
by the same two chicks (after 25 and 120 seconds).
When the two chicks that had eaten the larva on all
three occasions were tested anew they did so again,
this time within 20 seconds after exposure. When two
chicks that had attacked but not eaten (on all three oc-
casions) were tested again they repeated this behavior.
The last trials with this group took place 2 hours and
AO minutes after the start of the experiment.
In all cases when the larva was not eaten it was still
alive after the experiment (despite having been pecked
at), but on one occasion it was visibly damaged, and for
this reason it was killed by us.
Fifth-instar larvae. The older larvae were never
eaten by the chicks. In the initial trials attacks took
~l
Ns)
place in nine out of ten cases (Fig. 2). They were in the
form of pecks, in three cases followed by the chicks
lifting the larva and dropping it. In the next trial, which
took place less than an hour after the first, the chicks
behaved very differently. There were no attacks in the
form of pecks (Fig. 2). The decrease in attack fre-
quency between trials is statistically significant
(Fisher's exact test: two-tailed p < 0.05, one-tailed p <
0.01). In all cases except one (N = 10) the experimen-
tal chick inspected the comma larvae closely, some-
times for periods up to 15 seconds. In two cases at-
tacks were initiated but terminated before contact.
Larvae were alive and apparently undamaged after all
experiments, and they were returned to their host
plants where they soon continued to feed.
Fifth-instar larvae without spines. In this exper-
iment larvae from which the spines had been removed
were presented to the chicks. The first pair of chicks
did not eat the larva, but they showed no aversion and
handled the larva throughout the experiment. This
larva was presented also to the next pair of chicks, and
it was then eaten within 10 seconds. The three addi-
tional pairs of chicks also ate the spineless larvae
within seconds.
DISCUSSION
Five out of ten chicks in initial trials ate third-instar
larvae, but only two chicks continued to eat them in
subsequent trials. This suggests that they are not
highly palatable to birds; possibly the small spines con-
fer them a limited defense. On the other hand, chicks
showed no other signs of beginning to avoid third-in-
star larvae after having experienced them. They were
almost always attacked (even though the previous ex-
perience was less than an hour earlier), and two chicks
did eat them in all four trials within a time span of less
than three hours. Evidently no chemical defense ef-
fective against birds is present in the comma butterfly.
This conclusion is also supported by the fact that the
adults of the species are highly cryptic and also very
palatable to great tits (Parus major) in laboratory ex-
periments (SN & BST unpubl.). However, this evi-
dence is not conclusive, since butterfly larvae can be
unpalatable even when adults are palatable (Bowers &
Farley 1990, Dyer & Bowers 1996).
The chicks showed no initial aversion to the older
comma larvae, but very rapidly learned to avoid them.
It seems highly improbable that a chemical defense
should be present in old larvae but not in young larvae
or adults. More probably, the strong and sharp spines
are what defend older larvae, as demonstrated by the
result of removing the spines. In any case, the rapid
aversion learning suggests the possibility of an apose-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
matic function of the striking coloration. In the ab-
sence of color manipulations we cannot, however, rule
out the possibility that aversion was in fact to the sight
of the spines themselves, independently of color. In-
terestingly enough, the spines are colored in such a
way that might make them more conspicuous (Fig.
1B), so these two possibilities may be hard to separate.
A dual function is also possible, so that the coloration
indeed mimics bird droppings, as previously suggested
(Thomas 1986) but has an aposematic function once
the larva has been discovered. This would be useful if
some predators are not deterred by the spines, or if
larvae are often damaged by naive predators.
In this context it is, however, interesting to note that
the fifth-instar larvae were not damaged by the attacks
in the initial trials, when the chicks learned to avoid
them. This is in line with previous results from chemi-
cally defended aposematic insects (Boyden 1976, Jarvi
et al. 1981, Wiklund & Jarvi 1982, Wiklund & Sillén-
Tullberg 1985) but has not often been investigated in
mechanically defended prey (but see Carrick (1936)
for results from the related butterfly Aglais urticae).
The importance of such observations is that they
demonstrate that individual selection for aposematic
coloration is possible. In other words it is not necessary
to invoke kin selection or other types of indirect selec-
tion, as would be the case if some individuals must be
sacrificed before predators can learn aversion (Fisher
1930, Benson 1971). Individual selection is a more
parsimonious explanation of aposematic coloration
when direct benefits to the individual are present, be-
cause for indirect selection additional assumptions of
starting conditions are necessary regarding prey family
groupings and the movements and memory of preda-
tors. Indeed, even aposematic butterflies often have
cryptic pupae, and this is the stage of the life cycle that
is most likely to be killed by inspecting predators (Wik-
lund & Sillén-Tullberg 1985). Conversely, strong
spines, which (as demonstrated here) should give di-
rect benefits to the individual, are found very com-
monly in nymphalid larvae, often together with con-
trasting, bright colors or a jet-black color that should
provide little crypsis against green leaves (e.g., in other
Nymphalini and in the Kallimini and Argynniti).
Spines are absent in some chemically defended groups
such as the Danainae, with aposematic larvae, and also
in groups such as the Satyrinae and Apaturinae, which
have clearly cryptic larvae. In many cases, however,
the patterns are far less clear and the function of the
coloration uncertain (for instance, Ladoga larvae in the
Limenitidini have spines but cryptic coloration; Teshi-
rogi 1990).
The ontogenetic shift in defense tactics in P. c-album,
VOLUME 55, NUMBER 2
from cryptic coloration to aposematic coloration and
mechanical defense (seen also in A. urticae; Carrick
1936), is understandable in terms of the general in-
crease in size during larval growth, for several reasons.
First, it may not be possible for a small larva to have
spines large enough to deter birds and other verte-
brate predators. Second, small larvae may be more
vulnerable to attacks by vertebrate predators. If preda-
tors need to learn aversion, small larvae may not sur-
vive this process, and as a consequence it may not be a
profitable strategy to advertise a degree of unpalatabil-
ity (Gamberale & Tullberg 1996). Third, the increase
in size during growth of butterfly larvae has been
found to be coupled with a shift from predominantly
predation by invertebrates to predominantly predation
by vertebrates, such as birds (Kristensen 1994).
Hence, the need for defense against vertebrate preda-
tors should be largest in the last larval instars. Fourth,
it may be difficult to evolve a pattern that is effectively
aposematic given a very small size of the colored areas.
In tests with aposematic bugs (Lygaeidae), chicks
more readily learned aversion towards to the larger
late instars, even though the coloration is the same
(Gamberale & Tullberg 1996).
ACKNOWLEDGMENTS
This research was supported by grants from the Swedish Natural
Science Research Council (NFR) to SN and BST.
LITERATURE CITED
BENSON, W. W. 1971. Evidence for the evolution of upalatibility
through kin selection in the Heliconiinae. Am. Nat.
105:213-226.
Bowers, M. D. & S. FaruEy. 1990. The behavior of gray jays,
Perisoreus canadensis, towards palatable and unpalatable Lepi-
doptera. Anim. Behav. 39:699-705.
BoypDEN, T. C. 1976. Butterfly palatibility and mimicry: experi-
ments with Ameiva lizards. Evolution 30:73-81.
Carrick, R. 1936. Experiments to test the efficiency of protec-
tive adaptation in insects. Trans. R. Entomol. Soc. Lond.
85:131-140.
Cort, H. B. 1940. Adaptive coloration in animals. Methuen, Lon-
don.
Dyer, L. A. & M. D. Bowers. 1996. The importance of se-
questered iridoid glycosides as a defense against an ant preda-
tor. J. Chem. Ecol. 22:1527-1539.
Epmunps, M. 1990. The evolution of cryptic coloration, pp. 3-21.
In Evans, D. L. & J. O. Schmidt (eds.), Insect defences: adap-
tive mechanisms and strategies of prey and predators. State
University of New York Press, Albany.
FisHer, R. A. 1930. The genetical theory of natural selection.
Clarendon Press, Oxford.
GAMBERALE, G. & B.S. TULLBERG. 1996. Evidence for a peak-shift
inpredator generalization among aposematic prey. Proc. R. Soc.
Lond. B. 263:1329-1334.
GuiLrorb, T. 1990a. The evolution of aposematism, pp. 23-61. In
Evans, D. L. & J. O. Schmidt (eds.), Insect defences: adaptive
mechanisms and strategies of prey and predators. State Univer-
sity of New York Press, Albany.
1990b. Evolutionary pathw ays to aposematism. Acta Oe-
cologia 11:835-841.
JAv1, T., B. SILLEN-TULLBERG & C. WIKLUND. 1981. The cost of
being aposematic. An experimental study of predation on larvae
of Lope machaon by the great tit Parus major. Oikos
32:267-272.
KETTLEWELL, B. 1973. The evolution of melanism. The study of a
recurring necessity: with special reference to industrial
melanism in the Lepidoptera. Clarendon Press, Oxford.
KINGSOLVER, J. G. & W. B. Watt. 1983. Thermoregulatory strate-
gies in Colias butterflies: thermal stress and the limits to
adaptation in temporally varying environments. Am. Nat.
121:32-55.
KRISTENSEN, C. O. 1994. Investigations on the natural mortality of
eggs and larvae of the Large White Pieris brassicae (L) (Lep-
Pieridae). J. Appl. Entomol. 117:92-98.
MALcoL, S. B. 1990. Mimicry: status of a classical evolutionary
paradigm. TREE 5:57-62.
PouLToN, E. B. 1890. The colours of animals. Their meaning and
use. Especially considered in the case of insects. Kegan Paul,
Trench, Trubner, London.
Scott, J. A. 1986. The butterflies of North America. Stanford Uni-
versity Press, Stanford, California.
SCUDDER, S. H. 1889. The butterflies of the Eastern United States
and Canada with special reference to New England. Published
by the author, Cambridge.
SHaprrO, A. M. 1976. Seasonal polyphenism. Evol. Biol. 9:
259-333.
SILLEN-TULLBERG, B. 1988. Evolution of gregariousness in apo-
sematic butterfly larvae: a phylogenetic analysis. Evolution
42:293-305.
TrsHirocl, M. 1990. An illustrated book of the Japanese Nymphal-
idae. Tokai University Press, Tokyo.
Tuomas, J. A. 1986. RSNC guide to butterflies of the British isles.
Country Life Books/Hamlyn Publishing Group Ltd, Twicken-
ham.
WIKLUND, C. & T. JARVI. 1982. Survival of distasteful insects after
being attacked by naive birds: a reappraisal of the theory of
aposematic coloration evolving through individual selection.
Evolution 36:998-1002.
WIKLUND, C. & B. SILLEN-TULLBERG. 1985. Why distasteful but-
terflies have aposematic larvae and adults, but cryptic pupae:
evidence from predation experiments on the monarch and the
European swallowtail. Evolution 39:1155-1158.
Received 15 January 2000; revised and accepted 30 August 2001.
|
Journal of the Lepidopterists’ Society
55(2), 2001, 74-77
SEXUAL DIMORPHISM IN EYE MORPHOLOGY IN EUCHEIRA SOCIALIS (PIERIDAE)
NELL M. LUND, EMILY E. CWENGROS, AND RONALD L. RUTOWSKI
Department of Biology, Arizona State University,
Tempe, Arizona 85287-1501, USA
ABSTRACT. We examined the magnitude of sexual dimorphism in the eyes of madrone butterflies, Eucheira socialis (Pieridae). Via mi-
croscopic examination of the cornea, we determined the eye surface area, facet number, and facet diameter in 5 eye regions for males and fe-
males. Our analysis, which controlled for body size (forewing length), showed that in this species, as in most Lepidoptera, males have a signifi-
cantly larger eye surface area and more facets than females. Facet diameters vary with eye region but in an unusual way for the Lepidoptera:
the largest facets we observed were in the dorsal region of the male eye. While the results reveal interesting patterns of sexual dimorphism in
eye structure in E. socialis there is insufficient information about adult behavior to understand the behavioral and ecological implications and
causes of these patterns, especially those in facet diameter.
Additional key words: _ sexual dimorphism, eye structure, eye size, facet diameter.
Sexual dimorphism in eye structure is common in
the Lepidoptera (Yagi & Koyama 1963). Males usually
have eyes that are larger and have more and larger
facets than those of females. In response to a presen-
tation on sexual dimorphism in butterfly eyes by one of
us (RLR) at the 1999 meeting of the Lepidopterists’
Society, Dr. Arthur Shapiro suggested that sexual di-
morphism in eye structure was particularly pro-
nounced in the madrone butterfly, Eucheira socialis
Westwood (Pieridae). We investigated this claim by ex-
amining corneal structure in males and females of this
species.
Little has been reported about the behavior of E. so-
cialis adults that bears on the question of selection
pressures that might have shaped eye structure. Previ-
ous studies of E. socialis have focused on the social be-
havior of the larvae and the relationship between this
butterfly and madrone trees (Arbutus spp., Ericaceae)
on which the larvae feed (Kevan & Bye 1991, Under-
wood 1992, 1994). The adults live for approximately
one week and are apparently unable to feed because
the two halves of the proboscis do not anneal correctly.
Within 3 h of eclosion adult females generally travel
about 15-18 m to oviposit. A female lays her eggs in a
single large mass (20 to over 350 eggs) on the ventral
side of one madrone leaf (Underwood 1992). Males fly
around madrones and other trees in the vicinity only
when there is bright sunlight (D. Underwood pers.
comm.). The mating behavior of males and females is
essentially unknown.
MATERIALS AND METHODS
The E. socialis specimens used in this study were
obtained from Art Shapiro (University of California,
Davis) and Dessie Underwood (California State Uni-
versity, Long Beach) and reported to have been reared
in April 1991 from larvae found in two nests in the
Sierra Madre Occidental, Durango, Mexico. We re-
ceived them as papered specimens. We processed
each of 5 males and 5 females in the following way. We
measured the forewing length (FWL) to the nearest
0.1 mm using dial calipers. Then the head was re-
moved, bisected, and immersed in 10% NaOH solu-
tion until the soft tissue could be gently removed from
each cornea, approximately 5 minutes. Non-corneal
cuticle was trimmed away from the eye. Radial cuts—
two starting close together dorsally, one starting ven-
trally, and some starting both anteriorly and posteriorly
from the edges of the cornea—were made toward the
center of the cornea (Fig. 1). These permitted the
cornea to lie flat on a microscope slide and provided
references for identification of eye regions. To im-
prove the visibility of the cornea for handling, we ap-
plied acid fuchsia stain for a few minutes to the
cornea. Finally, the cornea was mounted in glycerol
and covered with a coverslip, the edges of which we
sealed with Cytosol 60 mounting medium (Stephen
Scientific).
Digitized images of each cornea were obtained by
video microscopy with a Nikon inverted microscope.
Approximately 15 images of each cornea made with a
10x objective were required to capture the entire
cornea. A composite image of each complete cornea
(for example, see Fig. 1) was created using Adobe
Photoshop (Adobe Systems Incorporated). For a size
reference a micrometer scale at 10x was photo-
graphed.
The numbers of facets in the cornea were manually
counted from the composite images. However, we
used image analysis software (Scion Image) to mea-
sure facet diameters and eye surface area (ESA) from
the composite images. The micrometer scale on each
image was used to calibrate the software. To obtain the
ESA of each image an outline tool was used. Two ESA
measurements taken by this method were averaged to
control for error in tracing the image; if the two mea-
surements differed by more than 5%, then a third
measurement was included in the average. Facet di-
VOLUME 55, NUMBER 2
Fic. 1. A micrograph of a representative comea from the eye of a male E. socialis made from multiple images. Dorsal is toward the top and
BeBONS
Batt ws
SOT Ogta scien,
age
x
~ tees FPR ee
“Bh (TH
ec Lace ms
me gO gO: ©
2 On On Oar j2ge,
ees
OG I BeaOe
2
On
in hea 0,
ae0g?,
anterior to the left in this image. Scale line = 0.25 mm.
cS S
Se.
fy
ose.
ose.
OSes.
Fic. 2. Schematic of the comea of E. socialis showing the facet
diameters (in mm; mean + standard deviation) measured for both
males and females in each of five regions of the cornea. Abbrevia-
tions: A, anterior; D, dorsal; P, posterior; V, ventral.
ameters were calculated by dividing the length of a
row of 10 facets by 10. Two measurements were taken
in each of the 5 eye regions (anterior, posterior, dorsal,
ventral, center), then averaged. If the two measure-
ments differed by more than 8%, then a third mea-
surement was included in the average. The measure-
ment points within each eye region were chosen based
on the clarity of the facet images within that region
(i.e. no wrinkles or tears in the cornea). All statistical
analyses were done on SYSTAT 9.0 (SYSTAT, Inc.).
RESULTS
The males in our sample were significantly smaller
than the females as measured by forewing length
(males: 25.7 + 1.22 mm:; females: 31.2 + 1.43 mm: t =
6.58, 8 df, p < 0.0002). Despite their smaller body size,
males have a significantly larger ESA than females
(males: 4.45 + 0.15 mm/?; females: 3.20 + 0.15 mm/?;
ANCOVA, with forewing length as covariate, p <
0.0001). ESA is not correlated with forewing length
within either sex (p = 0.135), although sample sizes
were small.
A larger ESA in males could mean that compared to
females they have more facets, larger facets, or both.
Males have significantly more facets (7781 + 85) than
females (6648 + 357: ANCOVA, with forewing length
as covariate, p = 0.011), but within the sexes there was
no significant correlation between facet number and
ESA (p = 0.117).
The mean facet diameters for each of the five eye
regions in males and females are illustrated in Fig. 2. A
two-way repeated measures ANOVA with facet diam-
eter as the dependent variable and region as the re-
peated measure showed that the effects of region, sex,
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
and the interaction between sex and eye region were
all significant (p < 0.001 for each). Therefore, variation
in facet diameters is explained by both sex and eye re-
gion, but there are significant differences between
sexes in the pattern of variation among regions. Facet
diameters of males were generally larger than those of
females but especially in the dorsal region. In fact,
only in the dorsal eye region was the difference in
facet diameter between males and females significant
(t = 8.69, 8 df, p < 0.0001). Facet diameters did not
differ significantly between males and females in any
other eye regions (anterior, t = 1.9, posterior, t = 0.368,
ventral, ¢ = 0.376; center, t = 2.157; all: 8 df, p > 0.05).
Another clear trend in the eyes of both males and fe-
males was that facets in the ventral region tended to be
smaller than those in the anterior, central, and poste-
rior regions along the equator of the eye.
DISCUSSION
Compared with conspecific females who are on ay-
erage larger in body size, E. socialis males have eyes
that are 1.39 times larger in surface area, have 1.17
times more facets, and have, at least dorsally, larger
facet lenses. Contrary to Dr. Shapiro’s initial impres-
sion, this sexual dimorphism in eye morphology is sim-
ilar in direction and magnitude to that seen in butter-
flies and other Lepidoptera (Rutowski 2000). For
comparison, Yagi and Koyama (1963) report male : fe-
male eye surface area ratios of 1.37 and 1.33 and facet
number ratios of 1.1 and 1.18 for two pierids, Colias
erate and Pieris rapae, respectively. Eyes in males that
are larger and have more ommatidia relative to body
size than those in females are generally interpreted as
indicating that acute vision in males is more important
to reproductive success than in females (Rutowski
2000, and in press).
In spite of this general similarity in eye structure,
the pattern of variation in facet diameters is quite dif-
ferent from that reported for another butterfly, Aster-
ocampa leilia. In A. leilia the largest facets are found in
the frontal and dorsofrontal regions of the eye regions
with facets in the dorsal region being smaller (Ziemba
& Rutowski 2000). Large facets indicate eye regions of
high resolution and sensitivity and imply that those re-
gions of the eye are frequently used in specific tasks
requiring high resolution or sensitivity such as when
males track females during rapid pursuit flights (Land
1989, 1997). The large facets in the dorsal part of the
male eye in E. socialis suggest that dorsal vision is bet-
ter than in other parts of the visual field. Little is
known. about the behavior of adult madrone butterflies
so the possible functions of this dorsal region of high
acuity are not clear, but it might be important in sexual
VOLUME 55, NUMBER 2
interactions. In some insects, acute dorsal vision pro-
duced by large facets is important in males for detect-
ing females against the sky (Land 1997). Some pierids
have been observed engaging in ascending flights dur-
ing courtship attempts. During an ascending flight a
male-female pair flies up together until one of them,
usually the male, terminates pursuit and drops rapidly
toward the ground (Rutowski 1978a, b). When Dessie
Underwood threw E. socialis females up in the air in
an attempt to pair them, she noted that E. socialis
males would only chase the female if she flew well (D
Underwood pers. com.). If ascending flights are a
common occurrence in E. socialis courtship, then the
male’s dorsal acute zone may be used for tracking the
female’s movements in this context.
ACKNOWLEDGMENTS
Our sincere thanks: to Dr. Arthur Shapiro for posing the question
that prompted our study; to Drs. Shapiro and Dessie Underwood for
providing the specimens, information on the behavior and natural
history of E. socialis, and helpful comments on an earlier draft of the
manuscript; to Kristine Ziemba for her consultation on cornea
preparation techniques; to Dennis McDaniel at the ASU Keck Lab-
oratory for instruction on video microscopy, Adobe Photoshop, and
Scion software; to Ian Gould and Faye Stump for use of their lab
and materials to make stains; and, finally, to NSF for financial sup-
port (Grant #97-23160 to RLR).
LITERATURE CITED
KEvAN, P. G. & R. A. Bye, 1991. The natural history, sociobiology,
and ethnobiology of Eucheira socialis Westw ood ( (Lepidoptera:
Pieridae), A unique and little known butterfly from Mexico. En-
tomologist 110(4):146—-165.
LAND, M. F. 1989. Variation in the structure and design of com-
pound eyes, pp. 90-111. In Stavenga, D. & I. Hardie (eds.),
Facets of vision. Springer-Verlag, Berlin.
. 1997. Visual acuity in insects. Ann. Rey. Entomol.
49:147-177.
Rutowskl, R. L. 1978a. The courtship of the small sulphur butter-
fly, Eurema lisa. Anim. Behav. 26:892-903.
. 1978b. The form and function of ascending flights in Col-
ias butterflies. Behav. Ecol. Sociobiol. 3:163—172
2000. Eye size variation in butterflies: Inter- aa intraspe-
eikie patterns. J. Zool. 252:187-195.
. Inpress. Visual ecology of adult butterflies. In Boggs, C. &
W. Watt (eds.), Ecology and evolution taking flight: butterflies
as model study systems. University of Chicago Press’ Chicago.
UnbDERWoOD, D. L. 1992. Factors influencing oviposition behavior
in the Mexican Pierid butterfly. Ph.D. Dyasamatton, University
of California, Davis, California.
. 1994. Intraspecific variability in host plant quality and
ovipositional preferences in Eucheira socialis (Lepidoptera:
Pieridae). Ecol. Entomol. 19:245-256.
Yact, N. & N. Koyama. 1963. The compound eye of Lepidoptera:
approach from organic evolution. Shinkyo Press, Tokyo.
ZIEMBA, K. S. & R. L. Rutowski. 2000. Sexual dimorphism in eye
morphology in a butterfly (Asterocampa leilia: Lepidoptera,
Nymphalidae). Psyche 103:25-36.
Received 3 November 2000; revised and accepted 28 August 2001.
GENERAL NOTES
Journal of the Lepidopterists’ Society
55(2), 2001, 78-79
OBSERVATIONS OF ADULT AND LARVAL BEHAVIOR IN THE WINTER SPHINGID,
ARCTONOTUS LUCIDUS (SPHINGIDAE)
Additional key words: _ phenology, hostplants, circadian habits, mate location, Bear sphinx moth.
Arctonotus lucidus (Boisduval), the Bear Sphinx, is a poorly
known, small green sphingid that occurs along the Pacific coast of
the United States and Canada. Part of the reason for the Bear
Sphinx’s anonymity is its very early season flight, ranging from De-
cember in southem California to April in Washington state and
British Columbia, when little else is flying. After its description by
Boisduval (1852), it was 90 years before Comstock and Henne
(1942) described the larva and adult phenology. Those descriptions
were based upon specimens reared from eggs on Oenothera dentata
Jepson. Osborne’s (2000) report of larvae on Clarkia breweri (A.
Gray) E. Greene and C. modesta (Jepson) represents the first con-
firmed record of larvae found on hostplants in nature.
Female specimens are rare in collections: there are only 7 of 285
(2.5%) field collected specimens in the Essig Museum (U.C.
Berkeley), the Bohart Museum (U.C. Davis), the Los Angeles
County Museum, and the San Diego County Museum combined.
The rarity of captured females and larvae explains why so little is
known about the species’ natural history. The purpose of this note is
to describe poorly known aspects of adult behavior including
circadian habits (flight times), mating, oviposition, and some larval
behavior.
On 2 February 1997 J. Kruse and I captured 30 males (no fe-
males) using mercury vapor and UV traps near Chinese Camp,
Tuolumne Co., California. On 7 February 1997 we were unsuccess-
ful in attracting any Arctonotus to the same traps at the same locali-
ties. However, that same evening we took two females and one male
at residential lights in the same general area.
Captured females were caged outdoors in Berkeley, Alameda
Co., California. Feeding was attempted by forcibly unrolling the
proboscis into sugar water (a technique that works with many Lepi-
doptera). I found that the proboscis was vestigial, flaccid and not ca-
pable of reaching the substrate on which the moth was perched.
Captive females became active at dusk (1745-1800 h PST) and laid
eggs singly or in pairs after brief flights around the enclosure. The
two females lived 5 days and laid a combined total of 380 eggs. Ap-
proximately half the eggs were maintained out of doors (4-21°C),
and the rest were brought indoors (16-21°C). Indoors the eggs
turned from pale green to yellow in 7 days and hatched 9 days fol-
lowing deposition. The eggs maintained outside hatched in 19 days,
10 days after those maintained indoors. Larvae were fed Fuschia
thymefolia Kunth, Clarkia unguiculata Lindley, C. amoena Lehm..,
and Oenothera species (all Onagraceae). Osborne (in press) provides
an excellent description of larval biology, to which I only add the ob-
servation that large (5 cm) 5th instar larvae regularly hide at the base
of the hostplant during non-feeding periods; mature larvae fed
mostly in the morning and late afternoon. The larvae range from
nearly all green with yellow spiracles to pink and black to all black
and this habit occurred regardless of color form.
Fic. 1. Female Arctonotus lucidus in
‘calling’ posture. Photo by Daniel Rubinoff.
VOLUME 55, NUMBER 2
Larvae began to burrow in soil in preparation for pupation 36—45
days after egg hatch. Mortality was high at this stage: of 45 larvae
that burrowed, only 18 pupated. Larvae pupated under dead leaves
and pieces of wood, just under the soil surface, and up to 16.5 cm
underground in the rootball of senescing hostplants. Pupation usu-
ally occurred in firmly packed ovate cells. The cremaster possesses a
bifureate tip as depicted by Osborne (1995) for Proserpinus clarkiae
(Boisduval) (Sphingidae).
The 18 pupae were maintained outside in a ventilated plastic tub
in Berkeley until November when they were placed in a refrigerator
at 1.7°C + 1°C. No development was evident in the pupae until they
had been moved from refrigeration to outside temperatures (be-
tween 8—20°C) for more than 45 days. J. Kruse (pers. com.) found
that daily cycling of pupae removed from a refrigerator (3°C) to
room temperature (18°C) for approximately 4-S hours, also induced
eclosion. The green coloration of the developing wings was visible
through the pupal cuticle for two days before the moths emerged.
The cuticle became very soft 24 hours before emergence.
Adults eclosed from 8-19 March 1998, usually between 1800 and
1900 h PST; they took 1-2 hours to dry their wings. Adults in cages
were active only from 1800 to 1930 h PST, though mating occasion-
ally lasted a few hours longer. Virgin females rested on the substrate,
everting and pulsating the papillae anales to disperse pheromone
(Fig. 1). When a male was placed in the same enclosure he rapidly
approached the female and mated. If no male arrived by 2000 h PST,
females stopped calling until the next sunset. One male fertilized
three females; those females laid 369, 397, and 401 eggs respectively.
Arctonotus lucidus pupae apparently are able to develop when sur-
face temperatures still regularly fall below freezing. Eggs hatch and
larvae begin development when most apparent hostplants are less
than 2 cm high, and night temperatures occasionally fall below 0°C.
Journal of the Lepidopterists’ Society
55(2), 2001, 79-80
79
Hodges (1971) stated that adults can be collected during the day
while nectaring on flowers. I suspect this report to be in error since
all collections throughout the range of the moth, that I have been
able to document, were made at lights, and none of the moths I
reared was active during the day. Moreover, the adults do not feed.
I thank M. Caterino, J. Herbeck, J. Kruse, F. Sperling, J. Tuttle
and especially J. Powell for comments which greatly improved the
quality of this work and J. Kruse for his assistance in the field. J.
Powell and J. DeBenedictis helped with museum tallies. This work
was supported in part by an ARCS foundation fellowship, the M. C.
Walker fund, and a grant from the California Agricultural Experi-
ment Station to F. Sperling.
LITERATURE CITED
BOIsDUVAL, J. 1852. Annales de la Societé Entomologique de
France, 2nd series 10:319.
Comstock, J. A. & C. HENNE. 1942. The early stages of Arctonotus
ene Bdy. (Lepidopt.). Bull. South. Calif. Acad. Sci.
41(3):167-171.
Hopces, R. W. 1971. Moths of North America, north of Mexico,
Fascicle 21(Sphingidae). 170 pp.
OsporNE, K. H. 1995. Biology of Proserpinus clarkiae (Sphingi-
dae). J. Lepid. Soc. 49(1):72-79.
OsporNE, K. H. 2000. Additional notes on Proserpinus clarkiae
and Arctonotus lucidus (Sphingidae) life histories from the Pa-
cific coast of North America. J. Lepid. Soc. 53(4):170-172.
DANIEL RUBINOFF, 201 Wellman Hall, Division of Insect Biology.
University of California, Berkeley, California 94720.
Received 25 February 2000; revised and accepted 23 July 2001.
EMERGENCE OF PARASITIC FLIES FROM ADULT ACTINOTE DICEUS (NYMPHALIDAE: ACRAEINAE) IN ECUADOR
Additional key words: _ parasitoid, adult Lepidoptera, neotropics, Arachidomyia.
A parasitoid is defined as ‘an organism which develops on or in
another single (host) organism, extracts nourishment from it, and
kills it as a direct or indirect result of that development (Eggleton &
Gaston 1990). In contrast parasites rarely kill their hosts and preda-
tors always consume more than one host. In addition, parasitoids
possess a free-living adult stage (whereas many parasites do not),
and do not reproduce inside the host (as do many parasites). Insects
with parasitoid life cycles are known from many taxanomic groups
including many families of Hymenoptera and Diptera (Eggleton &
Belshaw 1992), yet knowledge of host specificity and parasitoid life-
cycles remains patchy. Host relationships are known for only a small
percentage of parasitoid taxa in the tropics (e.g., references in Han-
son & Gauld 1995), and many parasitoid species remain unde-
scribed due to their often small size and highly specialized lifestyles
(Gaston 1991). The emergence of parasitoids from adult Lepi-
doptera is infrequently reported in the literature (e.g., Marshall
1896, Cockayne 1911, Edelsten 1933, DeVries 1979, Smith 1981,
McCabe 1998). The following record of sarcophagid flies emerging
from adult butterflies in Ecuador represents the first record of this in
several years, and only the second record involving Sarcophagidae.
On 8 December 1996 three female Actinote diceus Latreille
were collected at Cabamias San Isidro, located at around 2000 meters
elevation in north-eastern Ecuador. All were flying normally along a
road cut through disturbed cloud forest and cattle pasture. At the
time of collection all three butterflies were killed by a quick pinch to
the thorax, as described by DeVries (1987), placed together inside a
glassine envelope, and marked with the date and locality. The spec-
imens were then placed together in a plastic tub with other speci-
mens collected that day and returned to the lab. Upon arrival at the
lab and inspection of the specimens, two fly pupal exuviae were
found inside the envelope with the three Actinote females. Two
adult sarcophagid flies were also present inside the envelope. These
adults were identified using Shewell (1987) as belonging to the
genus Arachidomyia Toenceal Due to eclosion inside the glassine
envelope, both specimens were badly damaged and could not be
identified to species. Another empty puparium was found in an enve-
lope containing a fourth individual female A. diceus collected on the
same date, but no adult fly was recovered. Lepidopteran specimens
were retained in the collection of the senior author and the dipteran
specimens were deposited in the Tulane University collection.
Sarcophagid flies develop on a wide variety of food resources and
range in habit from detritivores to predators and parasitoids of in-
vertebrate and vertebrate hosts (Clausen 1940). The parasitoid habit
appears to have evolved on many separate occasions, and about half
80
of the described species can be considered parasitoids or cleptopar-
asitoids. Of these, approximately 750 species can be considered true
internal parasitoids (Eggleton & Belshaw 1992). Parasitoid Sar-
cophagidae are known to develop inside adult hosts of a variety of in-
sects and vertebrates including Orthoptera, spiders, gastropods, and
lizards (Shewell 1987, Allen & Pape 1996, Dial & Roughgarden
1996, Danyk et al. 2000, Pape et al. 2000). A number of Sarcophagi-
dae, including Arachidomyia, have been reared from lepidopteran
pupae (Clausen 1940, Shewell 1987, Parry 1995) and McCabe
(1998) reported Sarcophaga (=Arachidomyia) aldrichi Parker
emerging from adult moths, but large scale larval lepidopteran rear-
ing projects have found very few instances of parasitism by sar-
cophagids (Janzen & Hallwachs 1999, Dyer & Gentry 2001, Stire-
man & Singer unpublished data). In our area, little is known of the
life history of Actinote diceus. Adults and larvae are present year-
round and found in association with their larval food plant, Erato
polymnioides DC (Asteraceae) (Greeney et al. 2001). While their
adult lifespan is unknown adult females live only a few days in cap-
tivity (HFG unpublished data). McCabe (1998) argued that attack
by Arachidomyia occurred during the adult stage of the host, how-
ever in the present case this is unlikely due to the “fresh” appear-
ance of the adult butterflies and the rapidity with which the adult
flies appeared. We suspect that the parasitized individuals were at-
tacked as pupae.
Knowledge of parasitoid/host associations is necessary for under-
standing the structure and function of ecological communities, de-
veloping theories of parasite/host population dynamics, and estab-
lishing of sound biological control programs (Godfray 1994,
Hawkins 1994, Hawkins & Sheehan 1994, Jervis & Kidd 1996, Van-
Driesche & Bellows 1996). Reports of parasitism of adult Lepi-
doptera are rare, but extensive rearing programs, especially in poorly
studied tropical ecosystems, are needed to assess whether these re-
lationships tend to be facultative or obligate and generalized or spe-
cialized. Such rearing data is also needed to determine how para-
sitism of adults is achieved (i.e., via immature stages or directly) and
to assess the prevalence of this life history strategy and how it may
effect host populations. We hope that this report encourages further
research on tropical parasitoid-host relationships and the prevalence
of adult parasitism in Lepidoptera.
We thank P. DeVries, L. Dyer, and an anonymous reviewer for
thoughtful comments on earlier versions of this manuscript. We also
wish to thank J. E. O'Hara for examining the sarcophagid specimens.
Thank you to the PBNHS for their ongoing support of our field work.
LITERATURE CITED
ALLEN, G. R. & T. PAPE. 1996. Description of female and biology
of Blaesoxipha ragg Pape (Diptera: Sarcophagidae), a parasitoid
of Sciarasaga quadrata Rentz (Orthoptera: Tettigoniidae) in
Western Australia. Aust. J. Entomol. 35:147-151.
CLAUSEN, C. P. 1940. Entomophagus insects. Hafner, New York, N.Y.
Cockayne, E. A. 1911. A dipterous parasite bred from imago of
Nyssia lapponaria. Entomologist 44:253.
Danyk, T., D. L. JouNsON & M. MACKAUER. 2000. Parasitism of the
grasshopper Melanoplus sanguinipes by a sarcophagid fly, Blae-
soxipha atlanis: influence of a solitary and gregarious develop-
ment on host and parasitoid. Entomol. Exp. et Appl. 94:259-268.
DeVries, P. J. 1979. Occurrence of fly maggots in adult Morpho
thesus Deys. (Lepidoptera: Morphidae) females from Costa
Rica. Brenesia 16:223.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
. 1987. The butterflies of Costa Rica and their natural
history. Vol. 1. Papilionidae, Pieridae, and Nymphalidae.
Princeton University Press, Princeton, New Jersey.
DIAL, R. & J. ROUGHGARDEN. 1996. Natural history observation of
Anolisomyia rufianalis (Diptera: Sarcophagidae) infesting Anolis
lizards in a rain forest canopy. Environ. Entomol. 25:1325-1328.
Dyer, L. A. & G. L. Gentry, 2001. Caterpillars and parasitoids of
a Costa Rican tropical wet forest. (http:/Avww.caterpillars.org).
EDELSTEN, H. M. 1933. A tachinid emerging from an adult moth.
Proc. Entomol. Soc. Lond. 8:131.
EGCLETON, P. & R. BELSHAW. 1992. Insect parasitoids: an evolu-
tionary overview. Phil. Trans. R. Soc. Lond., Series B 337:1-20.
EGGLETON, P. & K. J. Gasron, 1990. “Parasitoid” species and as-
semblages: convenient definitions or misleading compromises?
Oikos 59:417-421.
GasTON, K. J. 1991. The magnitude of global insect species rich-
ness. Conservation Bio. 5:283-296.
Goprray, H. C. J. 1994. Parasitoids: behavioral and evolutionary
ecology. Princeton University Press, Princeton, New Jersey.
GREENEY, H. F., T. R. WALLA, & L. A. DyER. 2001. Caterpillars and
parasitoids of the Eastem Andes in Ecuador. (http://www.
caterpillars.org/ecuador).
Hanson, P. E. & I. D. GauLp (eds.) 1995. The Hymenoptera of
Costa Rica. Oxford University Press, New York.
Hawkins, B. A. 1994. Patterns and process in host-parasitoid inter-
actions. Cambridge University Press, Cambridge, Massachussets.
Hawkins, B. A. & W. SHEEHAN. 1994. Parasitoid community ecol-
ogy. Oxford University Press, New York.
JANZEN, J. D. & W. Hatiwacus. 1999. Philosophy, navigation, and
use of a dynamic database (“ACG Caterpillars SRNP”) for an
inventory of the macrocaterpillar fauna, and its food plants and
parasitoids, of the Area de Conservacion Guanacaste (ACG),
northwestern Costa Rica (http://janzen.sas.upenn.edu).
Jervis, M. & N. Kipp. 1996. Insect natural enemies. Chapman &
Hall, London.
MARSHALL, T. A. 1896. Survival of Acherontia atropos after being
struck by an ichneumon. Ent. Mo. Magazine 32:265.
McCabe, T. L. 1998. Dipterous parasitoids from the adults of
moths (Lepidoptera). Entomol. News 109:325-328.
Pape, T., S. C. MCKILLIP & R. V. MCKILLIP. 2000. Two new species
of Sarcophaga (Sarcorohdendorfia) Baranov (Diptera: Sar-
cophagidae), parasitoids of Littoraria filoso (Sowerby) (Gas-
tropoda: Littorinidae). Aust. J. Entomol. 39:236-240.
Parry, D. 1995. Larval and pupal parasitism of the forest tent
caterpillar, Malacosoma disstria Hubner (Lepidoptera: Lasio-
campidae) in Alberta, Canada. Can. Entomol. 127:877-893.
SHEWELL, G. E. 1987. Sarcophagidae. In J. F. McAlpine (ed.),
Manual of neartic diptera. Vol. 2. Research Branch, Agriculture
Canada. Monograph No. 28, pp. 1159-1186.
SmiTH, K. G. V. 1981. A tachinid (Diptera) larva in the abdomen of
an adult moth (Geometridae). Ent. Gazette 32:174-176.
VANDRIESCHE, R. G. & T. S. BELLOWS. 1996. Biological control.
Chapman & Hall, New York.
HAROLD F. GREENEY, Yanayacu Biological Station and Center for
Creative Studies, Cosanga, Ecuador, c/o Foch 721 y Amazonas,
Quito, Ecuador AND JOHN O. STIREMAN, Tulane University, 310
Dinwiddie Hall, New Orleans, Louisiana 70118, USA
EDITORIAL STAFF OF THE JOURNAL
Caria M. Penz, Editor
Department of Invertebrate Zoology
Milwaukee Public Museum
Milwaukee, Wisconsin 53233, USA
flea@mpm.edu
Pui DeVuies, Book Review Editor
Center for Biodiversity Studies
VS emaals Milwaukee Public Museum
es) ah “Milwaukee, Wisconsin 53233, USA
pjd@mpm.edu
- Associate Editors:
Gerarpo Lamas (Perw), Kenetm W. Puitie (USA), Rosert K. Rospins (USA),
Feuix Spertinc (Canada), Davin L. Wacner (USA), Cunister WikLUND (Sweden)
NOTICE TO CONTRIBUTORS
5 to the onic may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Techni-
Book Reviews, Obituaries, Feature Photographs, and Cover Illustrations. Obituaries must be authorized by the president
Requirements fi for Feature Photographs and Cover Illustrations are stated in Volume 44(2):111 and on the Society's web
tes er Journal Submissions to the editor at the shee address (or electronically to: eatainpes edu). Contributors should feel free to
mmend one or ek reviewers upon ete of their see pene ee to review pots for the Journal ; and ia re-
i: eae to: gaan utexas.edu). Before ciated moleuscripts or electronic ales ck iistraetions
7-50 or the Society's web site at http:/Avww.furman.edu/~snyder/snyder/lep/.
cripts in triplicate, typewritten, entirely double-spaced, with wide margins, on one side only of white letter-sized pa-
ipt according to the following instructions, and submit them flat, not folded. Submit original illustrations only af-
a cript has been revised and accepted, Authors should provide a idiskate copy of the final accepted manuscript in a stan-
aPC or Macintosh- -based word processing format.
“Include an informative abstract for Articles, Profiles, and Technical Comments.
y words: a to se see ose or terms not in the title should 2 accompany Articles, Profiles, General Notes, and
ne
Laie
Fi ures, “ath fine saving and. photoxaphs sist Re numbered consecutively in Arabic naan ea not use plate ae
ee. can be Seana Kinds of tats thie require caaehece frichude dese pHots of new taxa; life histories, host assaci-
at mmature morphology, and some experimental studies. —
ie - Proofs: | The edited manuscript and galley proofs will be mailed to the author for correction of printer's errors. Excessive author's
pe 0 ee will be charged to authors at the rate of $3.00 per line. A purchase order for reprints will accompany proofs.
3 eh For fos pee wis institutions, page ee are $50 ae Journal pees For caek s ewes msmtnOn
on. Aue of Book evens and Obituaries are ne from page tee
rrespondence: Address all matters relating to the Journal to the editor. Address book reviews directly to the book review
enue BY THE ALLEN PRESS, INC, LAWRENGE, KANSAS 66044 U.S.A.
:
af
Number 3
ISSN 0024-0966
THE LEPIDOPTERISTS’ SOCIETY
EXEcuTiveE Councin
Joun W. Brown, President Susan S. Borkin, L Vice President
Micnagr J. Smirn, Immediate Past President Mirna M. Casacranpe, Vice President
Manuer A. Batcazar-Lara, Vice President Davin C. Trrner, Treasurer
Ernest H. Wituirams, Secretary
: Members at large:
Ronald L. Rutowski M. Deane Bowers: George L. Balogh
Felix A. H. Sperling Ron Leuschner Andrew. V. Z. Brower
Andrew D. Warren’ Michael Toliver © Brian Scholtens ©
E:prroriAL Boarp
Rosert K, Rospins (Chairman), ), Joun W. Brown (Member at large)
Caria M. Penz ( Journal)
WituaM E. Mitier (Memoirs)
Puiu J. Scuarrerr (News)
Honorary Lir— MEMBERS OF THE SOCIETY
Cares L. Remincton (1966), E. G. Munror (1973), Ian F: B. Common (1987),
Joun G. Franciemonr (1988), Lincotn P. Brower (1990), Doucias C. Fercuson (1990),
Hon. Miriam Roruscuitp (1991), Craupe Lemame (1992), Freperick H. Rinpce (1997)
The object of The Lepidopterists’ Society, which was formed in May 1947 and formally constituted in December 1950, is
mote ene science of ama i in all its ae . to issue a pended! Aue other ee on ee to
sued, with addresses aiid special interests.
Active members—annual dues $45.00 within the U.S., $50.00 oeside theS.. Cutan
Affiliated members—annual dues $10.00 within the U.S.. $15.00 outside the U.S.
Student members—annual dues $20.00 within the U.S., $25.00 outside the U.S.
Sustaining members—annual dues $60.00 within the U.S., oes 00 outside the U. S.
Life members—single sum $1,800.00
Institutional subscriptions—annual $60.00
Airmail postage for the News $15.00
Sead remittances, payable to The Lepidopterists’ Society, to: Kelly M. Richers, Asst. Treasurer, 9417 Carvalho cae Bakersh Id
CA 93311; oe address changes to: Heit Z Dereinie, Nae ea Museum, 900 apace Blvd., Los eee! CA ABOUT 05
field; CA 93311.
The additional cost for members outside the U.S. is to cover mailing coe
Jodraal of The Lepidopterists’ Society (ISSN 0024- 0966) is published quarterly by The Lapdspredse Society, ‘fo Los At eS.
County Museum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, Cc
and at additional mailing offices. POSTMASTER: Send address changes to The Lepsepienss: Society, “Yo Natural Histaty Museu
900 HADES Blvd., Los Angeles, CA 90007-4057. Sy
Cover illustration: Head capsule of Memphis 5th instar caterpillar (Nymphalidae, Charaxinae), by C. M. Penz.
JOURNAL OF
tine LePpiporpTERISTS’ SOCIETY
Volume 55
Journal of the Lepidopterists’ Society
59(3), 2001, 81-84
2001 Number 3
A NEW SPECIES OF EUCOSMOMORPHA FROM NORTH AMERICA (TORTRICIDAE)
WILLIAM E. MILLER
Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108, USA (mille014@tc.umn.edu)
ABSTRACT. Eucosmomorpha nearctica, new species, is described from 19 adult specimens, 18 male and 1 female. It previously was
misidentified in North America as E. albersana (Hiibner). It differs from E. albersana in its more mottled forewing, smaller body size, more
prominent male hindwing anal pouch, and in details of female genital anatomy. Eucosmomorpha nearctica occurs widely, having been collected
in Kentucky, Michigan, Mississippi, North Carolina, and Saskatchewan. Although E. nearctica was thought to be an immigrant in North America
when reported as E. albersana, it now seems more likely that it is a native insect that escaped earlier recognition.
Additional key words: E. albersana, E. a. ussuriana, E. nearctica, Olethreutinae, Eucosmini.
Eucosmomorpha, up to now comprising four Pa-
laearctic species, is a structurally distinct but poorly
known genus tentatively included in the olethreutine
tribe Eucosmini (Horak & Brown 1991). The new
species described here already has a publication his-
tory in North America. I reported one male, captured
in Michigan in 1961, as the Palaearctic E. albersana
(Hiibner) (Miller 1983). I noted that it might prove to
be E. albersana ussuriana (Caradja); Caradja’s (1916)
description was insufficient to permit a more definite
determination. Additional reports of the insect fol-
lowed from Saskatchewan, Kentucky, and Michigan
(Dang & Parker 1990, Gibson 1993). Because of re-
cent unpublished finds in North America, as well as in-
creased interest in immigrant insects, I undertook to
resolve the insect’s identity.
MATERIALS AND METHODS
Forewing length was measured under a binocular
microscope at nominal 10x magnification to within 0.2
mm with an eyepiece micrometer. Wing venation was
examined in reflected light under a binocular micro-
scope after touching xylene to wings. Genitalia slides
were prepared by standard methods, and genitalia
double stained with chlorozole black E and saffranin.
Specimens mentioned without genitalia slide number
are undissected.
Character states are included in the description
which place the new species in the genus Eucosmo-
morpha as defined by Obraztsov (1961).
Collection and museum abbreviations are as follows:
JBS, J. B. Sullivan, Beaufort, North Carolina; LDG, L.
D. Gibson, Florence, Kentucky; MEM, Mississippi
Entomological Museum, Mississippi State, Missis-
sippi; MGAB, Muzeul de Istorie Naturala “Grigore
Antipa,” Bucharest, Romania; MSU, Michigan State
University, East Lansing, Michigan; UMSP, University
of Minnesota Entomology Museum, St. Paul, Min-
nesota; USNM, National Museum of Natural History,
Washington, DC.
IDENTITY OF EUCOSMOMORPHA ALBERSANA USSURIANA
As the identity of E. a. ussuriana was unclear from
Caradja’s (1916) description, I obtained the holotype
for study. Examination showed that it did not differ
structurally from typical E. albersana, and that its
forewing scale pattern differed only trivially (Figs. 1,
2). Thus E. a. ussuriana seems to represent no more
than individual or geographic variation. Moreover,
Caradja (1916) mentioned a specimen intermediate in
scale pattern between E. a. ussuriana and the typical
form. These observations confirm the appropriateness
of Kuznetsov’s (1989) treatment of E. a. ussuriana as a
synonym of E. albersana.
The early stages of what Caradja described as E. a.
ussuriana are unknown, but E. albersana is univoltine,
overwintering as a mature larva, the larva feeding on
Lonicera and Symphoricarpos (both Caprifoliaceae)
(Bentinck & Diakonoff 1968, Bradley et al. 1979, Han-
nemann 1961, Kuznetsov 1987, 1989, Razowski 1987).
a &
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics. 1-7. Wings and genitalia of Eucosmomorpha species. 1, Wings of E. albersana ussuriana holotype °. 2, Wings of E. albersana ° from
Potsdam, Germany. 3. Wings of E. nearctica paratype 3 from Franklin Co., Mississippi. a, subbasal fascia. b, medial fascia. 4, Genitalia of E. al-
bersana 6 from “Geelskov” (genit. slide WEM 299993). 5, Genitalia of E. nearctica paratype d from Jones Co., North Carolina (genie slide
WEM 55995). The aedeagus appears shorter than actual because of angle of mounting. 6, Genitalia of E. a. ussuriana holotype 2. The smaller
signum is circled. 7, Genitalia of E. nearctica paratype ° from Boone Co., Kentucky (genit. slide WEM 1310991). The smaller signum is circled.
The ductus bursae and corpus bursae became severed during dissection and are out of their natural positions.
Published forewing lengths, after conversion from
spans by an empirically derived equation (Miller
1977), range 5-7 mm, averaging 6 mm (Bentinck &
Diakonoff 1968, Bradley et al. 1979, Hannemann
1961, Kuznetsov 1987, Razowski 1987).
Specimens examined. EF. a. ussuriana: Holotype 2
[sex incorrectly given as d in original description and
on pin], Kasakewitch, Ussuri R., E. Siberia, Korb,
5729 Wlsm. 1908, Grapholitha albersana var. ussuri-
ana Car., forewing length 6.0 mm, genit. slide WEM
289995 (MGAB): E. a. albersana: 1 6, “Geelskovy,” 4
May 1895, P. albersana Hb., V. Kuznetsov det., genit.
slide WEM 299993 (MGAB): 1 2, Potsdam [Ger-
many], Z. 18/388, Lonicera, Henneby, genit. slide
WEM 289994 (MGAB): 1 d, 25.5.1882, Hamfelt Coll.
[of known European origin], genit. slide WEM 299991
(USNM); 1 6, Kent [England], 6.1913, H. C. Hay-
ward, genit. slide WEM 299992 (USNM). At other
times I have examined additional specimens not re-
counted here.
OTHER EUCOSMOMORPHA SPECIES
Review of the literature reveals three other de-
scribed species of Eucosmomorpha besides E. alber-
sana, all Asian: E. multicolor Kuznetsov, E. magnifica
Kuznetsov, and E. figurana Kuznetsov (Kuznetsov
1964, 1997). Nothing is known of these species beyond
their taxonomic descriptions. However, it is evident
from the published descriptions and illustrations that
all differ in forewing scale pattern and genital anatomy
from both E. albersana and the new species described
here. For example, unlike the valvae of E. albersana
VOLUME 55, NUMBER 3
and the new species, that of E. multicolor has a distinct
pollex, that of E. magnifica is parallel sided, and that of
E. figurana tapers gradually between the sacculus and
cucullus (Kuznetsov 1964, 1997).
Eucosmomorpha nearctica W. E. Miller,
new species
Eucosmomorpha albersana (not Hiibner, 1822);
Miller (1983, 1987), Dang and Parker (1990),
Gibson (1993).
Male (n = 18). Head. Middle front and vertex
brownish orange, lower front white with shorter scales,
a band of brown scaling crossing vertex; antenna
brownish dorsally, darker ventrally, flagellar scales no
longer than flagellomere, pecten apparently absent;
labial palpus white basally and ventrally, terminal
segment short, ~ 0.25 length of second segment,
brown, second segment expanded distally, subequal in
length to vertical eye diameter, scaled with patches of
orange and brown, brown distally; proboscis subequal
in length to labial palpus. Thorax. Mesonotum and
tegulae brownish orange, sternum shining white, legs
shining white between coxa and tibia, tibia and tarsi
banded white and grayish brown, paler on inner sides.
Forewing (Fig. 3). Costal fold absent; upper side
mottled brown and orange; subbasal and median
fasciae (a, b, respectively, in Fig. 3) brown, angling
outwardly from costa, expressed mainly on costa, the
median fascia distinct also on dorsum near tornus; 6 to
10 short white costal strigulae angling outwardly,
separated in outer one-third of wing by sinuate orange
striae; speculum consisting of three blackish brown
longitudinal dashes; fringe brownish orange distally,
paler basally; underside of wing grayish brown.
Hindwing. Veins M, and Cu, connate, base of M,
slightly closer to base of M,, all three subparallel;
upper and under sides grayish brown, fringe paler
except for a grayish brown line near base; basal two-
thirds of anal angle with wing edge thickened and
bowed, forming a pouch that appears aligned with the
hind tibia when the wing is spread. Abdomen. Shining
brown dorsally, shining white ventrally. Genitalia (n =
7) (Fig. 5). Sacculus broad basally; a long, thick seta at
apex of cucullus; uncus absent; socii pulvinate and
directed upwards in an uncus position; aedeagus with
two sinuous cornuti.
Female (n = 1). Exterior essentially as described for
male, except for absence of the hindwing anal pouch.
Genitalia (n = 1) (Fig. 7). Lamella antevaginalis
absent; apophyses anteriores and posteriores subequal
in length; ductus bursae short, encircled at the opening
to the ductus seminalis by a sclerotized ring subequal
83
in width to ductus bursae diameter; two unequal sized
signa on corpus bursae, the larger one cone shaped.
Diagnosis. I found no consistent differences in
male genitalia between E. albersana and E. nearctica
(Figs. 4-5). The taxa are distinguishable by other
characters detailed below. In brief, E. nearctica has a
distinctive forewing scale pattern, is smaller in body
size, the anal area of the male hindwing is more
extensively modified, and the female genitalia differ in
length of the ductus bursae and other structural
details.
The forewing of E. nearctica is more or less mottled
throughout (Fig. 3 here, and Fig. 1 in Miller 1983),
whereas that of E. albersana is dark purplish on the
basal two-thirds, and mostly pale orange or yellowish
on the distal one-third, a combination that creates an
overall bicolored appearance (Figs. 1, 2 here and
illustrations in Bentinck & Diakonoff 1968, Bradley et
al. 1979, Hannemann 1961, Razowski 1987).
Forewing length in E. nearctica of the combined
sexes ranges 3.8-5.5 mm, averaging 4.6 mm (n = 19).
The 4.6 average is three-fourths the corresponding 6
mm value derived from the literature for E. albersana,
but translates into only one-half of the E. albersana
body mass (Miller 1977).
The anal edge of the E. nearctica male hindwing is
thicker and more bowed than that of E. albersana,
thus creating a more prominent hindwing anal pouch
in E. nearctica. The apparent difference between the
taxa in aedeagus length in Figs. 4 and 5 is an artifact of
slide mounting absent in other preparations.
The ductus bursae in E. nearctica is only half as long
as that in E. albersana, is ringed with a sclerotized
band at the opening to the ductus seminalis which E.
albersana apparently lacks, and the smaller signum of
E. nearctica is larger than that of E. albersana (Figs. 6,
7). The smaller signum of E. albersana is but a speck
and is easily overlooked.
Types. Holotype ¢: Mississippi, Franklin Co., Trib.
of McGehee Crk., T6N, R4E, Sec. 26 SW, 31 Aug.
1992, J. MacGown, T. Schiefer, forewing length 4.9
mm, genit. slide WEM 299995 (USNM). Paratypes:
KENTUCKY: 1 2, Boone Co., Big Bone Lick State
Park, 4 Aug. 1989, L. D. Gibson, genit. slide WEM
1310991 (LDG); 1 4, same data, except 9 July 1991,
genit. dligle ILIDXE 102 (CLD). IMNCTSUIGANINe ilo,
Midland Co., 2 June 1961, R. R. Dreisbach, genit.
slide PJ 163 (MSU); 1d, Otsego Co., 13 June 1988, L.
D. Gibson, genit. slide LDG 095 (LDG).
MISSISSIPPI: 1 d, Scott Co., Bienville Natl. For.,
Caney Crk. Wildlife Mgt. Area, 2 mi [3.2 km] E.
Pulaski, 10 June 1988, D. & M. Hildebrandt, genit.
slide WEM 299994 (MEM): 1 ¢d, same data as
84
holotype (MEM). NORTH CAROLINA: 3 d, Jones
Co., N. of Stella, Haywood Landing, Croatan Natl.
For., hardwoods, 15-watt U-V trap, 18 July 1998, J. B.
Sullivan (JBS, UMSP, USNM); 2 d, same data as
preceding, except 2 Aug. 1997 (JBS, UMSP); 1 d, same
data as preceding, except genit. slide WEM 59995
(JBS); 1d, Jones Co., Island Walk, Croatan Natl. For.,
hardwoods, 15-watt UV trap, 17 June 1998, J. B.
Sullivan (JBS); 1 d, same data as preceding, except 30
April 1997 (JBS); 1 3, Craven Co., Croatan Natl. For.
Rd. 167, 21 June 1993, J. B. Sullivan, genit. in vial on
pin (JBS); 1 d, same data as preceding, except Rd.
3046, Gum Branch Rd., 25 April 1998 (JBS); 1 ¢,
Pender Co., Holly Shelter gamelands, 15-watt UV
trap, pine savannah, 26 August 1997, J. B. Sullivan
(USNM). SASKATCHEWAN: 1 ¢, Saskatoon, phero-
mone trap, 1984, Chisoholm (USNM).
DISCUSSION
Specimen and literature records of E. nearctica are
widely distributed: Kentucky, Michigan, Mississippi,
North Carolina, and Saskatchewan. Capture dates
from combined localities range from 25 April to 31
August, suggesting one to two generations per year.
Larval foodplants are unknown.
It is possible that E. nearctica is an immigrant in
North America as supposed when it was reported as E.
albersana (Miller 1983). However, a more straightfor-
ward interpretation of the information assembled here
is that it is a native American species that escaped pre-
vious recognition because of low population densities,
sparse collecting, and diminutive size. The species is
not known anywhere else than in North America, and
collection localities are inland, away from commercial
ports where immigrants usually are detected first.
ACKNOWLEDGMENTS
I thank D. Rusti (MGAB) for loaning the E. a. ussuriana holo-
type; J. W. Brown (USNM), R. L. Brown (MEM), L. D. Gibson
(LDG), and J. B. Sullivan (JBS) for loaning specimens in their care;
R. W. Holzenthal for use of photomicrographic equipment; R. L.
Brown for drawing my attention to recently collected specimens of
the new species which prompted this study; and J. W. Brown, R. L.
Brown, M. Sabourin, J. B. Sullivan, and L. D. Gibson for useful
manuscript reviews.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
LITERATURE CITED
BENTINCK, G. A., GRAAF & A. DIAKONOFF 1968. Nederlandse
Bladrollers (Tortricidae). Monogr. Nederland. Entomol.
Vereen. 3, 200 pp.
BRADLEY, J. D., W. G. TREMEWAN & A. SmiTH. 1979. British tortri-
coid moths, Tortricidae: Olethreutinae. Ray Society, British
Museum (Natural History), London. 336 pp.
CarapjA, A. 1916. Beitrag zur Kenntnis der geographischen Ver-
breitung der Pyraliden und Tortriciden des europaischen
Faunengebietes, nebst Beschreibung neuer Formen. Deut. En-
tomol. Z. “Iris” 30:1-88.
Danc, P. T. & D. J. PARKER. 1990. First records of Enarmonia for-
mosana (Scopoli) in North America (Lepidoptera: Tortricidae).
J. Entomol. Soc. Brit. Columbia 87:3-6.
Gipson, L. D. 1993. Additional North American records of Eucos-
momorpha albersana (Tortricidae: Enarmoniini). J. Lepid. Soc.
47:322-323.
HANNEMANN, H. J. 1961. Kleinschmetterlinge oder Microlepi-
doptera I. Die Wickler (s. str.) (Tortricidae). Pt. 48, Die Tier-
welt Deutschlands und der angrenzenden Meeresteile. Fischer,
Jena. 236 pp.
Horak, M. & R. L. Brown. 1991. Taxonomy and phylogeny, pp.
93-50. In L. P. S. Van Der Geest & H. H. Evenhuis (eds.), Tor-
tricid pests: their biology, natural enemies and control. Elsevier,
New York. 808 pp.
Kuznetsov, V. I. 1964. New genera and species of leafrollers (Lep-
idoptera, Tortricidae) from the Far East. Entomol. Rev.
43:445—-453.
. 1987. Family Tortricidae (tortricid moths), pp. 279-956.
In G. S. Medvedev (ed.), Keys to the insects of the European
part of the USSR (translation). U.S. Dept. Agr. & National Sci-
ence Foundation. 991 pp.
. 1989. Leaf-rollers (Lepidoptera, Tortricidae) of the south-
ern part of the Soviet Far East and their seasonal cycles, pp.
57-249. In O. L. Kryzhanovskii (ed.), Lepidopterous fauna of
the USSR and adjacent countries. Brill, Leiden. 405 pp. [trans-
lation].
. 1997. New species of tortricid moths of the subfamily
Olethreutinae (Lepidoptera, Tortricidae) from the South of
Vietnam. Entomol. Rey. 77:715—727.
MILLER, W. E. 1977. Wing measure as a size index in Lepidoptera:
The family Olethreutidae. Ann. Entomol. Soc. Amer. 70:253-256.
. 1983. Eucosmomorpha-albersana (Hiibner), a Palaearctic
species, collected in North America (Tortricidae, Grapholitini).
J. Lepid. Soc. 37:88-89.
. 1987. Guide to the olethreutine moths of midland North
America (Tortricidae). U. S. Dept. Agr. Agr. Handb. 660. 104 pp.
Opraztsov, N.S. 1961. Die Gattungen der palaearktischen Tortri-
cidae. II. Die Unterfamilie Olethreutinae. Part 4. Tijd. Ento-
mol. 104:51—70.
RazowskI, J. 1987. Motyle (Lepidoptera) Polski 7—Uzupelnienia I
Eucosmini. Monogr. Fauny Polski 15. 253 pp.
Journal of the Lepidopterists’ Society
55(3), 2001, 85-100
AN OVERVIEW OF STRYMON HUBNER (LYCAENIDAE: THECLINAE: EUMAEINI)
ROBERT K. ROBBINS
Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0127, USA
AND
STANLEY S. NICOLAY
1500 Wakefield Drive, Virginia Beach, Virginia 23455, USA
ABSTRACT. North American Strymon Hiibner was revised about 40 years ago, and the significantly larger Neotropical Strymon is now
incorporated into this classification. Strymon is characterized by anteriorly directed teeth on the posterior dorsal surface of the valvae. These
teeth were first noted by Clench and appear to be modifications of the sockets of setae that normally occur on eumaeine valvae. This charac-
terization is most consistent with past usage and appears to represent the best evidence for monophyly. As characterized, Strymon contains 48
described species. Variation in morphology of the male genitalia, female genitalia, wings, and head is documented, and male behaviors and lar-
val foodplant records are summarized. We tentatively divide Strymon into species groups, one of which is unusual in its use of Bromeliaceae as
its sole larval foodplant.
One recently described genus, Heoda Johnson, L. Miller & Herrera 1992, and one recently resurrected genus, Eiseliana Toledo 1978, are
made junior synonyms of Strymon Hiibner 1818. Six names recently described in Strymon are transferred to other genera: Strymon angulus Le
Crom & Johnson, 1997 to Thereus Hiibner; Strymon daplissus Johnson & Salazar, 1993 to Ministrymon Clench 1961; Strymon carmencitae Le
Crom & Johnson, 1997 and Strymon cryptogramus Johnson, Eisele & MacPherson, 1992 to Nicolaea Johnson; Strymon nivnix Johnson, Eisele
& MacPherson, 1990 to Calycopis Scudder; and Strymon additionalis Le Crom & Johnson, 1997 to Thecla F. The hindwings of Strymon nivnix
are designated a lectotype, and Strymon anthracaetus Salazar, Vélez, & Johnson, 1997 is regarded as a nomen dubium.
Additional key words: Bromeliaceae, foodplants, territoriality, Heoda, Eiseliana.
Strymon Hiibner is possibly the best-known New
World hairstreak genus (Lycaenidae: Theclinae: Eu-
maeini). It occurs from Canada to the temperate parts
of Chile and Argentina. Some Strymon are common
and well-known, such as S. melinus (Gray Hairstreak),
and some are pests on commercial pineapple, such as
S. ziba (Hewitson), S. megarus (Godart), and relatives
(Harris 1927, Carter 1934, Fonseca 1934, Zikan
1956, Guagliumi 1965, 1967, D’Araujo e Silva et al.
1967-1968, Beutelspacher 1972, Otero & Marigo
1990). The name Strymon has been widely used in
North America; the first extensive list of North
American Strymon species (Barnes & McDunnough
1917) contained about 40 taxa and was followed by
similar listings (Barnes & Benjamin 1926, McDun-
nough 1938, Klots 1951, Dos Passos 1964). Although
Ziegler (1960) and Clench (1961) rather drastically
changed the characterization of Strymon, 14 of the
48 described Strymon species currently recognized
(Appendix 1) are recorded from North America
(Opler & Malikul 1992, Opler & Wright 1999).
“Modern” taxonomic usage of Strymon began when
the genus was distinguished primarily by genitalic
structures (Ziegler 1960, Clench 1961). With in-
creased knowledge of the Neotropical eumaeine
fauna, however, it became clear that these characters,
as originally proposed, do not delimit Strymon. For
example, S. yojoa (Reakirt) has small anteriorly di-
rected teeth on the dorsal valva tips—a structure
mentioned in Clench’s generic diagnosis—but lacks a
tightly convoluted spiral of the ductus bursae (Fig.
20)—a structure noted in Ziegler’s generic character-
ization. Alternately, S. serapio (G. & S.) has the
tightly-convoluted spiral (Fig. 25), but also has a dou-
ble cornutus, not the single acuminate one described
by Clench (1961) (Fig. 16). To complicate matters,
the subsequently described genera Eiseliana Toledo
and Heoda Johnson, Miller, & Herrera possess some
genitalic structures that Ziegler and Clench used to
characterize Strymon. Finally, six species described in
Strymon since 1990 possess none of these characters.
The purposes of this paper are to characterize Stry-
mon, so that it will be clear which species belong to
Strymon, and to provide an overview of the compara-
tive morphology and ecology of the genus. Specifically,
this paper (1) outlines the nomenclatural history of
Strymon, (2) suggests that the best structure for dis-
tinguishing Strymon is the unique morphology of the
male genitalia valvae, which was first noted by Clench,
(3) describes and illustrates morphological variation
within the genus, (4) summarizes information on male
behavior, larval foodplant specificity, and habitat, (5)
preliminarily partitions Strymon species in nine
species groups, and (6) transfers six names from Stry-
mon to other genera. This work is intended to set the
stage for a species revision, including the description
of about five new species, mostly from the dry moun-
tains of Peru and southern Ecuador.
86
MATERIALS AND METHODS
The results in the this paper were based upon a
comparison of adult morphology using the 6,000+
specimens of Strymon in the National Museum of
Natural History (Smithsonian Institution, Washington,
DG, USA), of which 3,972 are Neotropical, plus many
specimens borrowed from other museums. This com-
parison employed standard entomological techniques
(Robbins 1991), including the examination of the male
and female genitalia (449 dissections) of all species
currently recognized im Strymon (Appendix 1), except
that we relied on genitalic figures of two species. For
those names that we could not identify from their
original descriptions, we examined their types or pic-
tures of the types. For only one species, Strymon an-
thracaetus Salazar, Vélez, & Johnson, could we not
identify the name or find its type (explained below). In
preparing a checklist of all Neotropical hairstreaks
(Robbins in press), RKR examined the adult morphol-
ogy of virtually all Neotropical species, although not in
the same detail as with Strymon. All genitalic terms
follow those in Klots (1970). All specific author names
for Strymon are listed in Appendix 1 and thus are
omitted from the following text.
Because relationships within the Eumaeini are still
poorly known, such as the genera that are most closely
related to Strymon, we characterize Strymon by a
complex and conspicuous trait that is unique within
the Eumaeini and that is phylogenetically consistent
with other traits that are unique within the Eumaeini.
We tentatively divide Strymon into species group on
the basis of many characters, but evidence for their
monophyly awaits formal phylogenetic analysis.
NOMENCLATURAL HISTORY
Hiibner (1818) described Strymon and included
two species, S. melinus and Hesperia acaciae Fabri-
cius. (Hesperia currently belongs to the Hesperiidae.)
A subsequent list of 13 Strymon species (Hiibner
1819) caused considerable confusion in the eventual
selection of a type species. Scudder (1872) selected
Hesperia titus F. from the 1819 list as the type (the
dates of Hiibner’s books were uncertain at the time),
but Riley (1922) invalidated this selection and re-
placed it with Strymon melinus. Finally, the Interna-
tional Commission on Zoological Nomenclature
(1959) placed Strymon on the Official List as Name
No. 1332 with Strymon melinus as type. Hemming
(1967) gives a more complete nomenclatural history.
STRYMON HUBNER
We characterize Strymon for the following discus-
sion of comparative morphology by setae on the poste-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
rior dorsal surface of the valvae that are modified into
anteriorly pointing “teeth” (Clench 1961) (Figs. 1-7).
This structure is immediately recognizable; either a
species has it or does not. The only exception is S. ziba
(Fig. 8), whose valva morphology and systematic posi-
tion are detailed below. Although the valva structure of
Strymon is most easily illustrated using a scanning
electron microscope (SEM), Clench discovered it us-
ing a light microscope. Almost all species that have
been placed in Strymon since 1960 (e.g., Ziegler 1960,
Clench 1961, 1964, Johnson et al. 1990, Johnson et al.
1992, Johnson & Kroenlein 1993, Johnson & Salazar
1993, Austin & Johnson 1997) have this valva struc-
ture. The only superficially similar valvae in the Eu-
maeini are those of Allosmaitia Clench, whose valvae
are needle-like posteriorly, unlike Strymon, and cov-
ered with teeth that are not anteriorly directed
(Clench 1964).
COMPARATIVE MORPHOLOGY
Male genitalia. The setae on the valvae of eu-
maeine hairstreaks are indistinguishable when viewed
with an SEM from those specialized setae that were
termed B-type trichoid sensilla (Ma & Schoonhoven
1973). In Strymon, these setae are modified. The an-
teriorly directed teeth on the posterior dorsal surface
of the valvae appear to be modifications of the sockets
of these setae. Supporting this interpretation, the setal
flagellum is still present, but is short (usually less than
1 um long) (Figs. 1-7). In S. ziba, some of the setal
sockets on the posterior dorsal valva tips are slightly
modified into anteriorly pointing teeth with a flagellum
that is not reduced in length (Fig. 8). As discussed be-
low, some evidence suggests that this structure is tran-
sitional between that in other eumaeines and in Stry-
mon while other evidence suggests that it is a further
modification of that which occurs in other Strymon.
We illustrate the male genitalia of nine Strymon
species to show the range of morphological variation
(Figs. 9-17). The tips of the gnathos in all Strymon
species are subterminally expanded and then sharply
tapered to a down-curved point (Figs. 9-17), but this
structure does not appear to be sufficiently distinct
from other eumaeines to distinguish the genus un-
equivocally. At least one cornutus is present within the
shaft of the penis unless the vesica is partially everted
(Figs. 12, 14, 15). If there is one cornutus, it is long
and slender (slightly wider in S. maritalis and S.
christophei) and sometimes barely sclerotized (S. yo-
joa, Fig. 11). If there are two cornuti, they are usually
paired and about the same size (Fig. 16) (first noted
and illustrated by Schwartz & Miller 1985). The pri-
mary exception is S. ziba, which has one cornutus con-
VOLUME 55, NUMBER 3 87
Fics. 1-8. Scanning electron micrographs (SEMs) of male genitalia valva tips in lateral and dorsal aspects. 1, S. melinus in lateral aspect; 2,
S. melinus at lower magnification; 3, S. acis; 4, S. ewrytulus in dorsal aspect (Arrow A — setal flagellum); 5, S. bazochii; 6, S. gabatha; 7, S.
megarus; 8, and S. ziba (Arrow B — slightly modified setal socket).
|
|
{)
|
t
88
siderably larger than the other (Fig. 17). Strymon male
genitalia are asymmetrical. For example, the penis is
twisted and down turned (Clench 1961) except for S.
ziba and S. sylea (Figs. 10, 17). The saccus is generally
asymmetrical to the right (Figs. 9b, 12, 14, in ventral
aspect, they are on the left side).
Strymon species have paired brush organs (sensu
Eliot 1973) that lie on the dorsal vinculum, but we
omit them in the figures for clarity. The vinculum is
not modified in structure, as it is in some eumaeines
(Robbins 1991), except for slight projections of the
vinculum in some species, such as S. tyleri and S.
crambusa. We are unable to distinguish the structure
of the brush organs in Strymon from those that occur
in Lamprospilus Hiibner, Electrostrymon Clench,
Ziegleria Johnson, and Calycopis Scudder. Males of S.
istapa in one part of its range (Florida to Puerto Rico)
may have or lack brush organs (Robbins & Nicolay
1999), and a similar dimorphism also appears to occur
in S. bicolor.
Other than the variation outlined above, the male
genitalia of Strymon seem to have few good structures
for distinguishing species. For example, the male gen-
italia of S. istapa, S. columella, S. limenia, and S. tous-
sainti are essentially indistinguishable except for small
differences in size (Robbins & Nicolay 1999) — these
species are distinguished by their wing pattern and fe-
male genitalia. Except for S. ziba and S. sylea, few Stry-
mon species appear to be authoritatively identifiable
solely on the basis of their male genitalic structures.
Female genitalia. The bursa copulatrix of Strymon
is asymmetrical, and we illustrate the range of varia-
tion (Figs. 18-26). All eumaeines with a sclerotized,
looped ductus bursae belong to Strymon., as first noted
by Ziegler (1960), but the exact shape of the loop
varies greatly interspecifically (Figs. 18-26). It is also
highly variable intraspecifically, as in S. cestri, where
some individuals lack the sclerotized loop of the duc-
tus bursae, some have it, and others are intermediate
between these extremes (Figs. 27-29). A few Strymon
species appear to always lack the loop (S. yojoa, S.
tegea, S. ohausi, S. sylea, and S. ziba), but the ductus
bursae is twisted at the point where the loop would
otherwise occur except in S. sylea (Figs. 19, 20, 23).
The signa, which are “boat”-shaped, occur in all Stry-
mon species, but may be small in some, such as S. li-
menia (Hewitson) (Fig. 24), and may lack the anterior
pointing spine (S. sylea, Fig. 19). Similar signa are
found on occasion in other eumaeines, such as Tricho-
nis Hewitson (Robbins 1987). There are two small
teeth inside the anterior ductus bursae (Fig. 18). Sim-
ilar teeth occur in other eumaeines, such as those il-
lustrated for Rekoa palegon (Cramer) (Robbins 1991).
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
The female Sth abdominal tergum of Eumaeini is
sclerotized, the sternum is membranous, and two
small circles lacking setae are also membranous (ex
illustrated by Field 1941). Since spiracles on the 8th
abdominal segment of the endoporian Ditrysia, in-
cluding the butterflies, are absent or vestigial (Dug-
dale 1974), we presume that these membranous cir-
cles are vestigial spiracles. In female Strymon, there
are two kinds of 8th abdominal segments. The first, of
which S. melinus is an example (Fig. 30), is like many
eumaeines with presumed vestigial spiracles at the
juncture between the tergum and sternum. The sec-
ond, of which S. bicolor is an example (Fig. 31), has
the presumed vestigial spiracles located within the ter-
gum, whose latero-posterior part has more furrows
than the first kind. Johnson et al. (1992) first described
the furrowed tergum as “rough” and used it as a distin-
guishing character of their new genus Heoda. How-
ever, this structure is difficult to discern and actually
occurs in all members of the S. istapa group (except
for S. acis, Appendix 1), including species that Johnson
et al. (1992) placed in Eiseliana and Strymon.
When the ductus seminalis does not arise from the
posterior tip of the corpus bursae in Strymon, such as
S. limenia, the tip of the corpus bursae, posterior of
the ductus seminalis, is lightly sclerotized (Fig. 24). All
Strymon species with this structure are a subset of
those whose females have an 8th tergum with the lat-
ero-posterior surface furrowed and with imbedded
presumed vestigial spiracles. The posterior end of the
corpus bursae of a few other species, such as S. ziba, S.
martialis, and S. christophei, is also sclerotized (Figs.
23, 26), but this structure appears to be analogous with
the structure in S. limenia and relatives because the
ductus seminalis arises from the posterior tip of the
corpus bursae in these species.
Wings. The wing venation of Strymon is typical of
the Eumaeini with 10 forewing veins (Eliot 1973). A
male scent patch (sensu Robbins 1991) occurs on the
dorsal surface of the forewing in the distal part of the
discal cell of most species, but is lacking in S. melinus,
S. avalona, S. sabinus, S. tyleri, S. rufofusca, S. cyano-
fusca, S. ohausi, S. christophei, S. oribata, and S.
legota. Because these species appear to belong to a
number of different species groups, as determined be-
low on the basis of many characters, we suspect multi-
ple losses of the forewing scent patch in Strymon.
Wing pattern, shape, and size vary greatly among
Strymon species (Figs. 34-53), and it is difficult to
characterize Strymon on the basis of these traits. Gen-
erally, the pattern on the ventral wing surface is not
sexually dimorphic, but males have more sharply pro-
duced forewing apices. Seasonal variation of the ven-
VOLUME 55, NUMBER 3
16
89
Fics. 9-17. Male genitalia of Strymon; teeth on valvae are inconspicuous at this magnification. 9, S. melinus in lateral (a) and ventral (b) as-
pects (Arrow D — acuminate cornutus); 10, S. sylea (Arrow C — tip of gnathos); 11, S. yojoa; 12, S. mulucha; 13, S. albata; 14, S. martialis:
15, S. limenia; 16, S. serapio (Arrow E — paired cornuti); and 17, S. ziba.
tral wing surface can be marked in some species, such
as S. melinus, but virtually absent in others, such as S.
bazochii. For example, individuals of S. melinus are
smaller and darker on average in early spring in North
America than they are in the middle of summer. The
dorsal surface of the forewing of most female Strymon
has a patch of black scales centered at the end of the
discal cell, which is sometimes mistaken for androco-
nia (Figs. 35, 40). Other eumaeines sometimes also
have a similar appearing patch of dark scales, such as
females of Tmolus venustus (Druce). Forewing length
varies from more than 2 cm (S. sylea, S. oreala, and S.
gabatha) to less than 0.8 cm (S. ohausi, S. ochraceus).
And some species, such as S. gabatha and S. serapio,
vary greatly in size intraspecifically, which is perhaps
related to their bromeliad larval foodplants (flowers of
Aechmea and Tillandsia, respectively).
Head. The antennal club of Theclinae, including
the Eumaeini, is generally cylindrical and incrassate,
but those of most Strymon are abrupt and flattened,
resembling those of Polyommatinae and Lycaeninae
(Eliot 1973). An antenna with an abrupt and flattened
club also occurs in a few other eumaeines, such as
Penaincisalia Johnson. The frons of S. melinus is cov-
ered with white and orange scales, and there are also
orange scales near the base of the antennae. Most
other Strymon share this head coloration, but a few
lack white or orange scales, such as S. bazochii (Go-
dart). However, a hairstreak with orange scales near the
base of the antennae, an orange-white frons, and an
abrupt, flattened antennal club is almost definitely a
Strymon, which allows field recognition of this genus in
most cases.
BEHAVIOR AND ECOLOGY
Male behavior. Male Strymon occupy mating ter-
ritories, usually on hilltops and along forest edges. The
males perch in the “territory,” fly at other butterflies
that enter this area, and then return to a perch very
close to the original one unless courtship has ensued
(e.g., Powell 1968, Robbins 1978, Alcock 1983, Alcock
& O'Neill 1986, 1987, Cordero & Soberén 1990). We
90
24a 24b
Fics. 18-26. Female genitalia bursa copulatrix of Strymon. 18, S.
sylea; 20, S. yojoa in ventral (a) and lateral (b) aspects; 21, S. mulucha (
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
melinus in ventral (a) and lateral (b) aspects (Arrow F — signum); 19, S.
Arrow G — teeth inside ductus bursae); 22, S. albata; 23, S. martialis; 24,
S. limenia in ventral (a) and lateral (b) aspects; 25, S. serapio in ventral (a) and lateral (b) aspects; and 26, S. ziba in ventral (a) and lateral (b) aspects.
VOLUME 55, NUMBER 3
27 imm
SEE
Fics. 27-29. Female genitalia variation of the bursa copulatrix
of S. cestri. 27, Peru; 28, Costa Rica; 29, Mexico.
have observed this behavior in the United States,
Panama, Venezuela, Ecuador, Peru, and Brazil for 15
Strymon species (RKR unpubl.). The males of most
species occupy mating territories in the afternoon. A
few species do so whenever the weather is favorable
(i.e., S. tyleri, S. davara), and one species only in the
early morning (males of S. serapio on hilltops in
Panama and southern Brazil between 0730 and 1000
hours). The following discussion is based on our un-
published observations.
When male Strymon perch in a mating territory,
they often open their wings 15-180 degrees, in addi-
tion to moving their hindwings back and forth as in all
Theclinae (Robbins 1980). We have observed the un-
usual wing opening behavior (Figs. 54, 55) in S. meli-
nus, S. mulucha, S. yojoa, S. davara, S. cestri, S.
bubastus, S. dindus, S. ochraceus, S. ziba, S. megarus,
S. azuba, and S. gabatha. Males of other hairstreak
genera generally do not open their wings when perch-
ing, but we have observed this behavior in some
species of Tmolus and Chlorostrymon. Another un-
usual male Strymon behavior is perching head down-
wards on tree-trunks, in addition to perching on
leaves. This behavior has been recorded in S. ziba, S.
gabatha, S. ochraceus, and S. megarus. Only once have
we seen a male hairstreak of another genus perch on a
tree-trunk (Chalybs janias |Cramer]).
Larval foodplants. The larvae of Strymon eat
plants in more than 30 families, ranging from gym-
nosperms to monocots and dicots, including Alstroe-
91
meriaceae, Amaranthaceae, Begoniaceae, Boragi-
naceae, Bromeliaceae, Cactaceae, Cannabidaceae,
Chenopodiaceae, Compositae, Convolvulaceae, Crassu-
laceae, Euphorbiaceae, Flacourtiaceae, Gesneriaceae,
Gramineae, Guttiferae, Haemodoraceae, Juglandaceae,
Labiatae, Leguminosae, Malvaceae, Melastomataceae,
Musaceae, Orchidaceae, Pinaceae, Polygonaceae, Por-
tulacaceae, Rosaceae, Sapindaceae, Sterculiaceae,
Strelitziaceae, Surianaceae, Ulmaceae, and Verbe-
naceae (RKR unpubl. plant family names follow Willis
1973). Some individual Strymon species, such as S.
melinus, are exceedingly polyphagous (Ehrlich &
Raven 1965, Tietz 1972), eating plant reproductive
structures in most of these families. Many of these
Strymon species, though, most frequently feed on
plants in the Leguminosae and Malvaceae, and some
are recorded as pests of beans and cotton (Ehrlich &
Raven 1965).
The only Eumaeini with larvae that eat plants in the
Bromeliaceae belong to Strymon (RKR unpubl.).
Some are serious pests of commercial pineapple (Har-
ris 1927, Carter 1934, Fonseca 1934, Zikan 1956, Gua-
gliumi 1965, 1967, D’Araujo e Silva et al. 1967-1968,
Beutelspacher 1972, Otero & Marigo 1990), but the
agricultural literature refers to lycaenid pineapple
pests as either Thecla echion L. (a misidentification of
S. megarus or S. ziba, cf. Honey & Scoble 2001) or
Thecla basalides (Geyer), a misspelling of Strymon
basilides (Geyer), which, in turn, is a junior synonym
of S. megarus (Appendix 1). Consequently, it is unclear
exactly how many Strymon species feed on pineapples,
but there are records for at least four species. The lar-
vae of S. ziba, unlike those of the others, eat plants in
a number of monocot families in addition to those of
the Bromeliaceae (e.g., Harris 1927, Robbins & Aiello
1982).
Habitat and range. Although Strymon occur in
habitats ranging from tropical wet lowland rainforest
to temperate climates, they are most diverse in xeric
and seasonally very dry tropical areas, which includes
most of the Pacific Coast of Mexico and Central
America, northern Colombia and northern Vene-
zuela, the mountains of Peru and southern Ecuador
(where there are a number of undescribed species),
the llanos of central Venezuela and surrounding
countries, and eastern Brazil in the cerrado and
caatinga life zones west to the Bolivian chaco. Stry-
mon melinus is the only widespread Strymon in tem-
perate North America while S. bicolor ranges from
Peru in the Andes to Chile’s temperate central valley
(Santiago and surroundings), and S. ewrytulus occurs
from Bolivia and southern Brazil south into Patagonia
(Argentina).
Fics. 30-33. SEMs of female 8th abdominal tergum in dorso-lateral aspect. 30, S. melinus; 31, S. bicolor; 32, S. eurytulus (Arrow I — fur-
row in tergum); 33, S. bazochii. Arrows H — presumed vestigial spiracles.
STRYMON AND ITS SPECIES GROUPS
The monophyly of Strymon, as we have character-
ized it, is supported by a complex and conspicuous
valva structure that is unique within the Eumaeini and
whose presence or absence is unambiguous except for
S. ziba. Other characters that are unique, or nearly so,
within the Eumaeini are restricted to subsets of Stry-
mon, adding further support to our characterization of
Strymon as a monophyletic genus. All species with a
scleortized looped ductus bursae belong to Strymon,
as do all eumaeines with a ventro-lateral surface of the
female 8th abdominal tergum that is furrowed with
imbedded presumed vestigial spiracles. Behaviorally
and ecologically, almost all eumaeine males that perch
with their wings open 15-180° belong to Strymon (a
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
behavior that may occur in all Strymon), as do all eu-
maeine males that perch on tree-trunks (one exception
noted above), and all species whose larvae eat plants in
the Bromeliaceae.
We have divided Strymon into nine species groups
(listed in Appendix 1) on the basis of many characters,
most of which were discussed above. Some of these
groups, such as the S. istapa and S. serapio groups, are
reasonably well-characterized while others, such as the
S. mulucha and S. albata groups, lack clear-cut derived
distinguishing traits. For example, some characters
suggest that S. cestri belongs to the S. yojoa species
group, and others that it belongs to the S. mulucha
species group. In the following paragraphs we charac-
terize the S. istapa and S. serapio species groups and
discuss the systematic positions of S. ziba and S. sylea.
VOLUME 55, NUMBER 3
93
Fics. 34-43. Dorsal and ventral wing surfaces. 34, male S. melinus (New Jersey, USA) (Arrow J — patch of black scales on female dorsal
forewing); 35, female S. melinus (New Jersey, USA); 36, male S. sylea (Loreto, Peru); 37, male S. yojoa (Panama Prov., Panama); 38, male S.
mulucha (Canal Area, Panama); 39, male S. albata (Canal Area, Panama); 40, female S. martialis (Florida, USA); 41, male S. limenia (Santiago,
Cuba); 42, male S. serapio (Santa Catarina, Brazil); and 43, male S. ziba (Canal Area, Panama).
We characterize those species with paired cornuti
that are about the same size as the Strymon serapio
group (Appendix 1). All recorded larval foodplants in
this group are plants in the Bromeliaceae. In fact, ex-
cept for S. ziba, all neotropical lycaenid larvae that use
Bromeliaceae, including all pests of commercial pineap-
ple, belong to this group. Except for S. ziba and one ob-
servation of a male Chalybs, all males that are known to
perch on tree-trunks belong to the S. serapio group.
We characterize the Strymon istapa group as those
species with a female 8th abdominal tergum whose
ventro-lateral surface is furrowed with imbedded pre-
sumed vestigial spiracles (Appendix 1). This tergum
structure is otherwise unreported in the Eumaeini.
Those species with a lightly sclerotized corpus bursae
between the origin of the ductus seminalis and the
posterior tip of the corpus bursae are a subset of this
group. In all other Strymon, the ductus seminalis
arises at the posterior tip of the corpus bursae. The in-
clusion of S. acis in this species group is provisional, as
explained in Appendix 1.
The systematic position of S. ziba is unresolved be-
cause of conflicting evidence. On the one hand, S. ziba
appears to belong to the S. serapio group. Both have
two cornuti, larvae that eat plants in the Bromeliaceae,
and males that perch on tree trunks. The ventral wing
pattern and male behavior of S. ziba and S. megarus
are nearly identical, suggesting that they are sisters. If
this systematic position is correct, then the unique
valva structure of S. ziba is a modification of the Stry-
mon valva. On the other hand, S. ziba appears to be
the sister to the remainder of Strymon. Evidence sup-
porting this systematic position is that the valva struc-
ture of S. ziba appears to be intermediate between
that of Strymon and other eumaeines. And unlike the
S. serapio group, S. ziba has paired cornuti of unequal
size, and its larvae eat plants in various monocot fami-
lies, not just Bromeliaceae. This evidence suggests that
S. ziba does not belong to the S. serapio group. Han-
ner and Robbins (in prep.) are trying to resolve this
conflicting evidence using mitochondrial DNA se-
quences.
94
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics. 44-45. Males perching with their wings open. 54, S. melinus (California, USA); 55, S. megarus (Veracruz, Mexico).
The systematic position of S. sylea is tentative be-
cause its penis is upturned and its ductus bursae is
straight, unlike virtually all other Strymon species. The
male genitalia valvae of S. sylea possess the unusual
basally directed teeth of Strymon when viewed with a
light microscope. The tips of the gnathos, scent patch,
and antennal club of S. sylea are consistent with Stry-
mon, and we cannot place S. sylea in any other eu-
maeine genus. Because S. sylea is an exceedingly rare
species in collections, we have not had the opportunity
to examine its valvae using an SEM, which would de-
finitively confirm that the valva teeth are indeed the
Strymon type. The placement of S. sylea in Strymon
appears to be the best option for now.
NOMENCLATURE
Generic synonymy. Using our characterization of
Strymon, based largely on that of Clench (1961), there
are five generic synonyms of Strymon, listed with their
type species in parentheses. Citations to all original de-
scriptions for this and the following list can be found in
Lamas et al. (1995).
Strymon Hiibner 1818 (Rusticus melinus Hiibner)
Callipareus Scudder 1872 (Strymon melinus Hiibner)
Callicista Grote 1873 (Callicista ocellifera Grote)
Uranotes Scudder 1876 (Strymon melinus Hiibner)
Eiseliana Toledo 1978 (Eiseliana koehleri Toledo)
Heoda Johnson, L. Miller & Herrera 1992 (Thecla heodes Druce)
Our characterization of Strymon appears to be
reasonable despite the uncertain systematic position of
S. ziba. If S. ziba belongs to the S. serapio group,
which is supported by some evidence, then S. ziba
clearly belongs to Strymon. If, on the other hand, S.
ziba is the sister to the remainder of Strymon, which is
supported by other evidence, then placing S. ziba in
Strymon avoids naming a monotypic genus for S. ziba.
The generic names Callicista, Eiseliana, and Heoda
could be applied to the S. istapa species group, but we
believe that our characterization of Strymon is more
reasonable and stable. Recognizing Callicista, Eise-
liana, or Heoda would leave the name Strymon for the
remainder of species with anteriorly directed teeth on
the dorsal valvae, but there is no evidence that this re-
maining group of species is monophyletic. And if a
generic name were also proposed for the S. serapio
group, there would still be no evidence for the mono-
phyly of Strymon. Lastly, our characterization of Stry-
mon is consistent with the way that Strymon has been
used in North America for the last 40 years (Ziegler
1960, Clench 1961).
Species groups. We list the 183 names that belong
to Strymon as we have characterized it, partitioned
into species groups (Appendix 1). We list the charac-
ters for each species group, but many characters are
homoplastic, and formal phylogenetic analysis is nec-
essary to establish the monophyly of these groups. The
original description of Strymon anthracaetus Salazar,
Vélez, & Johnson, 1997 was too poor to identify this
name or to determine whether it belongs to Strymon.
In August 2000 G. Lamas could not find the type at
the museum in Manizales, Colombia, where it was
supposed to be deposited. Consequently, we regard S.
anthracaetus as anomen dubium.
Names removed from Strymon. Many names
that were described in Strymon do not belong to Stry-
mon as we have characterized it. We confirm the pre-
vious transfer to other genera (Bridges 1988) of 23
VOLUME 55, NUMBER 3
names that were originally described in Strymon (Ap-
pendix 2).
We transfer the following six species, which were re-
cently described in Strymon, to other genera and give
brief reasons for the new generic placement. One is
transferred to “Thecla” because it belongs to a genus
that is yet undescribed.
1. Strymon angulus (Le Crom & Johnson, 1997) is
transferred to Thereus Hiibner, new combination.
Robbins (1991, 2000) characterized Thereus by a pair
of sclerotized invaginations on the membrane attached
to the ventro-lateral sides of the papillae anales, a pair
of ventro-lateral brush organs in addition to the pair of
dorsal ones, and sexual dimorphism in the antennal
club (>4 more nudum segments in the female than the
male). Although these characters are not mentioned in
the original description of S. angulus, the illustrated
holotype is a male of Thereus endera Hewitson, which
possesses the three synapomorphies of Thereus listed
above (Robbins in press).
2. Strymon daplissus Johnson & Salazar, 1993 is
transferred to Ministrymon Clench, new combina-
tion. The illustrated holotype of S. daplissus is a male
of Thecla clytie Edwards, which Clench (1961) placed
in Ministrymon.
3. and 4. Strymon carmencitae (Le Crom & Johnson,
1997) and Strymon cryptogramus (Johnson, Eisele &
MacPherson, 1992) are transferred to Nicolaea John-
son, new combinations. We characterize Nicolaea by
its male genitalia vinculum, which is strongly curved an-
teriorly in lateral aspect and which is flattened dorsally
in anterior aspect. These characters cannot be seen in
the original descriptions, but the illustrated holotypes of
Strymon carmencitae and Strymon cryptogramus are
specimens of Nicolaea fabulla (Hewitson, 1868) and N.
torris (Druce, 1907), respectively, which possess the
vinculum of Nicolaea as described (Robbins in press).
5. Strymon nivnix (Johnson, Eisele & MacPherson,
1990) is transferred to Calycopis Scudder, new com-
bination. The forewings of the holotype of S. nivnix
belong to a different genus than the hindwings (Rob-
bins in press), which appear to be the slightly aberrant
hindwings of Calcyopis cecrops F. We designate the
hindwings as the lectotype, which is placed in Calycopis
as characterized by Clench (1961) and Field (1967).
6. Strymon additionalis (Le Crom & Johnson, 1997)
is transferred to Thecla F., new combination. The
holotype is a male of “Thecla” emessa Hewitson, 1867,
which is characterized by the form of its scent patch on
the dorsal surface of the forewings, a white medial
stripe on the frons, and a penis tip with its ventral sur-
face flattened. Because the latter two characters were
not illustrated in the original description, our identifi-
cation is based upon the illustrated wing pattern and
scent patch.
ACKNOWLEDGMENTS
We dedicate this paper to the memories of J. B. Ziegler and H. K.
Clench, who together modemized the generic taxonomy of North
American hairstreaks. For the loan or gift of Strymon specimens over
the years, we are grateful to P. Ackery, C. Brévignon, C. Callaghan,
M. Casagrande, J.-Y. Gallard, J. Glassberg, G. Lamas, J. MacDonald,
L. Miller, Jacqueline Miller, James Miller, O. Mielke, J. Rawlins, B.
Sullivan, R. Vane-Wright, and J. Weintraub. We thank J. Glassberg
for allowing us to use his photographs of perching Strymon males.
We are grateful to V. Malikul for help with photography, S. Braden
with the SEM, and G. Venable with digitization and layout of the fig-
ures. For detailed comments on the manuscript, we thank J. Brown,
J. Burns, P. Gentili-Poole, J. Glassberg, J. Hall, D. Harvey, G.
Lamas, C. Penz, B. Sullivan, and the late B. Ziegler.
LITERATURE CITED
ALCOCK, J. 1983. Territoriality by hilltopping males of the great
purple hairstreak, Atlides ‘halesus (Lepidoptera, Lycaenidae):
convergent evolution with a pompilid wasp. Behav. Ecol. Socio-
biol. 13. 57-62.
ALCOCK, J. & K. M. O'NEILL. 1986. Density-dependent mating
tactics in the grey hairstreak, Strymon melinus (Lepidoptera:
Lycaenidae). Te Zool. Series A 209:105-113.
1987. Territory preferences and intensity of competition in
the grey hairstreak ‘Strymon melinus (Lepidoptera: Lycaenidae)
and the tarantula hawk wasp Hemipepsis ustulata (Hy-
menoptera, Pompilidae). Am. Midl. Nat. 118:128-138.
AuSsTIN, G. T. & K. JOHNSON. 1997. Theclinae of Rond6nia, Brazil:
Strymon Hiibner, with descriptions of new species (Lepi-
doptera: Lycaenidae). Insecta Mundi 11:201—235.
Barnes, W. & F. H. BENJAMIN. 1926. Notes on diumal Lepi-
doptera, with additions and corrections to the recent “List of
Diurnal Lepidoptera”. Bull. S. Calif. Acad. Sci. 25:88-98.
Barnes, W, & J. H. McDuNNoucH. 1917. Check list of the Lepi-
doptera of boreal America. Herald Press, Decatur, Illinois. 392 pp.
BEUTELSPACHER, C. R. 1972. Some observations on the Lepi-
doptera of bromeliads. J. Lepid. Soc. 26:133-137.
BripGEs, C. 1988. Catalogue of Lycaenidae & Riodinidae (Lepi-
doptera: Rhopalocera). Published by the author, Urbana, Illi-
nois. 788 pp.
CarTER, W. 1934. Notes on two pests of pineapple not known in
Hawaii. Proc. Haw. Entomol. Soc. 8:395-397.
CLEeNcH, H. K. 1961. Tribe Theclini, pp. 177-220. In P. R. Ehrlich
& A. H. Ehrlich, How to know the butterflies. Brown Co.,
Dubuque, Iowa.
. 1964. A synopsis of the West Indian Lycaenidae, with re-
marks on their zoogeography. J. Res. Lepid. 2:247-270.
CORDERO, C. R. & J. SOBERON. 1990. Non-resource based territo-
riality in males of the butterfly Xamia xami (Lepidoptera: Ly-
caenidae). J. Insect Behav. 3:719-732.
D’Araujo E Sitva, A. G., C. R. GoncALvEs, D. M. Gatvao, A. J. L.
GONGALVES, J. GOMES, M. DO NASCIMENTO SILVA & L. DE SI-
MONI. 1967-1968. Quarto catélogo dos insetos que vivem nas
plantas do Brasil. Ministerio da Agricultura, Rio de Janeiro, Part
I, Vol. 1, 422 pp., Vol. 2, 906 pp.. ‘Part IL, Vol. 1, 622 pp.. Vol. 2,
265 pp.
Dos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalo-
cera. Mem. Lepid. Soc., No. 1. 145 pp.
DucDaLe, J. S. 1974. Female genital configuration in the classifi-
cation of Lepidoptera. N. Ze: aland J. Zool. 1:127-146.
Euruicu, P. R. & P. H. RAVEN. 1965. Butterflies and plants: a study
in coevolution. Evolution 18:586—608.
96
Eurot, J. N. 1973. The higher classification of the Lycaenidae
(Lepidoptera): a tentative arrangement. Bull. Brit. Mus. (Nat.
Hist.) Entomol. 28:371—505.
FIELD, W. D. 1941. Notes on Erora laeta (Edwards) and Erora
quaderna (Hewitson) (Lepidoptera, Lycaenidae). Ann, Ento-
mol. Soc. Amer. 34:303-316.
. 1967. Preliminary revision of butterflies of the genus Ca-
lycopis Scudder (Lycaenidae: Theclinae). Proc. US Natl. Mus.
119:#3552, 48 pp., 126 figs.
FONSECA, J. P. 1934. Relagao das principais pragas observadas nos
anos de 1931, 1932 e 1933, nas plantas de maior cultivo no Es-
tado de S. Paulo. Arq. Inst. Biol. (S. Paulo) 5:263-189.
GUAGLIUMI, P. 1965. Contributo all conoscenza dell’entomofuana
nociva del Venezuela (coninuazione e fine). Riv. Agricol. Sub-
trop. Trop. 59:447-472.
. 1967. Insetti e arachnidi delle piante comuni del Vene-
zuela segnalati nel periodo 1938-1963. Relaz. Monogr. Agraria
Subtrop. Trop. (N.S.) 86:391 pp.
Harris, W. V. 1927. Ona lycaenid butterfly attacking pineapples in
Trinidad, B.W.I. Bull. Entomol. Res. 18:183-188.
HEMMING, F. 1967. The generic names of the butterflies and their
type-species (Lepidoptera: Rhopalocera). Bul. Brit. Mus. (Nat.
Hist.) Entomol., suppl. 9. 509 pp.
Honey, M. R. & M. J. ScoBLeE. 2001. Linanaeus’s butterflies (Lep-
idoptera: Papilionoidea and Hesperioidea). Zool. J. Linn. Soc.
132:277-399.
HUBNER, J. 1818. Zutriige zur Sammlung Exotischer Schmettlinge
[sic], bestehend in Bekundigung einzelner Fliegmuster neuer
oder rarer nichteuropiischer Gattungen. Jacob Hiibner, Augs-
burg 1:1-49.
. 1819. Verzeichniss bekannter Schmettlinge [sic]. Jacob
Hiibner, Augsburg. 2-8:17-128.
INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE.
1959. Opinion 541. Suppression under the plenary powers of
the generic names Chrysophanus Hiibner, 1818, and Bithys
Hiibner, 1818 (Class Insecta, Order Lepidoptera) (Opinion
supplementary to Opinion 165). Opin. Internat. Comm. Zool.
Nomen. 20:87—101.
JOHNSON, K., R. C. EISELE & B. N. MACPHERSON. 1990. The “hair-
streak butterflies” (Lycaenidae, Theclinae) of northwestern Ar-
gentina. II. Strymon, sensu stricto. Bull. Allyn Mus. 130:1-77.
Jounson, K. & K. R. KROENLEIN. 1993. Three remarkable new
species of Strymon Hiibner from Brazil and Ecuador (Lepi-
doptera, Lycaenidae, Theclinae). Rep. Mus. Nat. Hist., Univ.
Wisc. (Stevens Point). 35:1-8.
Jounson, K., L. D. MILLER & J. V. HERRERA. 1992. Eiseliana and
Heoda, high Andean and austral genera of neotropical Eu-
maeini (Lepidoptera: Lycaenidae). Acta Entomol. Chilena
17:107-146.
JoHNson, K. & J. A. SALAZAR. 1993. New species, statuses and
combinations in northern South American Strymon (Ly-
caenidae, Theclinae). Rep. Mus. Nat. Hist., Univ. Wisc.
(Stevens Point). 26:1-13.
Kors, A. B. 1951. A field guide to the butterflies of North America,
east of the Great Plains. Houghton Mifflin, Boston, 349 pp.
. 1970. Lepidoptera, pp. 115-130. In S. L. Tuxen (ed.), Tax-
onomist’s glossary of genitalia in insects. Munksgaard, Copen-
hagen.
Lamas, G., R. G. Rossins & W. D. FIELD. 1995. Bibliography of
butterflies. An annotated bibliography of the neotropical but-
terflies and skippers (Lepidoptera: Papilionoidea and Hesperi-
oidea). Atlas Neotr. Lepid. 124, 463 pp.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Ma, W. C. & L. M. SCHOONHOVEN. 1973. Tarsal contact chemosen-
sory hairs of the large white butterfly Pieris brassicae and their
possible role in oviposition behaviour. Entomol. Exp. & Appl.
16:343-357.
McDunnoucu, J. H. 1938. Check list of the Lepidoptera of
Canada and the United States of America Part 1. Macrolepi-
doptera. Mem. 5th Calif. Acad. Sci. 1:3-272 (1-3 [corrigenda]).
OpLeR, P. & V. MALIKUL. 1992. A field guide to easter butterflies.
Houghton Mifflin, Boston. 396 pp.
OPLER, P. & A. B. Wricut, 1999. A field guide to western butter-
flies. Houghton Mifflin, Boston. 540 pp.
Orero, L. S. & L. C. Marico. 1990. Butterflies: beauty and be-
havior of Brazilian species. Marigo Comunicagao Visual, Rio de
Janeiro. 128 pp.
POWELL, J. 1968. A study of area occupation and mating behavior
in Incisalia iroides (Lepidoptera: Lycaenidae). J. N. Y. Entomol.
Soc. 76:47-57.
RieEy, N. D. 1922. Notes on the generic names of Indian Thecli-
nae and Amblypodiinae (Lep. Rhop.). J. Bombay Natur. Hist.
Soc. 28:465-473.
ROBBINS, R. K. 1978. Behavioral ecology and evolution of hair-
streak butterflies (Lepidoptera: Lycaenidae). Ph.D. Disserta-
tion, Tufts University. 146 pp.
. 1980. The lycaenid “false head” hypothesis: historical re-
view and quantitative analysis. J. Lepid. Soc. 34:194-208.
. 1987. Evolution and identification of the New World hair-
streak butterflies (Lycaenidae: Eumaeini): Eliot's Trichonis Sec-
tion and Trichonis Hewitson. J. Lepid. Soc. 40:138-157.
. 1991. Evolution, comparative morphology, and identifica-
tion of the eumaeine butterfly genus Rekoa Kaye (Lycaenidae:
Theclinae). Smith. Contr. Zool. #498. 64 pp.
. 2000. The New World Hairstreak Genus Arawacus Kaye
(Lepidoptera: Lycaenidae: Theclinae: Eumaeini). Proc. Wash.
Entomol. Soc. 102:162-169.
. In Press. Eumaeini. In G. Lamas (ed.), Checklist: Part 4A.
Papilionoidea—Hesperioidea. In J. B. Heppner (ed.), Atlas of
Neotropical Lepidoptera. Scientific Publishers, Gainesville.
RosBINSs, R. K. & A.. AIELLO. 1982. Foodplant and oviposition
records for Panamanian Lycaenidae and Riodinidae. J. Lepid.
Soc. 36:65-75.
RosBINS, R. K. & S. S. NicoLay. 1999. Taxonomy and nomencla-
ture of Strymon istapa and S. columella (Lycaenidae: Thecli-
nae). J. Lepid. Soc. 52:318-327.
ScHwartz, A. & J. Y. MILLER. 1985. A new species of hairstreak
(Lycaenidae) from Hispaniola. Bull. Allyn Mus. 99:1-6.
SCUDDER, S. H. 1872. A systematic revision of some of the Ameri-
can butterflies, with brief notes on those known to occur in Es-
sex County, Massachusetts. Report, Peabody Acad. Sci.
4:94 83,
Tietz, H. M. 1972. An index to the described life histories, early
states and host of the Macrolepidoptera of the continental
United States and Canada. Vol. 1. Allyn Museum of Entomol-
ogy, Sarasota. 536 pp.
WILLIs, J. C. 1973. A dictionary of the flowering plants and ferns.
8th ed. revised by H. K. Airy Shaw. University Press, Cam-
bridge. 1245 pp.
ZIEGLER, J. B. 1960. Preliminary contribution to a redefinition of
the genera of North American hairstreaks (Lycaenidae) north of
Mexico. J. Lepid. Soc. 14:19-23.
ZIKAN, J. F. 1956. Beitriig zur Biologie von 12 Theclinen-arten.
Dusenia 7:139-148.
VOLUME 55, NUMBER 3
APPENDIX 1. Species groups, their characters, and
remarks. Character states in bold are especially useful in
delimiting that species group. Species synonymies follow
Robbins (in press). The subspecies concept was not used
in Robbins (in press), so we have designated geograph-
ically variable species as Synonyms and Subspecies.
STRYMON MELINUS GROUP. Characters: Male
lacking a scent patch on the dorsal surface of the FW,
no basal patch of white scales on the ventral surface of
the HW (except S. tyleri), penis tip down-turned with
a single slender cornutus, ductus bursae with simple
sclerotized loop, ductus seminalis arises from the un-
sclerotized posterior end of the corpus bursae, female
with Sth tergum unfurrowed and lacking imbedded
vestigial spiracles. Remarks: Some members, such as
S. melinus, S. tyleri, and S. rufofusca, can be exceed-
ingly common where they occur.
1. Strymon melinus (Hiibner, 1813)
SYNONYMS AND SUBSPECIES: Thecla hyper-
ici (Boisduval & Leconte, 1835), Thecla humuli
(Harris, 1841), Thecla pudica (H. Edwards,
1877), Strymon atrofasciata McDunnough, 1921,
Strymon setonia McDunnough, 1927, Strymon
meinersit Gunder, 1927, Thecla clarionensis
(Heid, 1933), Strymon youngi Field, 1936, Stry-
mon franki Field, 1938, Strymon caldasensis
Salazar, Vélez & K. Johnson, 1997
2. Strymon avalona (W.G. Wright, 1906)
3. Strymon tyleri (Dyar, 1913)
4. Strymon sabinus (C. Felder & R. Felder, 1865)
SYNONYM: Thecla promissa (Moschler, 1883)
5. Strymon rufofusca (Hewitson, 1877)
SYNONYMS AND SUBSPECIES: Thecla (Uran-
otes) valentina (Berg, 1896), Thecla lucaris (A.G.
Weeks, 1901), Thecla grisea (Dufrane, 1939),
Thecla nigriplaga (Dufrane, 1939), Strymon gua-
nensis Le Crom & K. Johnson, 1997
6. Strymon cyanofusca K. Johnson, Eisele &
MacPherson, 1990
STRYMON ALBATA GROUP. Characters: Male with a
scent patch on the dorsal surface of the FW, no basal
patch of white scales on the ventral surface of the HW
(except S. albata), penis tip down-turned with a single
slender cornutus, ductus bursae with simple sclerotized
loop, ductus seminalis arises from the unsclerotized
posterior end of the corpus bursae, female with 8th ter-
gum unfurrowed and lacking imbedded vestigial spira-
cles. Remarks: The only consistent difference with the
previous group is that the males have a scent patch.
7. Strymon albata (C. Felder & R. Felder, 1865)
SYNONYM: Thecla sedecia (Hewitson, 1874)
8. Strymon alea (Godman & Salvin, 1887)
SYNONYM: Callicista laceyi (Barnes & McDun-
nough, 1910)
9. Strymon bebrycia (Hewitson, 1868)
SYNONYMS: Thecla chonida (Hewitson, 1874),
Strymon buchholzi H.A. Freeman, 1950
STRYMON YOJOA GROUP. Characters: Male with a
scent patch on the dorsal surface of the FW (except S.
ohausi), no basal patch of white scales on the ventral
surface of the HW (except S. yojoa), penis tip down-
turned with a single slender cornutus that is barely
sclerotized, ductus bursae with a twist, but lacking
a sclerotized loop, ductus seminalis arises from the
unsclerotized posterior end of the corpus bursae, fe-
male with 8th tergum unfurrowed and lacking imbed-
ded vestigial spiracles. Remarks: Johnson et al. (1992)
noted that Thecla tegea Hewitson does not belong to
Strymon, but its male genitalia have anterior pointing
teeth on the valvae, and its male and female genitalia
are exceedingly similar to those of S. yojoa.
10. Strymon yojoa (Reakirt, 1867)
SYNONYMS AND SUBSPECIES: Thecla daraba
(Hewitson, 1867), Thecla beroea (Hewitson,
1868)
11. Strymon tegaea (Hewitson, 1868)
SYNONYM: Thecla seitzi (Spitz, 1931)
12. Strymon ohausi (Spitz, 1933)
STRYMON MULUCHA GROUP. Characters: Male
with a scent patch on the dorsal surface of the FW,
with basal patch of white scales on the ventral surface
of the HW, penis tip down-turned with a single slender
cornutus, ductus bursae with simple sclerotized loop
(except for some individuals of S. cestri), ductus semi-
nalis arises from the unsclerotized posterior end of the
corpus bursae, female with Sth tergum unfurrowed
and lacking imbedded vestigial spiracles. Remarks:
This species group is probably paraphyletic with re-
spect to the previous group.
13. Strymon mulucha (Hewitson, 1867)
SYNONYMS: Tmolus invisus (Butler & H. Druce,
1872), Strymon necjebus Le Crom & K. Johnson,
1997, Strymon necjabus Le Crom & K. Johnson
(missp.), 1997, Strymon novasignum Austin & K.
Johnson, 1997, Strymon clavus Austin & K. John-
son, 1997, Strymon implexus Austin & K. Johnson,
1997, Strymon inmirum Austin & K. Johnson,
1997, Strymon incanus Austin & K. Johnson, 1997
14. Strymon cestri (Reakirt, 1867)
SYNONYMS AND SUBSPECIES: Thecla cydia
(Hewitson, 1874), Thecla crossoea (Hewitson,
1874), Strymon chamiensis Salazar, Vélez & K.
98
Johnson, 1997, Strymon germana Austin & K.
Johnson, 1997
15. Strymon davara (Hewitson, 1868)
SYNONYMS: Thecla joannisi (Dufrane, 1939),
Thecla pallida (Dufrane, 1939)
16. Strymon crambusa (Hewitson, 1874)
17. Strymon astiocha (Prittwitz, 1865)
SYNONYMS: Thecla faunalia (Hewitson, 1868),
Thecla deborrei (Capronnier, 1874), Strymon ha-
los Austin & K. Johnson, 1997, Strymon consper-
gus Austin & K. Johnson, 1997
STRYMON MARTIALIS GROUP. Characters: Male
with or without a scent patch on the dorsal surface of
the FW, no basal patch of white scales on the ventral
surface of the, penis tip down-turned with a single
“wide” cornutus, ductus bursae with sclerotized loop
complex, especially in S. martalis (Fig. 23), ductus
seminalis arises from the posterior end of the
corpus bursae which is sclerotized dorsally, fe-
male with Sth tergum unfurrowed and lacking imbed-
ded vestigial spiracles. Remarks: A sister species rela-
tionship between the two included species is a new
hypothesis, but one that appears to be reasonably
strongly supported by the shape of the cornutus and
by the sclerotized patch on the corpus bursae. This
species group may be more closely related to the S.
melinus species group than its position here indicates.
18. Strymon martialis (Herrich-Schiffer, 1865)
19. Strymon christophei (W.P. Comstock & Hunting-
ton, 1943)
STRYMON ISTAPA GROUP. Characters: Male with a
scent patch on the dorsal surface of the FW (except for
S. oribata), no basal patch of white scales on the ven-
tral surface of the HW, penis tip down-turned with a
single slender cornutus, ductus bursae with sclerotized
loop simple except in S. acis, ductus seminalis arises
either from the unsclerotized posterior end of the cor-
pus bursae or from a point anterior of a sclerotized
patch on the dorsal surface of the corpus bursae, fe-
male with 8th tergum furrowed (except in S.
acis) and with imbedded presumed vestigial spir-
acles. Remarks: The inclusion of S. bazochii and S.
acis in this group are new hypotheses. The latter
species is included because it has imbedded presumed
vestigial spiracles and basal spots on the ventral surface
of the HW, which most of the species in this group
share. Its inclusion is highly tentative because its fe-
male 8th abdominal tergum lacks furrows and the
sclerotized loop of its ductus bursae is exceedingly
complex, unlike any other Strymon species.
20. Strymon istapa (Reakirt, 1867)
bo
bo
26.
Ne)
~l
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
SYNONYMS AND SUBSPECIES: Lycaena mod-
esta (Maynard, 1873), Callicista ocellifera (Grote,
1873), Thecla cybira (Hewitson, 1874), Thecla
arecibo (W.P. Comstock & Huntington, 1943),
Thecla clarionica (Vazquez, 1958), Thecla socor-
roica (Vazquez, 1958), Strymon clenchi Austin &
J.F. Emmel, 1998
. Strymon bazochii (Godart, 1824)
SYNONYMS: Hyreus thius (Geyer, 1832), Thecla
agra (Hewitson, 1868), hecla infrequens (A.G.
Weeks, 1901), Strymon gundlachianus M. Bates,
1935, Strymon diagonalis Austin & K. Johnson,
1997
. Strymon acis (Drury, 1773)
SYNONYMS AND SUBSPECIES: Papilio mars
(Fabricius, 1776), Thecla gossei (W.P. Comstock
& Huntington, 1943), Thecla bartrami (W.P.
Comstock & Huntington, 1943), Thecla casasi
(W.P. Comstock & Huntington, 1943), Thecla
petioni (W.P. Comstock & Huntington, 1943),
Strymon armouri Clench, 1943, Strymon leu-
costicha Clench, 1992
. Strymon columella (Fabricius, 1793)
SYNONYMS: Papilio dion (Schaller, 1788), Tmo-
lus erytalus (Butler, 1870), Thecla antigua (W.P.
Comstock & Huntington, 1943)
. Strymon limenia (Hewitson, 1868)
. Strymon toussainti (W.P. Comstock & Huntington,
1943)
SYNONYMS: Strymon andrewi K. Johnson & Ma-
tusik, 1988, Strymon rhaptos K. Johnson, Eisele
& MacPherson, 1990, Strymon amonensis D.S.
Smith, K. Johnson, J.Y. Miller & McKenzie, 1991
Strymon bubastus (Stoll, 1780)
SYNONYMS AND SUBSPECIES: Papilio
minereus (Fabricius, 1787), Thecla salona (He-
witson, 1868), Thecla sapota (Hewitson, 1877),
Thecla peruensis (Dufrane, 1939), Thecla ponce
(W.P. Comstock & Huntington, 1943), Strymon
vividus Le Crom & K. Johnson, 1997
. Strymon eurytulus (Hiibner, 1819)
SYNONYMS AND SUBSPECIES: Thecla ameri-
censis (Blanchard, 1852), Thecla argona (Hewit-
son, 1874), Thecla rana (Schaus, 1902, Thecla tu-
cumana (H.H. Druce, 1907), Thecla nigra (Lathy,
1926), Strymon peristictos K. Johnson, Eisele &
MacPherson, 1990, Heoda nivea (K. Johnson,
L.D. Miller & Herrera, 1992)
. Strymon flavaria (Ureta, 1956)
SYNONYM: Heoda erani (Benyamini & K. John-
son, 1996)
. Strymon ollantaitamba (K. Johnson, L.D. Miller &
Herrera, 1992)
VOLUME 55, NUMBER 3
30. Strymon colombiana (K. Johnson, L.D. Miller &
Herrera, 1992)
SYNONYM: Heoda lecromi (K. Johnson & Lugo,
1997)
31. Strymon bicolor (Philippi, 1859)
SYNONYMS AND SUBSPECIES: Thecla quadri-
maculata (Hewitson, 1874), Thecla heodes (H. H.
Druce, 1909), Thecla leptocosma (Hayward,
1949), Thecla tricolor (Ureta, 1949), Heoda ata-
cama (K. Johnson & L.D. Miller, 1992), Eiseliana
probabila (K. Johnson, L.D. Miller & Herrera,
1992), Heoda suprema (K. Johnson, L.D. Miller
& Herrera, 1992), Heoda shapiroi (K. Johnson,
L.D. Miller & Herrera, 1992)
32. Strymon wagenknechti (Ureta, 1947)
33. Strymon oribata (Weymer, 1890)
SYNONYMS AND SUBSPECIES: Thecla areni-
cola (Jérgensen, 1934), Thecla punona (Clench,
1944), Thecla rojasi (Ureta, 1956), Eiseliana
koehleri (Toledo, 1978), Eiseliana patagoniensis
(K. Johnson, L.D. Miller & Herrera, 1992)
STRYMON SYLEA GROUP. Characters: Male with a
scent patch on the dorsal surface of the FW, no basal
markings on the ventral surface of the HW, penis tip
up-turned with a single slender cornutus, ductus bur-
sae straight without a loop or twist, ductus seminalis
arises either from the unsclerotized posterior end of
the corpus bursae, female with 8th tergum unfur-
rowed and lacking imbedded vestigial spiracles. Re-
marks: The upturned penis tip and straight ductus
bursae, which is like most Eumaeini other than Stry-
mon, suggest that the single species in this group may
be sister to the remainder of the genus.
34. Strymon sylea (Hewitson, 1867)
STRYMON SERAPIO GROUP. Characters: Male with
a scent patch on the dorsal surface of the FW, no basal
patch of white scales on the ventral surface of the HW,
penis tip down-turned with paired cornuti, ductus
bursae with simple sclerotized loop, ductus seminalis
arises from the unsclerotized posterior end of the cor-
pus bursae, female with 8th tergum unfurrowed and
lacking imbedded vestigial spiracles. Remarks: All lar-
val foodplant records in this groups are Bromeliaceae,
as noted.
39. Strymon serapio (Godman & Salvin, 1887)
SYNONYMS AND SUBSPECIES: Thecla lemnos
(H.H. Druce, 1890), Thecla mesca (Dyar, 1914),
Thecla inconspicua (Lathy, 1930), Strymon gol-
bachi K. Johnson, Eisele & MacPherson, 1990,
Strymon trunctogen K. Johnson & Salazar, 1993,
Strymon altamiraensis K. Johnson & Kroenlein,
99
1993, Strymon henaoi Salazar, Vélez & K. Johnson,
1997, Strymon hurtadoi K. Johnson, 1997, Stry-
mon rosari Torres & K. Johnson, 1997, Strymon
originatus K. Johnson, Hernandez & Cock, 1997
36. Strymon glorissima K. Johnson & Salazar, 1993
SYNONYM: Strymon campbelli K. Johnson &
Salazar, 1993
. Strymon gabatha (Hewitson, 1870)
SYNONYMS: Thecla balius (Godman & Salvin,
1887), Strymon alexandra K. Johnson & Kroen-
lein, 1993, Strymon alicia Salazar, Vélez & K.
Johnson, 1997
38. Strymon monopeteinus Schwartz & J.Y. Miller, 1985
39. Strymon veterator (H.H. Druce, 1907)
SYNONYMS: Strymon lorrainea K. Johnson,
Eisele & MacPherson, 1990, Strymon coronos K.
Johnson, Eisele & MacPherson, 1990
40. Strymon oreala (Hewitson, 1868)
41. Strymon dindus (Fabricius, 1793)
42. Strymon ochraceus K. Johnson & Salazar, 1993
SYNONYM: Strymon baricharensis Le Crom & K.
Johnson, 1997
43. Strymon lucena (Hewitson, 1868)
SYNONYMS: Thecla cardus (Hewitson, 1874),
Thecla canitus (H.H. Druce, 1907), Strymon spe-
cialus K. Johnson, Eisele & MacPherson, 1997
44. Strymon legota (Hewitson, 1877)
45. Strymon azuba (Hewitson, 1874)
SYNONYMS: Strymon montevagus K. Johnson,
Hisele & MacPherson, 1990, Strymon rojos K.
Johnson & Kroenlein, 1993
46. Strymon eremica (Hayward, 1949)
SYNONYMS AND SUBSPECIES: Strymon lariy-
ojoa Kk. Johnson, Eisele & MacPherson, 1990,
Strymon barbara K. Johnson, Eisele & MacPher-
son, 1990, Strymon nicolayi K. Johnson, Eisele &
MacPherson, 1990
47. Strymon megarus ( Godart, 1824)
SYNONYMS AND _ SUBSPECIES: Tmolus
basilides (Geyer, 1837), Thecla basalides (W.F.
Kirby, 1871) (missp.), Thecla tigonia (Schaus,
1902), Strymon amphyporphyra K. Johnson,
Eisele & MacPherson, 1990, Strymon rotundum
Austin & K. Johnson, 1997, Strymon gallardi
Faynel & K. Johnson, 2000
STRYMON ZIBA GROUP. Characters: Male with a
scent patch on the dorsal surface of the FW, no basal
patch of white scales on the ventral surface of the HW,
penis tip up-turned with two large unpaired cornuti,
ductus bursae slightly twisted without a sclerotized
loop, ductus seminalis arises from the sclerotized pos-
terior end of the corpus bursae, female with Sth ter-
(ey)
~l
SE
omer aee
oe
eee
100
gum unfurrowed and lacking imbedded vestigial spir-
acles. Remarks: The single included species is wide-
spread and common. It is the only Strymon lacking the
clear-cut anterior pointing teeth on the valvae.
48. Strymon ziba (Hewitson, 1868)
SYNONYMS AND SUBSPECIES: Thecla thulia
(Hewitson, 1868), Thecla diaguita (Hayward,
1949), Strymon baptistorum K. Johnson, Eisele &
MacPherson, 1990, Strymon dondiego K. Johnson
& Adams, 1997, Strymon profusorubrus Le Crom
& K. Johnson, 1997, Strymon lecromi K. Johnson,
1997, Strymon spinatus Austin & K. Johnson, 1997,
Strymon latamaculus Austin & K. Johnson, 1997,
Strymon pallidulus Austin & K. Johnson, 1997, Stry-
mon tholus Austin & K. Johnson, 1997
APPENDIX 2. Alphabetical list of specific names that
were originally described in Strymon and that have
been transferred to other genera (Bridges 1988). For
those names that are junior synonyms, we note its se-
nior synonym.
1. Cyanophrys agricolor (Butler & Druce, 1872)
2. Satyrium aliparops (Michener & dos Passos, 1942),
a junior synonym of Satyrium liparops (Leconte)
3. Satyrium borealis (Lafontaine, 1969), a junior syn-
onym of Satyrium calanus (Hiibner)
4. Satyrium caryaevorus (McDunnough, 1942)
5. Satyrium chlorophora (Watson & Comstock, 1920),
a junior synonym of Satyrium saepium (Boisduval)
6. Lamprospilus coelicolor (Butler & Druce, 1872)
7. Satyrium coolinensis (Watson & Comstock, 1920),
a junior synonym of Satyrium acadica (Edwards)
8. Satyrium desertorum (Grinnell, 1917), a junior
synonym of Satyrium sylvinus (Boisduval)
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
9. Electrostrymon dowi (Clench, 1941), a junior syn-
onym of Electrostrymon angelia (Hewitson)
10. Satyrium fletcheri (Michener & dos Passos, 1942),
a junior synonym of Satyrium liparops (Leconte)
11. Satyrium godarti (Field, 1938), a junior synonym
of Satyrium calanus (Hiibner)
12. Calycopis gottschalki (Clark & Clark, 1938), a ju-
nior synonym of Calycopis cecrops (Fabricius)
13. Satyrium immaculosus (Comstock, 1913), a junior
synonym of Satyrium titus (Fabricius)
14. Satyrium kingi (Klots & Clench, 1952)
15. Satyrium montanensis (Watson & Comstock,
1920), a junior synonym of Satyrium acadica (Ed-
wards)
16. Satyrium muskoka (Watson & Comstock, 1920), a
junior synonym of Satyrium acadica (Edwards)
17. Satyrium okanagana (McDunnough, 1944), a ju-
nior synonym of Satyrium saepium (Boisduval)
18. Cyanophrys pastor (Butler & Druce, 1872), a ju-
nior synonym of Cyanophrys longula (Hewitson)
19. Satyrium polingi (Barnes & Benjamin, 1926)
20. Satyrium provo (Watson & Comstock, 1920), a ju-
nior synonym of Satyrium saepium (Boisduval)
21. Satyrium swetti (Watson & Comstock, 1920), a ju-
nior synonym of Satyrium acadica (Edwards)
22. Satyrium violae (Stallings & Turner, 1947), a junior
synonym of Satyrium favonius (Smith)
23. Satyrium watsoni (Barnes & Benjamin, 1926), a
junior synonym of Satyrium titus (Fabricius)
Journal of the Lepidopterists’ Society
59(3), 2001, 101-111
LEPIDOPTERISTS’ PERCEPTIONS OF A PROPOSED PERMITTING SYSTEM FOR
BUTTERFLY COLLECTING ON PUBLIC LANDS
KRISTINE C. MAZZEI
AND
ARTHUR M. SHAPIRO
Evolution and Ecology, University of California, Davis, California 95616, USA
ABSTRACT. There has been widespread and often rancorous discussion of the need for and desirability of regulation of insect collecting
on public lands in the United States. In order to gauge the potential success and utility of a model “No-Fee Permitting System” (NFPS) for but-
terfly collecting, an anonymous survey was sent to all of the members of the Lepidopterists’ Society in northern California, Oregon, and Wash-
ington in the spring of 1998. Over 52% of the surveys were returned. While many respondents feel they would use the system honestly, few of
the respondents felt other lepidopterists would do so. Correlation analysis between the ranked questions using the Gamma statistic shows that
those respondents who collect most often believe the NFPS would not be a useful tool for making management decisions. Likewise, the most
frequent collectors are least likely to utilize the NFPS and have a tendency to believe the NFPS would worsen the relationship between collec-
tors and government agencies. Respondents were invited to comment on the questions. The most widely discussed themes included: (1) 36%
of the respondents felt the permit would be a nuisance and too difficult to fill out prior to the collection activity; (2) 27% felt the permit was a
good idea and was worth trying; (3) 44% expressed a general mistrust of governmental regulation; and (4) 29% felt the permitting strategy is an
improper conservation focus. The combination of the quantitative and qualitative responses demonstrates that the NFPS would not be used
widely enough for it to be a worthwhile management tool. The relationship between collectors and federal agencies is laden with mutual mis-
trust, and the installation of a NFPS would only apparently amplify the tension between these two groups of people. We recommend that any
future regulation be developed in an atmosphere of consultation and open communication.
Additional key words: conservation, land management, The Lepidopterists’ Society, U.S. National Forests.
Garry Wills (1999) recently published A Necessary In 1965 Frederick H. Rindge, in his presidential ad-
Evil: A History of American Distrust of Government, a dress to the Twelfth Annual Meeting of the Pacific
book that shows that resistance to authority is truly “as Slope Section of the Lepidopterists’ Society, expressed
American as apple pie.” Such resistance is, moreover, the urgent need for butterfly collections as a tool for
closely tied to a characteristic animus against “cos- future conservation work (Rindge 1965). Today, most
mopolitanism” and “expertise” — a finding which lepidopterists agree that loss and modification of habi-
should not surprise readers of Richard Hofstadter’s tat have caused reductions in butterfly diversity and
classic Anti-Intellectualism in American Life (1963). abundance. Yet the role of butterfly collecting in this
These forces in the American polity have recently scenario is not well defined. Collecting may be viewed
come into play in an unlikely arena — government as either a source of distribution and abundance data
regulation of butterfly collecting on federal lands. critical for future work, or as a factor in the loss of di-
The publication of Stainton’s (1857) Manual of versity due to potential over-collection of sensitive
British Butterflies and Moths in the mid 19th century species.
may mark the beginning of butterfly collection as a Recently butterfly collection has been a focus of
hobby popular with common people as well as the aris- controversy among lepidopterists, academics, federal
tocracy. This manual was the first affordable, yet accu- and state agency employees, and conservationists. This
rate guide to butterflies in Great Britain (Kirby 1903). controversy has received a fair amount of publicity in
By the late 1800's, butterfly collecting was common in public media. In 1996 two articles describing federal
the United States, and several popular manuals were prosecution of lepidopterists involved in illegal collec-
in circulation (Edwards 1879, Maynard 1891, Scudder tion activities appeared in popular recreation maga-
1893, Holland 1898). The Lepidopterists’ Society was zines, Audubon (Williams 1996) and Outside (Alexan-
founded in 1947. According to the 1996 Statement on der 1996). Among lepidopterists, the controversy
Collecting Lepidoptera, the Society supports collec- boiled over in a heated exchange filling much of the
tion as “one of many legitimate activities enabling pro- April 1996 News of The Lepidopterists’ Society, play-
fessional and avocational lepidopterists to further the fully nicknamed “The Collecting Issue.” A variety of
scientifically sound and progressive study of Lepi- ideas and attitudes towards butterfly collecting were
doptera and education about Lepidoptera as well as addressed. Most hailed the activity as an honorable
encouraging interaction between professional and av- pastime providing much-needed distributional data
ocational lepidopterists” (Executive Committee of the (Kral 1996, Sun 1996), while Jeffrey Glassberg (1996)
Lepidopterists’ Society 1996). highlighted the importance of “non-consumptive”
butterfly enjoyment, such as photography and sight-
identification.
The issue of butterfly collection is especially perti-
nent for federal land managers. For example, in south-
western Oregon — an area known for its high butter-
fly diversity — butterfly enthusiasts may conduct the
majority of their activities on National Forest lands.
Currently, the United States Forest Service does not
require permits for the non-commercial collection of
insects on such lands, but retains the right to regulate
collecting administratively (Joslin 1998). On the Rogue
River National Forest in Oregon two areas have been
designated as “closed” to the collection of butterflies
since 1990 [Title 36 Coded Federal Regulation
261.53(a)], including Dutchman’s Peak and Observa-
tion Peak. This closure was based on anecdotal evi-
dence of decline of the Small Apollo, Parnassius phoe-
bus sternitzkyi. Forest Service biologists observed
heavy collecting of P. p. sternitzkyi in previous years
and had concerns about the extreme environmental
conditions of the mountain peaks inhabited by the
subspecies (B. Mumblo pers. com. ).
There has been no study specifically addressing the
effects of collection on population dynamics. In 1964
and 1965 heavy predation pressure was artificially ap-
plied to the Jasper Ridge Colony of Euphydryas editha
without significant decreases in population sizes in
1965 and 1966 (Ehrlich et al. 1975). Yet, in this ex-
ample of intended overcollection the authors concede
that they were unable to remove more than 5 to 25%
of the females from the population. Since the popula-
tion structure and viability of each species is likely to
vary, individual examples will not provide blanket an-
swers for butterflies as a whole. Due to the lack of
clear scientific guidance, conservationists and land
managers alike usually have taken a conservative
stance, thereby increasing the amount of regulation of
butterfly collection. For example, the National Park
Service has limited butterfly collection strictly to re-
search purposes (Code of Federal Regulations, Title
36, Part 2, Section 2.5). Some lepidopterists shared
with us their belief that information on species abun-
dance and distribution within state and national parks
has declined because amateur butterfly collectors can
no longer monitor the changes in these parks. It is thus
claimed on the one hand that over-regulation can stim-
ulate the withdrawal of potential data sources, while
under-regulation may allow the over-collection of sen-
sitive butterfly species.
In the spring of 1998, on our own initiative but with
the approval of biologists from the Rogue River Na-
tional Forest, we attempted to design a permitting sys-
tem that would be acceptable to lepidopterists and
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
useful to forest managers. In an age when conservation
is essential, it is crucial for managers to maintain the
most up-to-date information on at least the most sen-
sitive species within their domain. An effective per-
mitting system for butterfly collection might assist in
this daunting task by providing information on one
particularly charismatic group of organisms. But what
would such a system be like?
The development of this permitting system would
itself be an experiment. Instead of creating it by fiat
“from above,” we approached this policy issue in a
more inclusive manner. As described below, the pro-
posed permitting system was “floated” with lepi-
dopterists in the Pacific Northwest as a sort of trial bal-
loon to determine its reception and potential utility.
MATERIALS AND METHODS
Creation of the permitting system. The “No-Fee
Permitting System” (henceforth NFPS) was purpose-
fully designed as a “user-friendly” means for butterfly
collection within the bounds of forest service land. The
procedure for using the NFPS was described in gen-
eral terms in the belief that excess detail would merely
distract from the aims of the system. It would work as
follows: Upon arrival, a collector would voluntarily fill
out a two-copy form detailing the date, location, the
species desired and the quantity to be taken. One copy
of the form would automatically serve as a free permit
and would be retained by the collector. The second
copy would be deposited in a drop box on-site, and pro-
vide the agency with information about which locations
and species experience the most collection pressure.
Survey of the members of The Lepidopterists’
Society. All of the members of The Lepidopterists’
Society in northern California, Oregon, and Washing-
ton were surveyed. They were asked whether they
would use the NFPS and their opinion of it as a man-
agement and conservation tool. Northern California
was arbitrarily defined as any location north of Sacra-
mento, California. All 86 members of the society in the
defined region as of 1997 were mailed a packet con-
taining a cover letter, survey, and a pre-addressed,
stamped envelope in which to return their responses.
The cover letter was designed to establish a disinter-
ested position on the issue, detail the procedure for us-
ing the NFPS, and assure the respondent of anonymity
(Appendix 1). The survey was made up of eight ques-
tions and the majority of the answers were rankings
from one to five for particular, scaled responses (Ap-
pendix 2). The questionnaires were not coded in any
way for individual tracking and were separated from
their envelopes so that postmarks could not be used to
identify respondents. A few respondents elected to in-
VOLUME 55, NUMBER 3
wn
e
oO
me}
c
fo}
Qa
wn
oO
oc
ao)
(3)
Q
=
=>)
Zz
4 5
Never 2 A few times : > 20
each year times/year
Frequency of Collection
Fic. 1. Distribution of the survey responses to the first question,
“How often do you collect butterflies?”
clude signed comments, sometimes quite lengthy, but
none is identified in this paper.
Analysis of the survey response data. The re-
sponses to the quantitative rank questions were
graphed in order to detect trends in the replies. Since
the responses to the survey questions were ordered
categories it was necessary to use nonparametric sta-
tistics for the correlation analysis. The Gamma Statis-
tic (Siegel & Castellan 1988) was used to measure cor-
relations between the responses of the different
ranked questions. The Gamma statistic is most appro-
priate for calculating correlations with non-continuous
data and, similarly to other correlation statistics, it pro-
duces results ranging from —1 to 1. A Gamma statistic
close to | indicates a positive relationship between the
responses of two different survey questions, while
Gamma statistics approaching —1 indicate negative re-
lationships between the responses.
All of the comments from the qualitative questions
were compiled into individual documents and these
responses were read and evaluated separately. The re-
sponses to each question were reviewed several times
and “themes” that appeared in more than one answer
were labeled and tallied. Once this exercise was com-
pleted it was evident that there was a significant
amount of overlap in the responses to questions Al 7,
and 8. Thus, the responses to the three questions were
combined for the most frequent themes and the total
number of respondents expressing each particular
theme was recorded. Each survey reply was given an
identification number after being separated from its
envelope to avoid double-counting sentiments ex-
pressed twice by the same respondent.
RESULTS
Quantitative survey questions. The responses to
the survey were collected during the summer of 1998.
Overall, 45 surveys were received, yielding a return
rate of 52.3%. Responses to question | (Fig. 1), “How
103
Other
4%
Exchange
T%
Profit
0%
Enjoyment
22%
Research
22%
Personal
Collection
Museum 29%
Collection
16%
Fic. 2. The relative frequencies of the given responses to the
second survey question, “Why do you collect butterflies?”
often do you collect butterflies?” demonstrate the
presence of a wide variety of lepidopterists in our
population, from those who collect quite frequently to
others who do not collect at all. The second question,
on the reasons for butterfly collection (Fig. 2), clearly
points towards “personal collection” as the most widely
cited rationale. “Museum collection,” “enjoyment,”
and “research” all follow closely behind “personal col-
lection” as reasons for butterfly collecting. Not a single
respondent claimed to be collecting for “profit,” and
only a few respondents said they collect for “exchange”
purposes.
Question 3 asked respondents, “If you decided to
collect butterflies and you went to a location that im-
plemented a voluntary “No-Fee Permitting System” as
described in the letter, (a) would you use it; (b) would
you fill it out candidly and accurately; (c) do you think
other collectors would use it; (d) do you think other
collectors would fill it out candidly and accurately?”
While many respondents claimed they would use the
NFPS “every time” and would do so candidly and ac-
curately “every time,” they did not hold the same ex-
pectations of compliance for their fellow lepidopter-
ists. The most popular response for questions 3A and
3B is 5, signifying “every time” (Fig. 3). Notably, this
majority shifts to 3, signifying “sometimes,” for ques-
tions 3C and 3D (Fig. 4): in other words, many re-
spondents consider themselves better conservationists
or more ethical than lepidopterists in general.
When asked in question 5, “Do you think the NFPS
would be a useful tool for making management deci-
;
104
30 = —— —
Question 3A
B Question 3B
Number of Respondents
a
10 -
5
0 : St) IN =
1 2 3 4 5
Never : Sometimes Every time
Selected Frequency
Fic. 3. Distribution of the survey responses to questions 3A and
3B. These questions asked, “If you decided to collect butterflies and
you went to a facility or location that implemented a voluntary ‘No-
Fee Permitting System’ as described in the letter, (3A) would you
use it?” and (3B) “Would you fill it out CANDIDLY and ACCU-
RATELY?”
sions about butterfly collecting?” the responses were
extremely scattered. Almost an equal number of re-
spondents circled “yes” as “no,” and the majority of re-
spondents answered “maybe.” However, the responses
to question 6 were more revealing. Over 20 respon-
dents who answered the question, “How do you think
the NFPS would affect relations between collectors
and government agencies?” felt it would worsen the
relationship (either a 4 or 5), while only ten respon-
dents felt the NFPS would improve it (either a 1 or 2).
While the raw data are interesting to examine, the
correlations between responses to particular questions
reveal some of the most intriguing patterns in these
data (Table 1). It is clear that the respondents who
claim they would use the NFPS also express the likeli-
hood of using it candidly and accurately (p < 0.0001).
Likewise, those who felt other lepidopterists would
use the NFPS also tended to think they would use it
candidly and accurately (p = 0.0005). Since the re-
sponses to 3A and 3B are so tightly linked to each
other, the additional correlations are only tested using
comparisons with 3A with the understanding that a
correlation with 3A indicates a similar level of correla-
tion with 3B. This reasoning is also used for correla-
tions with 3C and 3D.
Several interesting trends emerge from the correla-
tion data. While not all of the trends produce results
with statistical significance, the correlations mentioned
here indicate potentially important relationships be-
tween survey responses. These trends include: (1) The
respondents who collect butterflies the most fre-
quently tend to be the ones who would not utilize the
NFPS (G = -0.35, p = 0.11) and believe others would
not use the permit either (G = —0.26, p = 0.21). (2)
The respondents who collect butterflies the least are
the ones who believe most strongly that the NFPS is a
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
30 - Se >
Question 3C
@ Question 3D
Np N
oS Ci
Number of Respondents
a
oO
|
|
|
|
|
1 2 3 4 5
Never 5 Sometimes Every time
Selected Frequency
Fic. 4. Distribution of the survey responses to questions 3C and
3D. These questions asked, “If other butterfly collectors went to a fa-
cility or location that implemented a voluntary ‘No-Fee Permitting
System’ as described in the letter, (3C) do you think they would use
it?” and (3D) “Do you think they would fill it out CANDIDLY and
ACCURATELY?”
useful tool for making management decisions about
butterfly collecting, while the most frequent collectors
tend to hold the opposite opinion (G = —0.46, p =
0.015). (3) The respondents who claim to be the most
likely to use the NFPS and believe others would do
the same are also the ones who feel the NFPS may be
a useful tool for making management decisions and
could potentially improve the relations between but-
terfly collectors and government agencies (all correla-
tions are significant at p = 0.05). (4) The collectors who
collect with the highest frequency have a slight ten-
dency to believe that the NFPS would worsen rela-
tions between butterfly collectors and government
agencies (G = 0.26, p = 0.18).
Finally (5), there is a tight relationship between the
respondents who feel the NFPS is a useful manage-
ment tool and those who feel it will improve relations
between the collectors and the agencies (p = 0.0054).
This also indicates a tight relationship in the other di-
rection; the respondents who feel the relationship will
be worsened also feel the NFPS is not a useful man-
agement tool.
Qualitative survey questions. Besides the ranked
survey responses, there were three questions that so-
licited comments from the respondents. Question 4
asked the respondent to “Explain your responses to
question #3.” Question 7 inquired, “What is your opin-
ion of the NFPS?” Lastly, question 8 asked the re-
spondent to write any additional comments they
wanted to discuss regarding the NFPS. The writer was
invited to attach additional sheets if necessary.
Since the qualitative questions were all fairly gen-
eral in what they were asking, there was a great deal of
overlap in the responses. Several themes continued to
emerge in the written response data, therefore it is fit-
ting to discuss the general trends detected in the re-
VOLUME 55, NUMBER 3
TaBLE l. Selected correlations between pairs of quantitative survey questions using the Gamma statistic (G
significant correlations are emphasized with the following notations:
Pairs of questions tested for correlation
ooo
represents p < 0.001; °° represents p < 0.01; and °
Gamma Statistic (G)
105
), which ranges from —1.0 to 1.0. Highly
represents p S 0.05.
p-value
3A: If you went to a location with the NFPS, would 3B: Would you fill it out candidly and 0.83 <().0001
you use it? 1 = never, 5 = every time accurately? 1 = never, 5 = ev ery time a
3C: Do you think other collectors would use the NFPS? 3D: Do you think other collectors would 0.85 0.0005
1 = never, 5 = every time fill it out candidly and accurately? eS.0
1 = never, 5 = every time
1; How often do you collect butterflies? 1 = never, 3A: If you went to a location with the NFPS, —0.35 0.11
5 = >20 times/yr would you use it? 1 = never, 5 = every time
3C: Do you think other collectors would use —0.26 0.21
the NFPS? | = never, 5 = every time
5: Is the NFPS a useful tool for making —0.46 0. Oi
management decisions about BC? 1 = No, 5 = Yes
6: How do you think the NFPS will affect
relations between BC's and Govt. Agencies?
1 = Strongly Improve, 5 = Strongly Worsen 0.26 0.18
5: Is the NFPS a useful tool for making management 3A: If you went to a location with the NFPS, 0.49 0.044
decisions about BC? 1 = No, 5 = Yes would you use it? 1 = never, 5 = every time =
3C: Do you think other collectors would use 0.56 0.027
the NFPS? | = never, 5 = every time 2
6: How do you think the NFPS will affect —0.61 0.0054
relations between BC's and Govt. Agencies? ch
1 = Strongly Improve, 5 = Strongly Worsen
6: How do you think the NFPS will affect relations 3A: If you went to a location with the NFPS, —0.59 0.013
between BC’s and Govt. Agencies? would you use it? 1 = never, 5 = every time 2
1 = Strongly Improve, 5 = Strongly Worsen
3C: Do you think other collectors would —0.58 0.026
use the NFPS? 1 = never, 5 = every time
sponses to all three questions (4, 7, and §) in order to
consolidate the findings of the study.
Thirty-six percent of the respondents expressed the
belief that the permit would be a nuisance and that it
would be too difficult or impractical to fill out ahead of
time. One respondent explains, “As you've described it,
most collectors have to indicate what species and how
many individuals they intend to take upon arrival .
MOST collecting does not happen that way — it is
rather a treasure hunt — you go out and see what you
can find.” Another says “quite often people go to a new
area, especially on vacation, without prior knowledge
of what can be found.” Finally, another comment sug-
gests a different approach to permits as a data source:
“When I go to collect, the reason is never to pick up a
certain number of a certain species, but rather to sam-
ple what is flying that day . . . the system you suggest is
useless for me unless I get to fill out the species and
count info after the fact — say at day end or even bet-
ter later by mail.”
Several replies included suggestions for alternate per-
mit systems. Two respondents expressed their belief
that the permit needs “teeth,” in other words, some reg-
ulations specifically geared towards the enforcement ae
the system. Four respondents suggested that the permit
have a third copy on which the actual species list can be
recorded and submitted at a later time.
On the other hand, several respondents (27%) be-
lieved the permit was a good idea and it was worth try-
ing to determine if lepidopterists would comply with
the new system. One person felt the NFPS “might help
gauge the pressure on a fragile population.” Although
12 respondents included comments leaning in this di-
rection, these responses tended to be brief and sup-
portive, yet not as emphatic in tone as the negatives.
One of the most consistent trends in the qualitative
data was a mistrust of government. Quotes from re-
spondents include:
“IT am very nervous about the NFPS being put
under the supervision of any state or federal
agency, (underline original)
106
“T will confess, my knee-jerk reaction anytime that
it is suggested that government become more
heavily involved in scientific endeavors, even if
they be hobbyist endeavors such as collecting
butterflies, that it is a bad idea.”
“I think the government screws up everything it
touches.”
“Bad idea to involve ‘the government’ in any more
of our lives than absolutely necessary.”
These types of sentiments were recorded in 20 of
the 45 returned surveys (44%).
Furthermore, many respondents (29%) felt that
butterfly collecting is an ineffective conservation focus:
“Collecting of several specimens of a population of
insects in the vast majority of cases has absolutely
no effect on the population whatsoever.”
“It is well known that the decline of species of any
kind has been due to two principle (sic) causes —
long term climate change and loss of habitat.”
“One issue is the immense hypocrisy perceived by
lepidopterists when the USFS bans collecting...
while permitting hunting, fishing, logging, bug
zappers, and the spraying of vast acreages of forests
with Baccilus thuringiensis, killing millions of leps.”
There were several reasons advanced as to why lep-
idopterists would be unlikely to comply with this sys-
tem. This opinion was expressed in 18/45 surveys
(40%):
“Some persons might feel guilty that they have
overcollected and not report all of their catches.”
“Anyone pursuing larger numbers of specimens for
whatever reason would be least likely to truthfully
report their activities.”
“There are some people in it for money, these
people seemingly have no thought about habitat,
populations, future of a species or subspecies, just
dollars.”
Clearly, many lepidopterists feel that the collectors
who take the highest numbers would be the least likely
to use the NFPS, a trend that aligns with the correla-
tion results from the quantitative data.
Several people worried that the data collected from
the NFPS would not be of a high quality because so
many different people with varying levels of skill in
species identification would submit it. In addition,
there seems to be a fear that agency personnel would
interpret the data inappropriately. Concerns over data
quality and interpretation were cited in six returned
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
surveys (13%). Some of these respondents felt that a
biologist working for the Forest Service may have little
or no understanding of what factors have the most in-
fluence on butterfly populations. For instance, de-
clines due to inclement weather may be wrongly asso-
ciated with a modest level of collection, especially by
agency personnel unfamiliar with insect (as opposed to
tree or vertebrate) population dynamics.
Six respondents (13%) expressed the fear that the
NFPS would not remain “No-Fee” after a few years.
Three lepidopterists pointed out that they believe
there are too few collectors to justify an effort. One re-
spondent explained, “there are nowhere near as many
collectors as the public is led to believe. In nearly 70
years of collecting, I rarely see another collector.” Fi-
nally, six respondents (13%) expressed the belief that
lepidopterists have an innate conservation ethic or
highlighted the importance of butterfly collection for
conservation purposes. They explain that amateur but-
terfly collectors are often the main group of lepi-
dopterists that provide accurate distribution data for
species occurrences on an annual basis. Further, many
of the dot-distribution maps widely used in Oregon
and Washington were based on specimens from pri-
vate collections (Dornfeld 1980, Hinchcliff 1996).
With only a few academic and professional research
lepidopterists in the region, it is imperative to augment
their data with the work of amateur collectors. Many
point out that collectors out on exploratory jaunts are
often the ones who discover new species and popula-
tions in an area. Regulation of this activity may have
negative impacts on the willingness of these people to
share their discoveries.
DISCUSSION
Analysis of the potential success and utility of
the “No-Fee Permitting System.” Based upon the
responses to the first few questions on the survey, it is
clear that this group of people was an appropriate
population to be surveyed. Most of the 45 respondents
collect several times per year with only three who
claimed to never collect at all (Fig. 1). Collectors usu-
ally go out into the field with the intention of taking
specimens, with the most widely cited rationale being
“personal collection” (Fig. 2). While no one admitted
to collecting for profit, 8/45 of the respondents (18%)
collect with the intention of exchanging specimens.
Collection for the purpose of “exchange” may imply
the taking of more rarer or harder-to-catch species in
order to increase their exchange value.
While approximately half of the respondents said
they would utilize the NFPS every time they went to a
participating location, an even greater proportion of
VOLUME 55, NUMBER 3
respondents predicted a lower level of participation
for other lepidopterists. Sixty percent of the respon-
dents felt that other collectors would only use the
NFPS “sometimes” at participating locations. Only
6/45 lepidopterists (13%) say they would “never” use
the proposed system. Since some of these are the most
frequent collectors, as indicated by the slight correla-
tion between question | and question 3A (Table 1),
this is a significantly negative response.
From the response data it is apparent that the reac-
tion to the NFPS is not positive enough to justify initi-
ating this sort of conservation strategy: it would, in
fact, be counterproductive. Furthermore, from the
comments and quantitative data it is apparent that the
relationship between lepidopterists and government
agencies is seriously in need of repair. While both
groups would benefit from improved communication
and cooperation, neither thoroughly understands the
predicaments and concerns of the other. Yet, without
cooperation, the future of butterfly conservation is
needlessly compromised.
The approach of this survey was to involve the per-
sons most affected by the model system in the deci-
sion-making process. We feel this is one of the first
steps to improving the relationship between the two
groups. It is critical that the U.S. Forest Service listen
to the concerns of the respondents and make its policy
decisions with these suggestions in mind: that is, only
extremely compelling reasons would justify failing to
heed the advice of the concerned public. In the Par-
nassius phoebus sternitzkyi case there are no such
compelling reasons.
It is clear from the ranked data that the lepidopter-
ists who collect most frequently have a tendency to be
the ones who do not consider the NFPS to be a useful
tool for making management decisions; they do not
think it will improve the relations between collectors
and government agencies, and they are the least likely
to comply with the system (Table 1). When these re-
sults are combined with the comments regarding the
success and utility of the NFPS, it becomes clear that
the system will be ignored and disliked by many, per-
haps most, frequent butterfly collectors. If imple-
mented with only weak support from lepidopterists
the NFPS would not only incite resentment, it would
also be nearly useless as a source of reliable data, mak-
ing it an essentially useless endeavor. In an era where
the sentiment towards government agencies is laden
with mistrust, agencies such as the US Forest Service
should strive to open channels of communication,
rather than close them by taking what would be per-
ceived as arbitrary bureaucratic action.
Suggestions for policy improvement and fu-
107
ture use. The NFPS was intended to improve the
amount of data available to land managers. Even if the
NFPS as we have described it would not be acceptable
to the collecting public, perhaps it would be helpful
for individual forests to set up voluntary data submis-
sion programs — carefully avoiding the word “permit”
or other threatening buzzwords. Titen lepidopterists
might feel they are doing their part for conservation
without the sense of unjustified regulation. This could
be a primary step toward improving the relationship
between collectors and government agencies. If legiti-
mate, well-documented conservation concerns ulti-
mately dictate some form of permitting system, it
would be much more likely to succeed if lepidopterists
understand its rationale and support its goals
ACKNOWLEDGMENTS
This study would not have been possible without the survey re-
spondents. Thank you for taking the time to think about our ques-
tions, write your comments, and mail the replies. Many people added
creative ideas to this project. We would like to thank Rich Van
Buskirk, Mikaela Huntzinger, Jim Fordyce, Nicole Jurjavcic, Tag Eng-
strom and Michelle Gadd for thinking about and discussing the
NFPS. We also appreciate the fe tbHBONS of two anonymous re-
viewers. Sy Schwartz facilitated an environmental policy seminar
that helped to shape the direction of the data interpretation. Barbara
Mumblo provided information on the recent history of butterfly col-
lection closures on the Applegate Ranger District of the Rogue
River National Forest. We gratefully acknowledge Joel E. Pagel.
who coordinated the funding of this research through the Rogue
River National Forest, Challenge Cost-Share Agreement.
LITERATURE CITED
ALEXANDER, C. 1996. Crimes of passion. Outside. January 1996:
29-32.
DornFELD, E. J. 1980. The butterflies of Oregon. Timber Press,
Forest Grove, Oregon. 276
Epwarps, W. H. 1879. The butterflies of North America: with col-
ored drawings and descriptions. Houghton, Mifflin and Co.,
Boston.
EHRLICH, P. R., R. R. WHITE, M. C. SINGER, S. W. MCKECHNIE & L.
E. GILBERT. 1975. Checkerspot butterflies: a historical per-
spective. Science 188:221—228.
EXECUTIVE COUNCIL OF THE LEPIDOPTERISTS’ SOCIETY. 1996. The
Lepidopterists’ Society Statement on Collecting Lepidoptera,
adopted by the Executive Council: 13 June 1996, Houston,
Texas. News of the Lepid. Soc. 38(4):108-110.
GLASSBERG, J. 1996. More on the Acom article. News of the Lepid.
Soc. 38(2):48—49.
HINCHLIFF, J. 1996. The distribution of the butterflies of Washing-
ton. The Evergreen Aurelians. Corvallis, Oregon.
HorstTaDTER, R. 1963. Anti-Intellectualism in American life.
Knopf, New York. 434 pp.
HOLLAND, W. J. 1898. The butterfly book: a popular guide to a
knowledge of the butterflies of North America. Doubleday &
McClure Co., New York. 382 pp.
Jostin, R. C. 1998. Noncommercial collection of insects on Na-
tional Forest system lands. News of the Lepid. Soc. 40(4):74.
Kirpy, W. F. 1903. The butterflies and moths of Europe. Caswell
and Company, Limited, London. 432 pp.
KRaL, T. W. 1996. Getting the facts straight: Thomas Kral speaks
out. News of the Lepid. Soc. 38(2):41-44.
108 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
MAYNARD, C. J. 1891. A manual of North American butterflies. De STAINTON, H. T. 1857. A manual of British butterflies and moths. J.
Wolfe, Fiske, and Co., Boston. 226 pp. Van Voorst, London.
RINDGE, F. H. 1965. The importance of collecting — Now. J. Lepid. SuN, S. K. 1996. On the federal regulation of insect collecting.
Soc. 19:193-195. News of the Lepid. Soc. 38(2):51-53.
SCUDDER, S. H. 1893. Brief guide to the commoner butterflies of WILLIAMS, T. 1996. The great butterfly bust. Audubon. March-
the northern United States and Canada; being an introduction April 98(2):30-37.
to a knowledge of their life-histories. H. Holt and Company, WILLS, G. 1999. A necessary evil: a history of American distrust of
New York. 206 pp: government. Simon and Schuster, New York. 365 pp:
SIEGEL, S. & N.J. CASTELLAN, JR. 1988. Nonparametric statistics for
the behavioral sciences. 2nd ed. McGraw-Hill, Boston. 399 pp.
APPENDIX 1 — Letter to Lepidopterists.
Evolution and Ecology
University of California, Davis
Davis, CA 95616
email: kemazzei@ucdavis.edu
June 17, 1998
Dear Member of The Lepidopterists’ Society,
I am a graduate student at the University of California, Davis, and I am interested in the
conservation biology of butterflies. Regulation of collecting has become a very controversial subject in
part because of poor communication about the justification and goals of the regulation. As part of my
Master’s thesis research I would like to determine how a new butterfly collection monitoring program
would be received by amateur, professional, and academic lepidopterists. Since the proposed system has
not been used before, the results of the enclosed survey will be of key importance when determining
whether such a program would be successful and worthwhile. Please read on!
As you may know, Apollo butterflies (genus Parnassius) are often under heavy collection
pressure. Intense butterfly collecting in a particular region has been claimed to contribute to irreversible
species decline. Both the collection and sale of Apollo butterfly species have been forbidden in Europe
and similar protective actions could occur in the United States. Lepidopterists are commonly aware of
“source” locations for particular species. The Siskiyou Mountains are the only habitat for a showy Apollo
subspecies, Parnassius phoebus sternitzkyi. Most of its habitats are on public lands.
At this time there are no hard data on how much collecting of Parnassius phoebus sternitzkyi is
occurring, or whether it is harming the populations. Because the subspecies has such a small geographic
range and a low reproductive rate, it may be proposed for protection of some kind. Such proposals should
not be made on pure instinct, however—protection might be urgently needed, but it also might be totally
unnecessary!
One way to monitor the amount of collecting on public lands would be the installation of a “No-
Fee Permitting System.” It would work like this: (1) Upon arrival, a collector would voluntarily fill out a
two-copy form detailing the date, location, the species desired and the quantity to be taken; (2) One copy
of the form would automatically serve as a free permit. The second copy, which would be deposited in a
drop box on-site, would provide the agency with information about which locations and species
experience the most collection pressure.
Please take a few minutes and complete the enclosed survey regarding the “No-Fee Permitting
System.” This survey is being sent to Lepidopterists’ Society members in Oregon, Washington, and
Northern California. The completed survey should be returned in the stamped envelope provided. |
would like to thank you for your honest participation in this important study. Let me assure you that there
are no identifying marks on the survey and that all replies will be kept absolutely anonymous. The
aggregate response may be published and/or provided to local, state, and federal agencies, and non-
governmental conservation organizations. It may shape the future of Parnassius phoebus sternitzkyi—
and of collecting on public lands. Thanks again!
Sincerely,
Kristine C. Mazzei
VOLUME 55, NUMBER 3 109
APPENDIX 2 — Survey on proposed “No-Fee Collection Permit System.”
Return to (stamped envelope has been provided):
Kristine C. Mazzei
Evolution and Ecology
University of California, Davis
Davis, CA 95616
email: kemazzei@ucdavis.edu
1998 Survey on Proposed “No-Fee Collection Permit System”
Please honestly respond to the following questions and feel free to explain any answers in the
“ADDITIONAL COMMMENTS?” section at the end of the survey. Thank you for your
participation. Please omit any information that might identify you or your location.
(1) How often do you collect butterflies (circle the appropriate number)?
Never A few times More than 20
each year times per year
1 2 3 4 5
(2) If you answered between 2 and 5 on question #1, please circle all of the reasons why you collect
butterflies:
Enjoyment Personal Museum Research Profit
Collection Collection
Exchange Other Reason:
(3) If you decided to collect butterflies and you went to a facility or location that implemented a
voluntary ““No-Fee Permitting System” as described in the letter,
(A) would you use it?
Never Sometimes Every Time
1 2 3 4 >
(B) would you fill it out CANDIDLY and ACCURATELY?
Never Sometimes Every Time
1 2 3 4 5
(continued on back)
110 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
(C) Do you think other collectors would use it?
Never Sometimes Every Time
1 Z 3 4 5
(D) Do you think other collectors would fill it out CANDIDLY and ACCURATELY?
Never Sometimes Every Time
1 2 3) 4 5
(4) Explain your responses to Question #3:
(5) Do you think the “No-Fee Permitting System” would be a useful tool for making management
decisions about butterfly collecting?
No Maybe Yes
1 2 3) 4 5
(6) How do you think the “No-Fee Permitting System” would affect relations between collectors and
government agencies?
Strongly No Effect Strongly
Improve Worsen
1 2 3 4 5
(7) What is your opinion of the “No-Fee Permitting System?”
(8) ADDITIONAL COMMENTS (please write as much as you like, use additional sheets if
necessary—but remember not to identify yourself).
‘Chank ou!
Journal of the Lepidopterists’ Society
55(3), 2001, 111-118
LIFE HISTORY AND LABORATORY HOST RANGE TESTS OF PARAPOYNX SEMINEALIS (WALKER)
(CRAMBIDAE: NYMPHULINAE) IN FLORIDA, U.S.A.
Gary R. BUCKINGHAM
Aquatic Plant Control Research, USDA-ARS, c/o Florida Biological Control Laboratory, P.O. Box 147100,
Gainesville, Florida 32614-7100, USA
AND
CHRISTINE A. BENNETT
Department of Entomology and Nematology, Institute of Food and Agricultural Sciences, University of Florida,
c/o Florida Biological Control Laboratory, P.O. Box 147100, Gainesville, Florida 32614-7100, USA
ABSTRACT. The native aquatic moth, Parapoynx seminealis (Walker), attacks big floating-heart, Nymphoides aquatica (S. G. Gmel.)
Kuntze, in Florida. Eggs are laid on the lower surface of floating leaves. Neonates bore into the relatively thick leaf or drop off the leaf on a
silken thread to feed on submersed leaves. Later instars build cases mostly by excising leaf pieces and attaching them to leaves. They feed from
the case or out of the case on the margin of the leaf and on the upper surface. Cocoons are made in the cases. There were three, or possibly four,
peak adult emergence periods in north-central Florida: June, August, mid-October, and possibly late March and April. Although the larvae are
specialists on floating-hearts, they fed and developed for relatively long periods on some non-host species in laboratory host range tests. The two
species most acceptable were Egeria densa Planchon and Hydrilla verticillata (L.fil.) Royle, both immigrant species in the non-related mono-
cot family Hydrocharitaceae.
Additional key words: _floating-heart, biocontrol, Nymphoides, Hydrilla, Egeria.
Aquatic moths in the genus Parapoynx (Crambidae: Big floating-heart ranges from New Jersey to
Nymphulinae) are of interest both to aquatic biologists Florida and west to Texas (Godfrey & Wooten 1981).
and to researchers on the biological control of aquatic It has heart-shaped leaves, 5-15 cm long, that are
weeds. The caterpillars feed on aquatic macrophytes green and smooth on the upper surface but purplish
and cover themselves with cases, portable or station- and rough or pitted on the lower surface due to an ir-
ary, which are made with the leaves of the host plant. regular layer of aerenchyma cells. The leaves are quite
They are the only aquatic caterpillars that have thick compared with leaves of most other native float-
branched gills on all body segments except the protho- ing species. Mature leaves were at least 1.5-3.0 mm
ax (Habeck 1974). Six species are native to North thick with about 50-66% of the thickness composed of
America. All have been reported from Florida (Mon- spongy aerenchyma cells. Several leaf petioles arise
roe 1972) although records for two species are suspect from the tip of a stem near the surface of the water.
[P. badiusalis (Walker) and P. curviferalis (Walker)]. A The white flowers arise just below the petioles and of-
seventh species, P. diminutalis Snellen, is an immi- ten a cluster of short, stout, fleshy roots forms among
grant in Florida (Buckingham & Bennett 1996). No or below the flower stalks (Godfrey & Wooten 1981).
host plants have been recorded for P. curviferalis, but These roots are colloquially called “bananas” and are
the other species, except the subject of this study, P. sold in the aquarium trade. “Bananas” excise and drop
seminealis (Walker), have hosts in at least two genera to the hydrosoil where they produce very thin sub-
and three have hosts in multiple families (Habeck mersed leaves and later new plants. Submersed leaves
1974). are also produced by rooted plants.
Parapoynx seminealis reportedly develops only on Several species of Parapoynx and other Nymphuli-
floating-heart, Nymphoides, and specifically on big nae have been of interest for the biological control of
floating-heart, N. aquatica (S. G. Gmel.) Kuntze submersed aquatic weeds (Buckingham 1994), but
(Menyanthaceae). A second native floating-heart, N. none has been purposefully released in any country
cordata (EIl.) Fern., occurs within the range of the (Julien & Griffiths 1998). We became interested in P.
moth. It has not been reported as a host, possibly be- seminealis while awaiting the results of foreign surveys
cause of lack of collecting or because often only the for natural enemies of submersed weeds. We planned
host genus is recorded. Herlong (1979) mentioned to conduct host range tests in quarantine with the po-
that a P. seminealis larva “on occasion was found on a tential control agents to determine their safety for re-
nearby leaf of Nymphaea odorata” Aiton (Nym- lease in the United States. Because of its reported host
phaeaceae), but he did not indicate whether the larvae specificity, the native P. seminealis appeared to be an
were actually feeding and developing. excellent test subject for developing techniques to rear
and study aquatic caterpillars and for comparing the
laboratory physiological host range with a known eco-
logical (or realized) host range. We report below our
studies on the life cycle and the host range of this in-
teresting aquatic moth.
MATERIALS AND METHODS
Insects and plants. Insects and host plant mate-
rial, big floating-heart, were collected by boat from
Santa Fe Lake, Alachua County, Florida, near Mel-
rose, one to three times a month from June 1980 to
June 1981 except during December, February, April,
and May. Damaged leaves were collected for observa-
tion and for recovery of immature stages. Undamaged
leaves were collected for a food source. Anecdotal ob-
servations were made of leaf abundance and damage
at the various sites.
Rearing. Undamaged leaves were held in outdoor
pools or in a refrigerated temperature cabinet. Rearing
was conducted in floating screen cages in the outdoor
pools and in plastic pans in the laboratory (22—24°C).
The large size of most floating leaves, 10-15 cm wide,
precluded rearing in small cages. New leaves were
added often because the leaves broke down, possibly
because of disease or because they were excised from
the roots. Efforts to transplant plants from the lake
were not successful, although some new plants were
grown from “bananas.” New adults were collected
from the cages, paired in the laboratory for mating,
and then placed into new cages. Usually eggs or
neonates were placed back into the cages after they
were counted, but some were removed for studies.
Initially, leaves were dipped in 0.01N potassium per-
manganate for 45 seconds to one minute to surface
sterilize them and then were washed with water. How-
ever, even treated leaves continued to break down
within a couple of days and this treatment was discon-
tinued early in the study.
Life cycle studies. Insects were held for these
studies (1) in the laboratory with natural lighting from
a window and with fluorescent lighting during work
hours (ca. 0700-1750 h), (2) in the quarantine labora-
tory with only fluorescent lighting during work hours,
and (3) in a quarantine greenhouse with natural light-
ing supplemented with fluorescent lighting (16 h pho-
tophase). Laboratory temperatures usually ranged
from 22-24°C and greenhouse temperatures from
26-32°C. Larval and pupal development was deter-
mined in a temperature cabinet at 27°C and 16 h pho-
tophase. Head capsules of living larvae were measured
at 12x and of preserved larvae at 50x with an ocular
micrometer in a dissecting microscope. All measure-
ments are reported as mean + standard error (number,
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
range). Specimens have been deposited in the Florida
State Collection of Arthropods (FSCA), Gainesville,
Florida, and in the U.S. National Museum of Natural
History (USNM), Washington, D.C.
No-choice host range tests. Test plants were field
collected and were held in pools for a few days up to
several weeks until used or in a refrigerated tempera-
ture cabinet for a few days. Tests were conducted in
the laboratory at 22-24°C with eggs containing active
larvae just prior to eclosion and with neonates. The
cages for these tests were 30 ml plastic cups with their
bottoms replaced by nylon organdy and capped with
either organdy or plastic lids. The cups sat on coarse
sand in a shallow pan filled partially with water aerated
by an aquarium pump. The water was aerated because
neonates had died within five days on floating-heart in
a preliminary test with closed cups and no aeration.
Pieces of test plants were cut to fit into the 30 ml cups,
which were 30 mm wide at the bottom and 40 mm at
the top and 45 mm deep. Stem tips or sections cut
from broad leaves were used depending on the growth
form of the plant. Each cup received one egg or
neonate. There were two no-choice tests, each with six
treatments (Table 1). Each test had two control treat-
ments: one without plant material and one with float-
ing-heart. The other treatments were common aquatic
plant species. Both tests were initiated with 20 repli-
cates in each treatment except the cage without plant
material in test B, which had ten replicates. Replicate
numbers (n) less than 20 in Table 1 are due to losses
during handling. Test A was terminated when all larvae
died. Test B was terminated at 22 days because float-
ing-heart leaves were lacking. The cups were exam-
ined for larval survival every three to seven days in test
A and four to seven days in test B. Plant material was
changed as needed. Presence of feeding was recorded
at each examination, but the amount was not esti-
mated.
No-choice tests were also conducted with medium-
sized larvae, 1 cm or greater in length, reared in the
laboratory on floating-heart (test C — larval age was 63
days after oviposition, test D — larval age was 49 days
after oviposition). Larvae were removed from their
cases and placed individually on test plants in 177 ml
Styrofoam drinking cups in the laboratory at 22-24°C.
Plants were cut to fit into the cups. Pieces of elongate
plants, ca. 9-18 cm long, and sections of large leaves,
ca. 5-8 cm diameter, were used. Initially, there were
eight replicates per treatment in test C and three
replicates in test D, except in the treatment without
plant material, which had five. One replicate in test C
was lost in handling, and after six days Vallisneria
americana Michaux (Hydrocharitaceae) was added to
VOLUME 55, NUMBER 3
TABLE |.
Plant species!
Cage without plant material
No-choice feeding tests with Parapoynx seminealis neonates.
Plant family
Egeria densa Planchon Hydrocharitaceae
Hydrilla verticillata (L. fil) Royle Hydrocharitaceae
Nymphoides aquatica (J. G. Gmel.) Kuntze Menyanthaceae
Sagittaria subulata (L.) Buch. Alismataceae
Vallisneria americana Michaux. Hydrocharitaceae
Cage without plant material =
Limnobium spongia (Bosc.) Steud. Hydrocharitaceae
Najas guadalupensis (Sprengel) Magnus Najadaceae
Nuphar luteum (L.) Sibth. & Smith Nymphaeaceae
Nymphoides aquatica Menyanthaceae
Potamogeton illinoensis Morong
Potamogetonaceae
Test Longevity (days) No. larvae*
symbol? mean SE range n 1 2-3. >4
A (3) 0.3 7-12 18
A 61.2 9) 7-89 15 4 3 10
A 24.3 6.4 7-102 = 20 ] 2 6
A 65.5 2, 7-89 19 0 4 16
A 8.2 0.5 7-12 17 3 0 0
A 9.5 0.8 7-12 10 3 0 0)
B 6.8 0.2 5-7 10
B 14.4 1.0 5-22 20 11 4 0
B 11.9 1.0 5-22 19 13 2 0
B 10.4 0.7 5-18 19 2 ] 0
B 19.7 BD 5-22 17 2 4 12
B 11.5 0.8 5-18 20 10 ] 0
Is
No. observations
with damage®
'N. aquatica, big floating-heart, is the field host plant.
> Cages were 30 call plastic cups with organdy bottoms sitting in aerated water in the laboratory at 22—24°C. There were 20 larvae per treat-
ment, Test A was conducted until larvae died. Test B was conducted for 22 days until the control plant, big floating-heart, was no longer avail-
able. Only three of 78 larvae were alive in test B at 22 days on test plants (L. spongia and N. guadalupensis) versus 12 of 20 on big floating-heart.
Numbers (n) less than 20 are due to handling losses, except the “cage without plant material” treatment in test B, which was started with only
10 larvae.
5 Cages were observed for feeding damage every three to seven days in test A, first observation at seven days, and four to seven days in test B,
first observation at five days. Feeding was noted but not estimated. Plant material was changed as needed.
4Number of larvae (replicates) in the category: 1 — only one observation with damage, usually the first; 2-3-two or three observations with
damage, >4 — greater than 4 observations with damage. When the total number of larvae is less than n, there were larvae that did not feed; when
total numbers are greater than n, some larvae fed before they were killed or missed during handling.
the five replicates of the treatment without plant ma-
terial in test D. Frass pellets on the floor of the cup
were counted as an indication of the amount of feed-
ing. In test C, the cups were examined at eight days
and at 15 days when the test was stopped. In test D,
the cups were examined mostly daily from Monday to
Friday until the larvae died. Survival and the number
of frass pellets were recorded at each examination, the
frass was removed, and plant material was changed as
needed.
RESULTS
Phenology. Based on larval sizes and larval and pu-
pal numbers, it appears that peak adult emergence oc-
curred in June, early August, and mid-October. There
were many large larvae at the end of March and some
pupae, which suggests that there was probably also an
emergence peak in April when we did not sample. Pu-
pae might have been absent during winter since we
found none in January, but we are unable to confirm
that because we did not sample in February. Only two
adults, one each in October and November, were ob-
served in the field, but some were undoubtedly pre-
sent most of the time based on our collections of other
stages. Grass along the shore was swept with a net in
October when pupae were common and the tempera-
tures still warm, but without success. This suggests
that adults rest away from the waterway during the day
or move to emergent plants elsewhere in the water-
way. However, we examined and collected emergent
plants at times throughout the year for studies with
other insect species, but we saw no adults.
Parapoynx seminealis eggs were found in August,
October, and November 1980, and June 1981. All sizes
of larvae were present from August 1980 to June 1981.
Larvae were common at the beginning of the project
in June and July 1980, but we did not record their sizes
when we placed the unopened larval cases in the rear-
ing cages. In late November there were small larvae
among the “bananas” attached to the plants, and there
were many small and large larvae among attached “ba-
nanas” in mid-January. There were also many at that
time in cases on the leaves. “Bananas” that had fallen
from the plants were common on the soil surface be-
neath plants in shallow water during a visit on 27 Oc-
tober 1999, but none were infested.
Plant damage. Larvae damaged the plants heavily
throughout the year. Large sections were cut from the
leaves by multiple larvae until there was often little re-
maining leaf material (Fig. 1 and Fig. 1 in Habeck
1974). Damaged leaves appeared to be more suscepti-
ble to pathogens especially Psewdomonas marginalis
(Brown) Stevens pv marginalis (Brown) Stevens which
caused the leaves to Henan mushy. Another patho-
cere te EE
114
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fic. 1. Parapoynx seminealis feeding damage on floating leaves of big floating-heart, Nymphoides aquatica. Arrow points to larval case on
lower left margin of the large leaf. Fic. 2. Mature larva of P. seminealis.
around the middle of the pupa. Fic. 4. P. seminealis male.
gen, Cercoseptoria nymphaeacea (Cke. & Ell.)
Deighton, was first recorded for Florida from our ma-
terial. Damage from neonatal boring produced small
holes in the leaves, which added to the leaf decline.
We had five sites that varied greatly in available leaf
material because of the damage from varying caterpil-
lar populations.
Eggs. The newly deposited egg is a flattened, ellip-
tical disc, length 0.90 + 0.01 mm (17, 0.84—0.94),
width 0.65 + 0.01 mm (0.62-0.66). If the floating leaf
was mature, each egg was deposited in a pit in the
arenchyma layer on the lower surface that has been
described as “dark-punctate or pitted” (Correll & Cor-
rell 1972). The margins of the eggs often overlapped
on the raised margins of the pits. Younger leaves with-
out a well-developed aerenchyma layer are smoother
and the eggs did not overlap as much. The center of
the fresh egg was pale yellow, but the margins were
clear. The chorion appeared roughened with longitu-
dinal ridges. The entire egg turned yellow as it aged,
but the color disappeared as the visible embryo ma-
tured. The light brown head capsule and dark pronotal
Fic. 3. P. seminealis pupa, note shiny air layer in the cocoon silk
shield signaled the approaching eclosion. Eggs devel-
oped in six days at 27°C and in eight to ten days at
about 24°C. The swollen egg was ovate when the em-
bryo was developed, length 0.80 + 0.01 mm (8,
0.76-0.81), diameter 0.55 + 0.01 mm (0.52-0.58).
Eggs were placed in loose rows, two to five eggs deep,
along the leaf margin. The rows were 0.9 to 4.7 mm
from the margin. The heads of most embryos were at
the end of the egg closest to the leaf margin, but a few
were oriented at the other end. Field egg masses were
relatively small, 14.6 + 2.6 eggs (15, 2-28).
Neonates. Upon eclosion, the neonate burrowed
into the thick leaf or dropped from the leaf on a silken
thread. The leaf entrance hole was often marked by
light green frass. The first instar was distinguished by
the lack of tracheal gills and by the dark brown prono-
tal shield and frontoclypeal triangle of the head cap-
sule. Most of the head capsule was light brown like
that of later instars. The larva tunneled either just be-
neath. the lower epidermis or from there to the upper
epidermis. Viewed from below, the tunnel beneath the
lower epidermis appeared green from the frass;
VOLUME 55, NUMBER 3
viewed from above, the tunnel beneath the upper epi-
dermis appeared as a dark blotch. However, if the leaf
were held towards a light, the upper tunnel appeared
transparent. The epidermis over the tunnel often
broke down forming a hole in the leaf through which
fungi invaded. Dark gut contents could be clearly seen
in the whitish, almost transparent, neonate that was of-
ten visible in the tunnel. Younger floating leaves were
sometimes too thin for a tunnel and the larva fed in a
trench on the lower surface. It also fed on the petiole
by scraping the surface or by boring into it. Submersed
leaves are much thinner than floating leaves and the
larva fed by scraping away all the material from the
lower surface. This created windows in the leaf. Older
first instars made small cases on the thin submersed
leaves by bending the leaf margin inwards along a cut
and fastening it to the leaf. Some fed on the “bananas”
and on the flower buds, both of which are attached just
below the floating leaves.
A larva dropping from a leaf through the water
could be manipulated by intercepting the silken
thread attached to the plant. The larva also attached a
thread to the forceps whenever handled. If food were
not available, larvae crawled actively around the con-
tainer and even crawled out of 30 ml plastic cups into
the air where they perished. When confined in capped
45 ml cups without water but with moist filter paper,
they lived from 5.5 to 114.5 h (mean = 58.1 + 7.0 h;
median = 64.5 h; n = 29). In cups with dry filter paper
all were dead at 2 h. Head capsules of living first in-
stars were 0.34 mm wide and the larvae were 1.8—2.4
mm long. Development time was four to 12 days with
most developing to second instars in five to eight days.
Older larvae. Second and later instars were distin-
guished readily from first instars by the presence of
tracheal gills. They also were whitish and somewhat
transparent with the gut easily seen. The head capsule
was very light brown except the last instar’s which was
pale yellowish to whitish. Second instars fed in tunnels
and externally on the lower surface and petiole, but
they also began making small cases on the floating
leaves and fed in channels on the upper surface of the
floating leaves. As the larvae matured, the case-making
habit intensified, but some larvae were also found
feeding without cases. Cases were formed in several
ways. A piece was cut from the edge of the leaf with
the size weakly related to the size of the larva. This cir-
cular to semicircular piece was either attached to the
upperside of the leaf by attaching the rough lower sur-
face of the piece to the smooth upper surface of the
leaf, or attached to the underside of the leaf by upper
surface to lower, or lower to lower. Attachment to the
upperside of the leaf was most common. Next most
common was attachment to the underside by upper
surface to lower surface. With both of these common
orientations, the excised piece was camouflaged as part
of the leaf.
To excise a piece, the larva started eating from the
edge of the leaf and ate a cut inward, revolving its body
around like the hand of a clock until reaching the mar-
gin again. The thickness of the leaf prevented the larva
from merely cutting the leaf as reported for the P.
maculalis (Clemens) case-making behavior on yellow
waterlily, Nuphar (Welch 1916). The larva then held
onto the excised piece with the hind legs and pulled it-
self onto the main leaf, dragging the piece behind it.
The piece was then attached to the leaf with silk. The
case was completed within an hour by medium to
large larvae. Small larvae formed a silken gallery inside
the case that pulled away from the leaf piece and re-
mained around the larva on the main leaf when the
case was pulled apart. Older larvae placed the silk in
two parallel rows thus forming a tunnel with the two
leaf pieces. When the case was pulled apart, the silk
remained attached to both leaves leaving the larva
naked. More silk was placed on the rough lower leaf
surface than on the smooth upper leaf surface. Large
larvae also webbed together the overlapping portions
of two leaves to make a case without cutting pieces.
When medium and large larvae made detached cases
the two excised pieces were usually placed lower sur-
face to lower surface. The case was thus green on both
sides. The first pieces cut from leaf margins were
roughly circular to semicircular, but as the leaves were
cut up, the shape of the excised pieces became highly
variable with multiple angles.
The size of the cases was also highly variable. Some
examples of the sizes of excised pieces used in field
collected cases of mid-instar larvae (10-15 mm long)
were 10 x 10, 10 x 15, 10 x 20, 10 x 25, 15 x 25, 20 x
20, 25 x 35, 40 x 75 mm; and of large larvae (20-25
mm long) were 15 x 25, 20 x 25, 20 x 35, 20 x 40, 25 x
30, 30 x 30, 30 x 35, 30 x 40, 30 x 45 mm. All of these
pieces were attached to floating leaves. Larvae in both
attached and detached cases fed along the margins of
the leaves and on the surfaces of the leaves near their
cases. They also exited from their cases to feed naked
on the leaf surface. Those on the upper surface ate
down to the lower epidermis forming patches or chan-
nels and occasional holes. Larvae also fed somewhat
on the inside of the cases. Feeding appeared to be
mostly nocturnal, but larvae were observed during the
day out feeding on the leaves, especially during the
cooler period of autumn and winter. Most cases were
attached along the leaf margin (Fig. 1) but many were
away from the margin near the center of the upper
116
surface. A thin layer of water from small holes in the
leaf or from wash over the sides usually surrounded
these cases. The larva often wiggled its body side to
side which apparently moved water through the case
as reported for P. maculalis (Welch & Sehon 1928).
The gills of the mature larva are very long, white, and
beautifully delicate (Fig. 2). Mueller and Dearing
(1994) described similar case construction on
Nymphaea ampla by Parapoynx rugosalis Méschler in
Costa Rica.
A graph of measurements of the head capsules of
233 preserved larvae from both laboratory experi-
ments and field collections did not have well defined
separations between instars. It suggested that there
might be as many as 10 or 11 instars in the field popu-
lations. These measurements and the presumed in-
stars were 0).32—0.36 mm (n = 17) I°, 0.40-0.44 (n = 9)
II°, 0.46-0.52 (n = 31) III°, 0.56-0.64 (n = 39) IV°,
0.66-0.74 (n = 32) V°, 0.76-0.90 (n = 30) VI°,
0.94-1.00 (n = 8) VII°, 1.04-1.22 (n = 30) VIII’,
1.28-1.40 (n = 14) IX°, 1.44-1.54 (m = 15) X°,
1.60-1.66 (n = 6) XI° (males?), 1.78-1.86 (n = 6) XI°
(females?). Measurements of living larvae that were
measured just before or after ecdysis confirmed the
sizes of the first nine instars. However, the maximum
head capsule size prior to pupation in the sample of
live larvae was 1.40 mm. Possibly host plant quality
was lower in the excised laboratory leaves and thus lar-
vae pupated sooner that they did in the field. Most of
the measurements above 1.40 mm in the preserved
specimens were from larvae collected directly in the
field. The approximate body lengths of larvae varied
from 3.2 mm for II° to 24.2 mm for mature larvae.
The larval development period at 27°C was 41.1 +
1.1 days (7, 36-44) and the development period from
neonate to adult was 51.4 + 1.4 days (5, 47-54).
Pupa. The mature larva formed a pupal chamber by
tying the case together completely around the edges. A
silken cocoon was formed in the case. The newly
formed pupa (Fig. 3) moved actively when disturbed,
but moved less as it matured. The central portion of
the cocoon appeared silvery because of air trapped in
the silk. Air was provided to the cocoon through small
elliptical feeding spots in the surface of the leaf made
before pupation. These spots, six to ten in a group,
were near the center of the cocoon and the silk di-
rectly above them was the most silvery. The spots were
0.40-0.60 mm long and 0.30-0.40 mm wide. Some of
the ridges on the lower aerenchyma surface of the at-
tached excised piece also had similar feeding spots.
There was a distinct prepupal stage when the light
yellow larva contracted into the three body regions
with very obvious abdominal segments. The newly
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
formed pupa was light yellow with three obvious or-
ange spiracles on projections of the abdomen (2.0-2.5
mm long on segments A2—A4); as the pupa matured,
the eyes darkened and the wing pads, legs, and anten-
nae turned white before darkening. The afternoon of
emergence, the pupa was brown with visible wing
markings. The female pupa was distinguished by an-
tennae that were obviously shorter than the wing pads
and by the mesothoracic legs, which terminated just
beyond the wing pads in the same body segment, A5.
The male’s antennae were as long or longer than the
wing pads, and the mesothoracic legs ended in the an-
terior portion of segment A6. The abdomen of the new
female pupa exceeded the hindlegs, but that of the
new male was usually shorter than the hindlegs. Near
emergence the male’s abdomen lengthened and ex-
ceeded the hindlegs. The widest part of the body was
between the spiracles of A3 and A4. The pupal size
was: female, length = 12.2 + 0.1 mm (9, 11.4-12.7),
width = 3.3 + 0 mm (3.2—3.5), width at spiracle = 3.8 +
0.1 mm (3.5-4.2); male, length = 10.4 + 0.1 mm (8,
10.0-11.0), width = 2.7 + 0 mm (2.7-2.8), width at
spiracle = 3.1 + 0.1 mm (2.8-3.3). Pupal development
time at 27°C was: female, 10.0 and 11.0 days (2): male,
10.0 + 0.8 days (5, 7-11).
Adults. Forewings of the adults are sexually dimor-
phic. Females have more or less unicolorous reddish
brown forewings compared with males that have gray-
ish forewings with a white stripe parallel to the side
and hind margins (Fig. 4). Both sexes have hindwings
with two transverse black stripes on a white ground-
color and tan or orangish markings along the hind mar-
gin (for a detailed description see Monroe 1972).
Adults emerged in outdoor cages all day although
probably mostly at night because the largest numbers
were present in early morning. Observed night emer-
gence times were 2000 h, 2100 h, and 2100-2245 h for
three females and 1800-2100 h for one male.
The newly emerged female calls the male by raising
her abdomen above her resting wings and extending
the tip. One mating was at 0200 h and lasted less than
an hour. Colony females generally mated the next
evening after they emerged. Females typically rested
with their wings folded over their body. Males often
rested with their wings extended so that the hind
wings were visible, although they also rested with
folded wings, especially after a disturbance. Adults
were not strongly attracted to light in the laboratory.
When they escaped, they flew erratically, landing on
the walls rather than flying to the window or overhead
light. .
Oviposition was observed at 2000 h in an outdoor
tank, and at 2100 h and later in laboratory cages. Fe-
VOLUME 55, NUMBER 3 117
TaBLE 2. No-choice feeding tests with medium-sized larvae of Parapoynx seminealis.
No. frass
Test Longevity (days) pellets/day *
Plant species! symbol? Mean range n Mean SE
Egeria densa C 15.0 15.0 8 9.3 2.3
Hydrilla verticillata C 15.0 15.0 W 5.0 0.5
Nymphaea odorata Aiton C 15.0 15.0 8 8.0 7
Nymphoides aquatica C 15.0 15.0 8 62.4 6.8
Potamogeton illinoensis C 8.9 0.9 8-15 8 0.5 0.1
Cage without plant material D 6.0 6.0 5 0.4 0.2
Egeria densa D 51.3 29.6 6-107 3 1252, 3.5
Hydrilla verticillata D 46.7 20.9 13-85 3 13.6 1.8
Limnobium spongia D 18.7 4.7 14-28 3 10.7 3.8
Nymphaea odorata D 26.7 8.1 12-40 3 14.7 2.0
Nymphoides aquatica D 66.3 14.0 47-93 3 98.5 0.6
Sagittaria subulata D 17.3 43 13-26 3 3.6 3.0
Vallisneria americana D 23.7 2.3 19-26 3 13.6 1.4
Vallisneria americana D 18.0 4.1 7-28 5 9.7 3.6
‘ Nymphaea is in the Nymphaeaceae, N. aquatica is the field host plant.
Cages were 177 ml styrofoam cups with medium-sized larvae reared in the laboratory at 22—24°C on big floating-heart. Test C was termi-
nated after 15 days; test D terminated when all larvae died.
5Individual frass pellets were counted and removed every one to three days. Plant material was changed as needed.
males sat at the edge of the leaf perpendicular to the
margin with the fore and mid legs on the leaf and the
abdomen curled under the leaf. They also sat halfway
on the leaf parallel to the margin with the abdomen
curved sideways under the leaf. Realized fecundity in
the laboratory was 293.3 + 25.5 eggs (17, 56-412).
Thirteen of the females laid the largest number of eggs
during their first night of oviposition, which was one to
three nights after emergence, and eight of those laid
the majority of their eggs during the first night after
emergence. None oviposited during the night they
emerged or after the fifth night. Longevity in the labo-
ratory was 4.0 + 0.4 days (16, 2-6) for females and 3.4
+ 0.4 days (14, 2-5) for males.
Host range tests. Some neonates fed briefly on
each test plant species, but most feeding was before the
first change of plant material (Table 1). Notable feeding
and longevity was observed in test A on Egeria densa
Planchon and Hydrilla verticillata (L. fil.) Royle, both in
the family Hydrocharitaceae. Longevity and feeding
were similar among the other test species in both tests.
Larvae did not complete development in test A on float-
ing-heart because of the lack of fresh plant material at
the end of the field season. Although test B was termi-
nated at 22 days because of lack of floating-heart, that
had little impact on the longevity data for the test plants
because only three of 78 test larvae were still alive at 22
days. It did, however, greatly reduce the longevity for
floating-heart, which had 12 of 20 larvae still alive. Small
differences in longevity among test plants might have
been masked by the relatively long intervals of three to
seven days between changes of plant material.
Medium-sized larvae that had developed on floating
heart until tested survived longer than neonates and
fed more on some of the same plant species (Table 2).
Again, longevity on test plants was highest on E. densa
and H. verticillata (test D), but feeding (as indicated
by mean number of frass pellets per day) was similar
on all test plants except on the little eaten Potamoge-
ton illinoensis Morong (Potamogetonaceae) and Sagit-
taria subulata L. Buch (Alismataceae). In test C, most
feeding on N. odorata was during the first eight days,
13.2 frass pellets per day, compared with 2.0 during
the final seven days. It was vice versa in the other four
species.
DISCUSSION
The life cycle of P. seminealis is quite similar to that
reported for P. maculalis on Nuphar (Nymphaeaceae)
(Welch 1916). One difference is that P. maculalis pre-
ferred to oviposit through oviposition holes of Donacia
leafbeetles (Chrysomelidae) (Welch 1916), which were
not present on big floating-heart.
This study has confirmed the limited observations
on larval behavior and feeding reported by Forbes
(1910), Habeck (1974), and Monroe (1972). Kimball
(1965) reported adults collected throughout Florida in
every month of the year, but not in north Florida dur-
ing December and January, which agrees with our
phenology data. This is the only species of Parapoynx
recorded feeding on floating-hearts in North America,
but a polyphagous relative, Synclita obliteralis
(Walker) (Crambidae: Nymphulinae) has also been
recorded from it (Habeck 1991). In Europe, the re-
ee
oe
i;
|
118
lated floating-heart, N. peltata (Gmel.) O. Kuntze, is
attacked by Nymphula nymphaeata L. (Crambidae:
Nymphulinae) and possibly occasionally by Cataclysta
lemnata, (L.) (Crambidae: Nymphulinae) (Van Der
Velde 1979). We were not surprised that the host feed-
ing range in the laboratory was wider than that ob-
served in the field, but we were surprised that the two
plant species most heavily eaten were in the monocot
family Hydrocharitaceae. Both species, E. densa and
H. verticillata, are immigrants in North America and
are thus new associations with P. seminealis. No host
records were found for P. seminealis on the related
North American genus Elodea. Interestingly, John and
Nanjappa (1988) in India reported that Parapoynx
diminutalis (Snellen), a native moth common on H.
verticillata, fed upon and destroyed Nymphoides
cristata (Roxb.) O. Kuntze. This moth, an immigrant
in Florida, did not eat big floating-heart in our labora-
tory host range tests with it (Buckingham & Bennett
1989). The current study with P. seminealis demon-
strates both the difficulty of accurately assessing in the
laboratory the narrowness of the feeding host range of
a biological control agent and the importance of good
field host range data in the native range for interpret-
ing laboratory data. We did not demonstrate that
caterpillars could complete development on the test
plants. However, if field data were lacking, the
amounts of laboratory feeding we found might pre-
clude use of even a truly specialized species like this as
an introduced biological control agent.
ACKNOWLEDGEMENTS
We thank Ms. Bonnie Ross for help in all phases of this study and
Dr. Dale H. Habeck and Dr. Sanford Porter for review of the man-
uscript. Plant pathogens were identified by the Bureau of Plant
Pathology and test plants by the Botany Section, Division of Plant
Industry, Florida Department of Agriculture and Consumer Ser-
vices, Gainesville, Florida. Partial funding was provided by the Wa-
terways Experiment Station, U.S. Army Engineers, Vicksburg, Mis-
sissippi. This study was conducted cooperatively by the ARS-USDA
and the University of Florida-IFAS through a Research Support
Agreement.
LITERATURE CITED
BUCKINGHAM, G. R. 1994. Biological control of aquatic weeds,
Chapter 22, pp. 413-479. In D. Rosen, F. D. Bennett & J. L.
Capinera (eds.), Pest management in the subtropics: biological
control—the Florida experience. Intercept Ltd., Andover,
Hampshire, United Kingdom.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
BUCKINGHAM, G. R. & C. A. BENNETT. 1989. Laboratory host range
of Parapoynx diminutalis Snellen (Lepidoptera: Pyralidae), an
Asian aquatic moth adventive in Florida and Panama on Hy-
drilla_verticillata (Hydrocharitaceae). Environ. Entomol.
18:526-530.
. 1996. Laboratory biology of an immigrant Asian moth,
Parapoynx diminutalis (Lepidoptera: Pyralidae), on Hydrilla
verticillata (Hydrocharitaceae). Florida Entomol. 79:353-363.
CorRELL, D. S. & H. B. Correuu. 1972. Aquatic and wetland
plants of southwestern United States. Vol. 2. Stanford Univ.
Press, Stanford, California. 1777 pp.
ForBes, W.T. M. 1910. The aquatic caterpillars of Lake Quinsiga-
mond. Psyche 17:219-227.
Goprrey, R. K. & J. W. WooreEN. 1981. Aquatic and wetland
plants of the southeastern United States. Univ. Georgia Press,
Athens, Georgia. 933 pp.
HaBeck, D. H. 1974. Caterpillars of Parapoynx in relation to
aquatic plants in Florida. Hyacinth Contr. J. 12:15-18.
. 1991. Synclita obliteralis (Walker), the waterlily leafcutter.
(Lepidoptera: Pyralidae: Nymphulinae). Division of Plant In-
dustry, Florida Department of Agriculture and Consumer Ser-
vices, Gainesville, Florida, Entomol. Circ. 345. 2 pp.
HERLONG, D. D. 1979. Aquatic Pyralidae (Lepidoptera: Nym-
phulinae) in South Carolina. Florida Entomol. 62:188-193.
Joun, G. M. & C. Nanyappa. 1988. 26. Parapoynx diminutalis
Snellen (Pyralidae: Lepidoptera) as a serious pest of
Nymphoides cristatum in Keoladeo National Park, Bharatpur,
Rajasthan. J. Bombay Nat. Hist. Soc. 85:637.
JuLiEN, M. H. & M. W. GrirritHs. 1998. Biological control of
weeds: a world catalogue of agents and their target weeds. 4th
ed. CAB International, Wallingford, United Kingdom. 223 pp.
KIMBALL, C. P. 1965. The Lepidoptera of Florida, an annotated
checklist. Division of Plant Industry, Florida Department of
Agriculture, Gainesville, Florida. 363 pp.
Monrok, E. 1972. Pyraloidea, Pyralidae (Part). Fasc. 13.la In
Dominick, R. B. et al. (eds.), The Moths of North America
north of Mexico. E. W. Classey Ltd. and R. B. D. Public., Lon-
don. 134 pp.
MUELLER, U. G. & M. D. DEARING. 1994. Predation and avoid-
ance of tough leaves by aquatic larvae of the moth Parapoynx
rugosalis (Lepidoptera: Pyralidae). Ecol. Entomol. 19: 155-158.
VAN DER VELDE, G. 1979. Nymphoides peltata (Gmel.) O. Kuntze
(Menyanthaceae) as a food plant for Cataclysta lemnata
(L.)(Lepidoptera, Pyralidae). Aquat. Bot. 7:301-304.
WELCH, P. S. 1916. Contribution to the biology of certain Lepi-
doptera. Ann. Entomol. Soc. Am. 9:159-190.
WELCH, P. S. & G. L. SEHON. 1928. The periodic vibratory move-
ments of the larva of Nymphula maculalis Clemens (Lepi-
doptera) and their respiratory significance. Ann, Entomol. Soc.
Am. 21:243-258.
GENERAL NOTES
Journal of the Lepidopterists’ Society
53(3), 2001, 119
NYCTEOLA FRIGIDANA WALKER (NOCTUIDAE: SARROTHRIPINAE)
REPORTED AT AN UNORTHODOX BAIT
Additional key words: | Gadway Barrens, New York, Salix bebbiana, insect remains.
On 15 July 1998, while attempting to compare the ef-
ficacy of two different types of bait, I noted a common-
place occurrence. Yellow jackets (Hymenoptera: Vespi-
dae; Dolichovespula arenaria (Fab.)) were feeding on
the fresh spattered insect remains on the front of my ve-
hicle. An hour later, at dusk, I hung out two 30—meter
long strands of cotton clothesline rope that were soaked
in different bait formulas: the more traditional
beer/sugar/molasses bait (Holland 1903) was being com-
pared to a simple bait of red wine saturated with sugar. I
ran the trials through uniform jack pine/blueberry habi-
tat on the Gadway Barrens, Clinton County, New York.
The vehicle I used for transportation was parked in
a 10-meter gap between the bait trials. I would pass
the front of the vehicle each time I traversed the two
trials. No apparent differences in habitat existed in the
immediate sample area.
Five Nycteola frigidana (W|k.) were observed prob-
ing the fresh remains of insects spattered over the
windshield and front of the vehicle over the course of
the night. One Caripeta piniata (Pack.)(Geometridae)
and one Catocala gracilis Edw. (Noctuidae) were also
recorded probing the insect remains. The insect re-
mains on the parts of the vehicle where the N. frigi-
dana were observed feeding were determined to be
mostly Diptera and definitely not that of Lepidoptera.
The two baited ropes produced many Idia aemula
Hbn., I. americalis (Gn.), I. lubricalis (Gey.), Catocala
similis Edw., Apamea amputatrix (Fitch), A. lignicol-
ora (Gn.), Phlogophora periculosa Gn., Apharetra
dentata (Grt.), Pseudaletia unipuncta (Haw.), Leuca-
nia pseudargyria Gn., Agrotis ipsilon (Hufn.), Noctua
Journal of the Lepidopterists’ Society
55(3), 2001, 119-121
pronuba L., Graphiphora auger Fab. (all Noctuidae),
and Caripeta piniata (Pack.)(Geometridae), but no N.
frigidana or Catocala gracilis.
Nycteola frigidana comes poorly to both bait and
light. I have taken only the occasional specimen at tra-
ditional bait on the Gadway Barrens. In June, Nycteola
caterpillars can be found commonly on Salix bebbiana
Sarg. (Salicaceae) at this site. Additional information
on the range and systematics of Nycteola can be
gleaned from several sources (Fletcher 1959, McDun-
nough 1943, Rindge 1961). It is apparent that alterna-
tive methods of sampling are possible for these diffi-
cult-to-attract moths.
As an aside, I detected no significant differences in
numbers or composition of species at the two types of
bait being tested.
I thank Chris Weber for assistance in the field and
in rearing caterpillars. Kathy Schneider and Ed Stain-
ton introduced me to the Gadway Barrens site.
LITERATURE CITED
FLETCHER, D.S. 1959. Notes on North American species of Nycte-
ola (Lepidoptera, Noctuidae). J. N.Y. Entomol. Soc. 67:51-53.
HOLLAND, W. J. 1903. The moth book. New York: Doubleday, Page
& Co. 479 pp.
McDunnoucu, J. 1943. Phalaenid notes and descriptions (Lepi-
doptera). Can. Entomol. 75:59-62.
RINDGE, F. H. 1961. A synopsis of the genus Nycteola from North
America, including a new species from Arizona (Lepidoptera:
Noctuidae). J. N.Y. Entomol. Soc. 69:203-206.
TimoTHy L. McCCaBE, New York State Museum,
Cultural Education Center, Albany, New York 12230,
USA.
HEPIALUS CALIFORNICUS (HEPIALIDAE) OVIPOSITION PREFERENCE ON THE LUPINE
LUPINUS ARBOREUS
Additional key words: dispersal, tanglefoot, aerially-dispersed eggs.
One of the consequences of complete metamorpho-
sis in Lepidoptera is that larvae and adults experience
very different environments and selective pressures.
Adult Lepidoptera are far more mobile than larvae, al-
lowing use of a larger portion of the habitat. Adults
make important decisions regarding host plants and the
location of oviposition sites on this larger scale, deci-
sions that greatly affect larval survival (Setamou et al.
1999). While many Lepidoptera demonstrate specificity
in host plant oviposition sites (e.g., Haribal & Renwick
1998), it is less clear whether Lepidoptera that aerially
disperse their eggs are similarly selective. Falling into
the latter category is the ghost moth Hepialus californi-
cus (Lepidoptera: Hepialidae), a nocturnal moth native
to the West Coast of the United States. Males of H. cal-
ifornicus perch on vegetation and release pheromones,
forming leks at dusk and dawn to which females are at-
tracted (Wagner 1985). Gravid females find a host plant
for the larvae and release their eggs while hovering over
the vegetation. Other species of Hepialidae, notably
Korscheltellus gracilis, also exhibit similar ‘hovering
oviposition behavior (Wagner et al. 1989, Wagner &
Rosovsky 1991). The primary host plant for H. californi-
cus at Bodega Bay is the bush lupine Lupinus arboreus,
but larvae are polyphagous and have occasionally been
found on Eriophyllum staechadifolium, Helenium pu-
berulum, Rumex sp. and Rubus sp. (Wagner 1985).
Wagner (1985) suggested that females scatter their
eggs over a wide range of potential host plants and sur-
faces, an assertion bolstered by a documented case of H.
californicus ovipositing over asphalt. Widely-dispersed
egg-laying by H. californicus at BML is questionable in
light of: (1) the larval dependence on the bush lupine
host plant for survival; and (2) the ability of Lepidoptera
to detect the CO, signal of host plants (Stange 1997).
Wagner (1985) and others (Strong et al. 1995, 1996)
noted that H. californicus eggs are deposited into the
detritus underneath lupine bushes, where the larvae
hatch and have to find a lupine stem or root in which to
burrow. Since the larvae are small (less than 2 mm at
hatching) and extremely vulnerable, and the vast major-
ity of H. californicus larvae at BML are found on bush
lupines, being deposited closer to the lupine’s stems and
roots would seem to improve the chances of larval sur-
vival. In addition, larger lupine bushes have larger stems
and roots, and H. californicus larvae are found on the
largest stems and roots within a bush (pers. obs.). In this
note I present data on non-random H. californicus
oviposition within bush lupine, along with a technique
for collecting aerially-dispersed eggs.
I tested the hypothesis that H. californicus oviposits
non-randomly and has an increased frequency of ovipo-
sition near the center of lupine bushes. I collected H.
californicus eggs at two sites at the Bodega Marine Re-
serve, Bodega Bay, Califomia, USA. One was a large
patch of lupine bushes east of the marine station (Upper
Draw) and one was to the west of the marine station on
the lee side of a hill (Mussel Point). At each site, I iden-
tified 14 lupine bushes for sampling (28 bushes total). I
measured each bushes’ length, width, and height in me-
ters. Each bush had four 22.2 cm diameter white plastic
plates, covered in “tanglefoot” sticky trap, placed at
ground level beneath it. The four plates were allocated
to ‘interior’ and ‘perimeter’ sites as shown in Fig. 1. Ini-
tially, I also placed traps one meter outside the bushes to
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
check for outside-bush oviposition; however, after three
trapping dates there had been no eggs laid on the out-
side-bush plates and I concentrated on within-bush sam-
pling. There were a total of 56 traps/site x 2 sites, for a
total of 112 traps. The traps were placed under bushes
before sunset (between 1730 and 1900 h, depending on
date) and collected in the morning starting at 0645 h. I
examined each trap for ghost moth eggs, which were re-
moved after being counted. The numbers of eggs on the
two ‘perimeter’ plates were added to get each bush’s
perimeter number of eggs, and two ‘interior’ plates were
added to get the interior number of eggs. I sampled dur-
ing the mating and oviposition period of H. californicus
over six dates, from late March through early May, 2000.
Over six sampling dates I recovered 187 eggs: 125
eggs in interior traps and 62 in perimeter traps. This
difference between the two areas was significant (chi-
square: df = 1, p < 0.001). This trend was significant
across both sites (Fig. 2) and was supported by an
analysis of how many traps at each location caught
eggs. Of the 23 (out of 28) traps that caught eggs, 13
had interior > perimeter eggs, 7 had perimeter > in-
terior eggs, and 3 had equal numbers — a significant
difference (chi-square: df = 2, p < 0.01). In compari-
son, an analysis of bush height, volume (length x
width x height), or ‘ground coverage’ (length x width)
on the total number of H. californicus eggs laid found
no significant correlation (height: df = 1, 26, F = 2.19,
<< While \oltmnanes Che = 1 AS, 18 = Boi, jo « W.2
ground coverage: df = 1, 26, F = 2.59, p < 0.12).
Despite variation in the numbers of eggs oviposited
between sites and bushes, I did note a preference by
H. californicus for oviposition in the center of lupine
Lupine Bush °
Gina
Fic. 1. Trap placement in lupine bushes. The two perimeter and
interior traps are added together for the number of perimeter and
interior eggs collected per date, respectively.
VOLUME 55, NUMBER 3
. Bush Interior
Bush Perimeter |
Mussel Point Upper Draw
Site
Fic. 2. Total eggs caught per site vs. placement. At both sites,
more eggs were caught in interior versus perimeter traps.
bushes. This conclusion is supported by comparisons
of both the total numbers of eggs laid as well as the
number of bushes with more eggs in the interior than
on the perimeter. This preference is reasonable given
the additional risk incurred by H. californicus larvae
that have to travel extensively through a predator-filled
zone like the detritus (D. Strong pers. com.). Females
capable of detecting the thickest part of a lupine bush
and releasing their eggs closer to it would thus in-
crease larval survival.
The above logic might also be applied to oviposition
that depends on the size of the lupine bushes; how-
ever, there was no evidence of any correlation be-
tween oviposition and several indices of bush size. It
may be that intraspecific competition for lupine stems
and roots is so low due to larval mortality that bushes
have approximately the same chance of supporting
larvae; however, this is impossible to evaluate without
quantifying larval mortality in the detritus. Wagner
(1985) suggested that female moths may preferen-
tially oviposit on the lee- versus wind-ward side of
bushes, something that may be especially important
given BMLs strong coastal winds. This hypothesis,
however, was not tested in the current study.
The sampling technique used here makes it possi-
ble to quantify the amount of “egg rain’ that lupine
bushes experience due to H. californicus, an impor-
tant factor in establishing a life-cycle model of H.
californicus for addressing larger ecological issues.
Some of the questions that could! be answered by
this technique in conjunction with additional experi-
ments include: do different sites have different num-
bers of eggs, or at different times? Does the location
of leks influence where females oviposit? What is the
survival rate of eggs deposited close to roots versus
at the perimeters of bushes? All of these questions
could be answered with further applications of this
technique, providing a better sense of the ecological
and population dynamics of Hepialus californicus.
I would like to thank Rick Karban, Don Strong, Lynne
Adler and the Bodega Marine Lab support staff. This
project was supported by in-class funds from the UC-
Davis Population Biology and Entomology departments.
LITERATURE CITED
Haripa., M. & J. A. RENWICK. 1998. Differential postalightment
oviposition behavior of monarch butterflies on Asclepias
species. J. Insect Beh. 11:507-538.
SETAMOU, M., F. SCHULTHESS, N. BOSQUE-PEREZ, H. POEHLING,
& C. BORGEMEISTER. 1999. Bionomics of Mussidia ni-
grivenella (Lepidoptera: Pyralidae) on three host plants. Bull.
Ent. Res. 89:465—471.
STANGE, G. 1997. Effects of changes in atmospheric carbon diox-
ide on the location of hosts by the moth, Cactoblastis cacto-
rum. Oecologia (Berlin) 110:539-545.
STRONG, D. R., J. L. Maron, P. G. CoONNors, A. WHIPPLE, S. HaR-
RISON & R. L. JEFERIES. 1995. High mortality, fluctuation in
numbers, and heavy subterranean insect herbivory in bush
lupine, Lupinus arboreus. Oecologia (Berlin). 104:85-92.
STRONG, D. R., A. V. WHIPPLE & A. L. CHILD. 1996. Notes from
underground: Heavy insect herbivory and potent natural en-
emies. Bull. Ecol. Soc. Am. 77:429.
WacneER, D.L. 1985 The biosystematics of Hepialus F. s. lato: with
special emphasis on the californicus-hectoides species group,
UC Berkeley Ph.D. thesis, 391 pp.
Wacn_er, D. L. & J. Rosovsky. 1991. Mating systems in primitive
Lepidoptera, with emphasis on the reproductive behavior of
Korscheltellus gracilis (Hepialidae). Zool. J. Linnean Society.
102:277-304.
WacNneER, D. L., D. R. Tost, B. L. PARKER, W. E. WALLNER & J. G.
LEONARD. 1989. Immature stages and natural enemies of Ko-
rscheltellus gracilis (Lepidoptera: Hepialidae). Ann. Ent. Soc.
Am. 82:717-724.
EVAN PREISSER, Population Biology Graduate
Group, University of California at Davis, Davis,
California 95616, USA. Phone: (530) 756-4586; Email:
elpreisser@ucdavis.edu.
wo a
PTS TT
Journal of the Lepidopterists’ Society
55(3), 2001, 122-123
STATUS OF PIERIS VIRGINIENSIS (PIERIDAE) IN NEW YORK STATE
Additional key words: conservation, Brassicaceae.
Since the introduction of the cabbage butterfly, Pieris
rapae (L..) (Pieridae), into Quebec around 1860 (Scud-
der 1889), many changes in the distributions and abun-
dances of members of the genus Pieris have been ob-
served in northeastern North America. Pieris rapae has
spread through most of the continent, while P oleracea
Harris and P. virginiensis Edwards disappeared from
many localities. Competition from P. rapae was thought
to be the cause for the decline of the native species
(Scudder 1889, Forbes 1926), but P. rapae does not fre-
quent forested habitats and the two native species pre-
fer to remain beaneath the forest canopy (Klots 1935,
Chew 1981). The likely cause of the decline of the na-
tive species was the reduction of forest habitat (Klots
1935, Chew 1981, Cappuccino & Kareiva 1985), possi-
bly exacerbated by the introduction of non-native Bras-
sicaceae which are lethal to larvae (Bowden 1971, Chew
1978) but to ovipositing females are not distinguished
from the natural hosts (Chew 1980).
I attempted to clarify the current distribution of P.
virginiensis in New York State to assess the need for
attention by the conservation community. The present
distribution and abundance of P. virginiensis in New
York State is poorly understood, partly due to the long-
standing synonymy with P. oleracea which is sympatric
with P. virginiensis in New York State (e.g., Hovanitz
1963, dos Passos 1965, 1966). Historic reports without
specimens for verification are not reliable. However,
the loss of well-known P. virginiensis colonies and a
paucity of modern reports are seen as symptoms of a
general decline within New York State (Forbes 1926,
Shapiro 1974). Similar concerns in Ontario proved un-
warranted after previously unknown populations were
located in the 1980s (Layberry et al. 1998).
I searched the literature for historic (before 1974)
and recent (1974-1999) reports of P. virginiensis
within New York State. The dividing point between
“historic” and “recent” was based on reports that
Shapiro (1974) would have been aware of. In addition
to the literature, the collections at the New York State
Museum and Cornell University were examined, the
New York Natural Heritage Program database was
consulted, and localities with appropriate habitat were
visited during the flight periods in 1998-1999.
Pieris virginiensis is broadly distributed in New York
(Fig. 1), having been documented from 28 counties. Re-
cent accounts were found from 12 counties and historic
accounts were found from 22 counties. During field sur-
veys I encountered the species at 12 localities within 7
counties. Five recent localities were documented in the
News of the Lepidopterists’ Society Season Summary
(1974, 1977, 1987, 1997). Two recent reports were found
in the New York Natural Heritage Program database.
Sixteen counties have reports dating only from before
1974, while six have strictly recent accounts. Only two lo-
calities have reports dating from both time periods.
Localities in which I searched forested habitat for
populations included areas in the vicinity of historic
records and areas near localities where I found extant
populations. Regions from which historic documenta-
tion of the species exists but where I did not locate ex-
tant populations were the southern Catskill Mts. in
Sullivan and Ulster Counties, and the area around
Trenton Falls in both Oneida and Herkimer Counties.
Extensive deciduous forest remains in these areas, but
most of the potential habitat is inaccessible. I did not
search in the eastern portions of the Catskill Moun-
tains or the western counties.
The narrow overlap of historic and recent accounts
of P. virginiensis within New York State suggests that
the information available is insufficient to elucidate
changes in distribution. Newly discovered localities on
the Tug Hill Plateau and central and western counties
do not necessarily represent newly colonized habitat.
The presence of P. virginiensis in southern Ontario
and one historic specimen from Potsdam (St.
Lawrence County, New York) (Shapiro 1974) suggest
the species has gone unnoticed in the Tug Hill region
Fic. 1. Map of New York State showing old (before 1974) and re-
cent (1974-1999) collection localities for Pieris virginiensis (open
square, old; solid square, recent; x, both time periods).
VOLUME 55, NUMBER 3
until my explicit attempts to locate it there. The possi-
bility also exists that historic records of P. oleracea in-
cluded individuals of P. virginiensis, although all spec-
imens I have seen were correctly identified.
Populations in western counties may also have long
escaped notice. Pieris virginiensis was last reported
from the Buffalo metropolitan area in 1873 (Forbes
1926), but has recently been found in adjacent Chau-
tauqua County (Season Summary 1987). Reports from
the vicinity of Olean in Cattaraugus County spans both
time periods considered here (Shapiro 1974, Season
Summary 1993) but could have included an extirpa-
tion followed by recolonization, as was documented in
McLean Bog (R. Dirig, Season Summary 1974).
The possibility also exists that P. virginiensis popula-
tions have shifted hosts and are moving into areas previ-
ously unoccupied. Courant et al. (1994) and Porter
(1994) discussed the strong selective pressures associ-
ated with the use of non-native Brassicaceae by native
pierines. Porter (1994) presented three possible out-
comes for a population encountering the spread of non-
native Brassicaceae, including the initial decline of the
native Pieris population as the lethal effects of the new
host remove individuals feeding on the lethal host, fol-
lowed by an increase due to two potential responses. Se-
lection would favor females that recognize the lethal
host and refuse to oviposit, or selection would favor lar-
vae capable of utilizing the new host. The third outcome
would be loss of populations as the non-native, lethal
host expands its range and dominance across the land-
scape. The host association of populations at most of the
localities in which I found P. virginiensis could not be
ascertained. All sites where large colonies were found
were associated with native Dentaria. The relationship
between P. virginiensis and the native and non-native
Brassicaceae continues to be a question requiring re-
search. The populations in and around Tompkins and
Cortland Counties, which include the recolonized
McLean Bog, may offer an arena for such observation.
The loss of forest continues to be the greatest threat
to the viability of P. virginiensis populations. Historic
extirpations typically have been noted in areas of urban
expansion. Colonies are occasionally found in wood-
lands within suburban areas. These habitats are more
suitable to P. rapae, but the responses of native Pieris
species to habitat conversion are not understood.
Although P. virginiensis is likely less common than in
the past (Cappuccino & Kareiva 1985), it remains in
many forested areas throughout New York State. How-
ever, no conclusion regarding its long-term viability can
be made until additional information on the population
dynamics within a series of localities in New York State is
gathered. The number of unknown populations I found
during only a few days within two flight seasons indicates
that additional surveys will find this species more wide-
spread than is made apparent by the distribution map in-
cluded here (Fig. 1), particularly in the Catskill Moun-
tains and the vicinity of Allegany State Park. Additional
survey efforts should also be given to localities with his-
toric documentation but lack recent specimens.
I would like to thank James Liebherr for permitting
access to the collection at Cornell University, Tim Mc-
Cabe for supplying label data from specimens in the
New York State Museum, Kathy Schneider for supply-
ing copies of reports in the New York Natural Heritage
Program database, and F. Chew and A. Porter for their
reviews of this manuscript. Vouchers taken during
field studies were deposited in the collection of the
New York State Museum in Albany.
LITERATURE CITED
BowDEN, S. R. 1971. American white butterflies and English food-
plants. Journal of the Lepidopterists Society 25:6-12.
Cappuccino, N. & P. KareIva. 1985. Coping with a capricious en-
vironment: a population study of a rare pierid butterfly. Ecology
66:152-161.
CHEW, F. S. 1978. Introduced plants as food resources for native
cabbage butterflies. Atala 5:13-19.
. 1980. Foodplant preferences of Pieris caterpillars (Lepi-
doptera). Oecologia 46:347-353.
. 1981. Coexistence and local extinction of two pierid but-
terflies. American Naturalist 118:655-672.
CourRANT, A. V., A. E. HOLBROOK, E. D. VAN DER REIJDEN & F-. S.
CueEw. 1994. Native pierine butterfly (Pieridae) adapting to
naturalized crucifer? Journal of the Lepidopterists Society
48:168-170.
bos Passos, C. F. 1965. Review of the Nearctic species of Pieris
“napi” as classified by androconial scales and description of a
new seasonal form (Lepidoptera: Pieridae). Journal of the New
York Entomological Society 73:135-137.
. 1966. Pieris narina oleracea (Harris) in New Jersey (Lepi-
doptera: Pieridae). Journal of the New York Entomological So-
ciety 74:222-223.
Forbes, W. T. M. 1926. Lepidoptera. In M. D. Leonard (ed.), A
List of the Insects of New York, Memoirs of the Cornell Uni-
versity Agricultural Experiment Station 101.
Hovanitz, W. 1963. The relation of Pieris virginiensis Edw. to
Pieris napi L. Species formation in Pieris? Journal of Research
on the Lepidoptera 1:124—134.
Kiots, A. B. 1935. On the life history of Pieris virginiensis Edws.
Journal of the New York Entomological Society 43:139-142.
LayBERRY, R. A., P. W. HALL & J. D. LAFONTAINE. 1998. The but-
terflies of Ganndas Toronto: University of Toronto Press.
Porter, A. 1994. Implications of introduced garlic mustard (Al-
liaria petiolata) in the habitat of Pieris virginiensis (Pieridae).
Journal of the Lepidopterists Society 48:171-172.
SCUDDER, S. H. 1889. The butterflies of the eastern United States
and Canada. Cambridge, MA.
SEASON SUMMARY. 1974. 1977. 1987. 1997. Annual supplement to
News of the Lepidopterists’ Society.
SHAPIRO, A.M. 1974. Butterflies and skippers of New York State.
Search 4:1-59.
Epwarb J. STANTON, 29726 Avenida de Cortez, Sun
City, California 92586, USA.
BOOK REVIEWS
Journal of the Lepidopterists’ Society
53(3), 2001, 124
FLYING JEWELS—BUTTERFLY IMAGES, by Yasutaka
Murata & Daisaburo Okumoto. 2001. Publisher:
Shueisha Inc, 5-14, Sarugaku-cho, 1-chome, Chiyoda-
ku, Tokyo, 101-8050 Price: 4,410 Japanese yen
[written in Japanese] ISBN4-08-532058-0.
This slim book is a collection of 76 butterfly por-
traits photographed in natural habitats. The photo-
graphic localities range from Japan, China, Indonesia,
Europe, North America, Mexico, Central and South
America. Each photograph is accompanied by minimal
text describing the location and behavior of the butter-
flies, and occasionally, the impression of the authors. I
confess that, given the number of pretty picture books
in the world dedicated to butterflies, I did not antici-
pate being taken by Flying Jewels—Butterfly Images.
However, it became clear that this book includes im-
ages that are truly unique, beautiful and show a patient
understanding of both photography and subject.
In particular I was impressed by those images taken
with a 15 mm lens. These not only provide an aston-
ishing butterfly’s view of the world, but considering the
balance of light, the user hostility of a 15 mm lens, and
the color balance that is involved in film photography,
these images are mini-miracles. How does Morpho
amathonte, Teinopalpus aureus or Papilio glaucus see
the world when flying along a forest edge? What does
the world look like to puddling males of Agehana
maraho or Trogonoptera trojana? What view of the
world does Lycaena phaleas have when feeding on a
sunflower, or Polygonia c-aureum when feeding on a
persimmon fruit that still hangs from the tree? Thanks
to the images in this book, we have come much closer
to knowing the answers.
Following these extraordinary images there is a sec-
tion (in Japanese) stressing the worldwide need for
conservation to preserve butterflies in particular, and
natural habitats in general. The final text encompasses
brief technical descriptions of where each photo was
taken and what equipment was employed. Although
Flying Jewels — Butterfly Images is decidedly not a
scholarly work, the images in this book form a connec-
tion between the reader and the projected sense of
wonder of the photographer for the subject. And that
is natural, reflective art.
P. J. DeVriEs, Center for Biodiversity Studies,
Milwaukee Public Museum, 800 West Wells St.,
Milwaukee, Wisconsin 53233, USA.
Journal of the Lepidopterists’ Society
55(3), 2001, 124-126
SESIDAE—CLEARWING MOTHS. HANDBOOK OF PALAE-
ARCTIC MACROLEPIDOPTERA. VOLUME 1. Karel Spa-
tenka, Oleg Gorbunoy, Zdének La&ttivka, Ivo ToSevski,
and Yutaka Arita, 1999. Watercolor illustrations by
Bohumil Stary, Ruth Holzinger, and Franti¥ek Gregor.
Genitalic illustrations by Oleg Gorbunov and Ivo
ToSevski. Clas M. Naumann, Managing Editor.
Published by Gem Publishing Company, Brightwood,
Brightwell cum Sotwell, Wallingford, Oxfordshire, UK
OX10 OQD. ISBN 0-906802-08-3. xv + 569 pp., 487
color illustrations, 504 male and female genitalic
illustrations, and 309 distribution maps; all reproduced
by off-set printing. Hardbound with color jacket. Price
£120, $170.00 (US). Shipping, £5, $7.12 (US) for first
copy, £3, $4.30 (US) for additional copies; overseas
shipping, £10 for first copy, £5 for additional copies.
Subscriptions to The Gem Publishing Company are
accepted by check or banker's draft in sterling or US
dollars. Payment may also be made directly into Giro
Account No. 467-6912.
In 1991, an editorial board for The Handbook of
Palaearctic Macrolepidoptera (HPM) was formed. As
stated in Volume 1, HPM “is not meant to cover the
entire field of the butterflies and the larger moths,”
but to “provide a basis for the treatment of those
groups where critical, reliable and experienced taxon-
omists are available and willing to devote themselves
to preparing modern revisions to their special groups.”
Moreover, “HPM will not compete with Microlepi-
doptera Palaearctica.” Given the above, and the fact
that the placement of Sesiidae within the Microlepi-
doptera is unlikely to be challenged, it is difficult for
this reviewer to ascertain how the editorial board of
HPM has determined taxonomic coverage of its publi-
cations. Clearly, there appears to be a taxonomic in-
consistency between what is being advertized (by
HPM) and what is being published. That said, I think
that volume 1 on Sesiidae is one of the more compre-
hensive and superbly illustrated works published on
any lepidopteran taxon.
This book is divided logically into several parts in
the following order: introduction; check-list of
Palaearctic Sesiidae; known host-plants; keys to the
subfamilies, tribes, and genera; a taxonomic treatment,
including Insertae sedis; bibliography; color and black-
and-white illustrations of the imagos; and indices to in-
sect names and technical terms, and plant names.
The introduction contains several concise, well writ-
ten parts that include an historical review, diagnostic
characters of the family, morphology, biology, natural
VOLUME 55, NUMBER 3
history, mimicry and behavior, distribution, economic
importance, collecting, and rearing. Although a phylo-
genetic classification of the tribes within Sesiidae is
presented (Fig. 10a), no summary of supportive char-
acters is provided.
The check-list of Palaearctic Sesiidae appears com-
plete with all species arranged to reflect some degree
of relatedness. The authors have provided all syn-
onymies. In addition, a list of host-plants is arranged
alphabetically by the species-group names. All moth
species are arranged, as within the check-list, within
their respective taxonomic categories, with all authors
and dates given. In addition, all associated host-plants
are provided with their respective family names in an
easily readable table.
Keys to the subfamilies, tribes, and genera empha-
size features of the antennae, wing venation and de-
gree of wing transparency, legs, and male and female
genitalia. The couplets are dichotomous, easy to read,
and emphasize features that are relatively easy to diag-
nose. The keys to the species are a pleasure to read be-
cause they emphasize body maculation. However, in a
group such as the Sesiidae, the authors recommend,
especially when trying to identify a “rubbed” speci-
men, that the genitalia be used to confirm the identity.
The taxonomic treatment for all species included
provides the bulk of the text. It is divided into several
sections that include a complete synonymy, diagnosis,
variation, including sexual dimorphism, male and fe-
male genitalia, bionomics, habitat, and distribution
maps. Because the male and female genitalia of the
type-species of Zhuosesia Yang, 1977 could not be ex-
amined, its subfamilial and generic placement could
not be made; and consequently it is placed Insertae
sedis.
Although this volume has several coauthors, its de-
scriptions are surprisingly uniform. However, there are
some general inconsistencies to be noted. First, rela-
tive lengths given for the apophyses posteriores and
antrum in females, are too general. Terms like “com-
paratively long” or “quite short” are not informative.
On occasion, lengths relative to some other structure
are given, e.g., “the antrum is twice the length of the
eighth segment.” It may have been better to use these
ratios or some other ratio throughout the text more
consistently. In addition, the position of the ductus
seminalis in the female is inconsistently illustrated (ei-
ther present or absent), and its position relative to the
ostium bursae and the inception of the corpus bursae
would have been valuable to include in the description
of the female genitalia. Other inconsistencies include
the description of the “thorns” on the distal part of the
aedeagus in the male and the “bifid” nature of the co-
ecum penis of the aedeagus in the male; are probably
due to reduction of the original illustration and the ori-
entation of the specimen when it was illustrated.
Several errors of omission or interpretation were
noted during a detailed review of the species descrip-
tions of Tinthiini and Sesiini. For example, in Tinthia
tineiformis (p. 38) the corpus bursae is described as
having “a small rounded signum in upper part,” while
the associated text Fig. 276 lacks a signum. The uncus
in males of Tinthia myrmosaeformis and T. hoplisif-
ormis are described as being “hooked and bifurcate,”
but these features are not illustrated in text Figs. 13
and 15, respectively. The text includes a description of
the female genitalia of Tinthia mianjangalica (p. 42),
but the text figure is omitted. The description of Para-
doxecia gravis (p. 50) indicates that the corpus bursae
“has a sclerotized field,” yet it appears only partially il-
lustrated. Moreover, the corpus bursae appears to
have a long crescent-shaped signum, which is not
mentioned in the description. Descriptions of the fe-
male genitalia for Sesia repanda and S. przewalskii er-
roneously indicate that the antrum is V-shaped. In
these cases, it is the anterior margin of the ostium that
is V-shaped.
An extensive bibliography contains 830 references,
including the most important references dealing with
the Palaearctic Sesiidae. However, many faunistic, bi-
ological, and other references were prudently ex-
cluded from an already useful bibliographic listing.
There are 489 beautifully reproduced color illustra-
tions of the imagos with several accompanied by illus-
trations of the head and its appendages and/or other
diagnostic body features. The original illustrations
were rendered in water color. The reproductions are
2x original size. Some of the illustrations of species
that have heads and other body appendages illustrated
separately could have been made larger as their small
size restricts the reader from seeing diagnostic color
patterns.
Genitalic features represented by 276 male illustra-
tions and 228 female illustrations were rendered in
black ink. One male illustration (text fig. 5) and 1 fe-
male illustration (text fig. 7) are accurately labeled to
offer the user a clear understanding of the terms given
in the descriptive text. Five male illustrations and 3 fe-
male illustrations appear to be added to accommodate
8 species that were discovered after the position of the
bulk of the illustrations of the book plates were already
“fixed.” Although, the addition of a section to accom-
modate these additional species appears awkward, it
does not detract from the work, but adds to its com-
pleteness. Easy-to-read distribution maps for all 309
species are provided.
Two indices are provided, one for insect names and
technical terms and one listing the species of host-
plants included. I found it very convenient that for
each moth species listed in the index that the primary
locator pages are in bold, the page the species is found
in the species key is indicated, and pages for all plates
and text figures are given. Additionally, a square sym-
bol precedes the page number on which each distribu-
tion map for a given species is found. In the host-plant
index, I found it very useful to have the family name
provided for each species of plant listed.
In conclusion, I find this volume a very useful con-
tribution to the taxonomic understanding of Palaearc-
tic Sesiidae and of Lepidopterology in general. If a
phylogenetic classification had been included with a
presentation and discussion of supportive characters,
the volume would have been a standard for future
works edited by any organization. The taxonomic com-
pleteness and depth, along with the fine color and
black-and-white line illustrations greatly overshadow
the few errors and inconsistencies found in the text.
Davip ADAMSKI, Systematic Entomology Labora-
tory, USDA, c/o National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560-
0168, USA (email: dadamski@sel.barc.usda.gov).
Journal of the Lepidopterists’ Society
55(3), 2001, 126-128
MARIPOSAS ARGENTINAS: GUIA PRACTICA E ILUSTRADA
PARA LA IDENTIFICACION DE LAS PRINCIPALES MARIPOSAS
DIURNAS Y NOCTURNAS DE LA PROVINCIA DE BUENOS
AIRES, by Andrés E. Varga. 2000. Published by the
author. 148 pp. Paper. ISBN: 987-43-1671-3.
Approximately US $60. Price available from the author:
museo@mariposasdelmundo.com; fax (54) (11) 4664-
2108; mail address Calle Italia 650, San Miguel 1663,
Pcia. Buenos Aires, Republica Argentina. (In Spanish.)
What do you know about the Southern Cone—the
south-temperate zone comprising Chile, Argentina
and Uruguay? (It is cone-shaped; look at a map.) You
probably have some notions of tango and gauchos and
military dictatorships, but did you know that the re-
gion embraces some of the greatest natural beauty on
the planet, an incredible array of biomes, and a unique
flora and fauna? Long better-known to Europeans, the
Southern Cone only recently has begun to attract sig-
nificant eco-tourism from the USA.
What do you know about the butterflies there? For
most people, Lepidopterists or not, South America
equals big, shimmering blue Morphos and gaudy swal-
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
lowtails. But things change beyond the Tropic of
Capricorn, and for most of us the Southern Cone is
terra incognita. The biggest problem has been the lack
of a field guide or faunistic treatment for any of the
countries. Here is a paradox. All three countries have
strong entomological traditions going back to the mid-
dle of the nineteenth century, with indigenous journals
and abundant Lepidopterological literature—but until
very recently, nothing one can actually use afield.
There are taxonomic catalogues of all three faunas.
They are antiquated, not very accessible, and typically
not illustrated. Uruguay has the least interesting fauna.
The country is small, nearly flat, and mostly agricultur-
alized. Virtually everything found there occurs in Ar-
gentina too.
The first breakthroughs to a popular field guide
occurred in Chile. Butterfly studies in Chile were
dominated for half a century by two arch-rivals, the
academic José (“Pepe”) Herrera G. and the accom-
plished amateur Luis E. (“Lucho”) Pefia G. Pefia was
better-known outside Chile: he sold specimens of the
Chilean entomofauna worldwide and collaborated in
the filming of nature documentaries for the BBC and
others. He prepared a small pocket guide to Chilean
butterflies that was distributed as a subscription pre-
mium by a newspaper in the 1970s. This was later ex-
panded into a real book, Las Mariposas de Chile (with
coauthor Alfredo J. Ugarte P.), appearing in 1997 (Ed-
itorial Universitaria, Santiago) after Pefia’s death.
Even as this work was in preparation, many new
species of lycaenids were being discovered by the Is-
raeli Dubi Benyamini and others, and described by
Kurt Johnson, Zsolt Balint, and Benyamini. These
were worked into the manuscript. Thus we have an
up-to-date and effectively complete reference work
available for Chile.
The Argentine situation is much more daunting.
The Argentine fauna is perhaps an order of magnitude
richer. Unlike Chile, Argentina extends into the humid
lowland tropics. Elements of the enormous fauna cen-
tered on Amazonia extend through the Argentine
northeast and as far south as the subtropical forests of
Tucumén. Argentina is a land of great physiographic,
climatic and ecological complexity. The lowland tropi-
cal fauna articulates with the tropical Andean and alti-
plano fauna in the dissected yungas of Salta and Jujuy.
In places one can drive from rain forest (selva) through
Andean-alpine habitats to mesquite and columnar cac-
tus desert in four hours. To the south lie zones of sub-
tropical thorn scrub (espinal), creosote-bush desert
(monte), subhumid to subarid bunchgrass prairie
(pampa), cool Patagonian desert-steppe (including
southern juniper woodland) and the cool to cold mon-
VOLUME 55, NUMBER 3
tane forests of the Patagonian Andes and southwestern
Tierra del Fuego. What a country! Unlike Chilean
Patagonia, Argentine Patagonia is relatively dry and
sunny, fostering a rich butterfly fauna. And recent
studies have revealed tremendous evolutionary dy-
namism there and along the steppe-forest ecotone, a
veritable “ferment of variability.” How to deal with
such a fauna?
In 1973 Kenneth Hayward, an Englishman long res-
ident in Argentina, published a taxonomic catalogue of
the butterflies and skippers of the entire country. It
contained hardly any biology, had no range maps or
plates, and was rife with errors, but it has had to serve
as the foundation of modern Argentine butterfly stud-
ies. Hayward was also responsible for an extraordinary
four-volume work on Argentine butterflies, in oversize
format with magnificent lithographic plates in the style
of Edwards and Scudder. Published between 1948 and
1967 under the pompous series title “Genera et
Species Animalium Argentinorum,” this rare and pon-
derous work was intended to initiate a complete zoo-
logical inventory of the country. It represents the acme
of Peronist pretension; indeed, Juan Domingo Perén
himself patronized the initial volumes. This work does
contain some biology, but it is hardly useful as a field
guide, nor is it widely or readily accessible even within
the country.
Andrés Eugenio Varga has undertaken to change
the situation for the better. Argentina is an entrepre-
neurial country, and in the Buenos Aires suburb of San
Miguel, about 30 km from the city center, Varga has
opened a Museum of Butterflies of the World display-
ing some 70,000 mounted specimens in uniquely-con-
structed concave display cases (intended for ease in
viewing) under filtered light. There is a biology lab, a
rearing facility and a library. The museum—first of its
kind in the Southern Cone—is open to guided tours
only Monday through Friday, and on a drop-in basis to
the general public on Saturdays. It has a support
group, the Friends of the Museum, and has initiated
its own journal. Meanwhile, Varga has published the
first in a projected series of four volumes intended to
cover the butterfly fauna of the entire country. Unsur-
prisingly, it focuses on the Province of Buenos Aires.
Volumes 2 through 4 will cover (in order) “Meso-
potamia” (the humid tropical northeast), the pampa
and cuyo (the desert-Andean contact zone centered on
Mendoza), and Patagonia and Tierra del Fuego. He’s
made a good, if not perfect, start.
Mariposas Argentinas is lavishly produced, in 20 x
28 cm format, with color illustrations on nearly every
page. The color reproduction is remarkably true—
even capturing the elusive near-UV purple sheen on
the upper surfaces of Colias lesbia (p. 27). There is an
introductory essay by Varga emphasizing global change
and the biodiversity crisis. This is followed by six pages
of basic butterfly biology and anatomy and a résumé of
basic taxonomy (the Linnean hierarchy, binomial
nomenclature and the Code—cladistics is not men-
tioned). The main part of the book—the species ac-
counts, about which more anon—occupies pp. 17-94.
This is followed by a very detailed 20-page section on
collecting and preserving butterflies. Some may find
this material retrograde in emphasis, or at least incon-
sistent with the author's conservationist rhetoric.
There is a 2-page overview of the Museum and its pro-
jects, a 4-page glossary (84 terms), a 7-page bibliogra-
phy (118 items), and 2 pages of maps. The bibliogra-
phy contains both Argentine and foreign references.
Of the former group three were new to me, and one of
those—in an obscure agricultural extension venue—is
not in Lamas, Robbins and Field’s Annotated Bibliog-
raphy of the Neotropical Butterflies and Skippers (Sci-
entific Publishers, Gainesville, Florida, 1995) although
it dates from 1968. Varga has published most of the
material other than species accounts previously as
freestanding articles in Argentine magazines.
The species accounts cover 171 butterflies and skip-
pers, most of which are illustrated. Many show sexual
and seasonal dimorphism, and both dorsal and ventral
surfaces. The nature of the variation is usually explicit
in the captions; an exception is the pierid Tatochila
vanvolxemti (p. 25), which has seasonal forms analo-
gous to our Pieris napi, but they are not identified as
such. In addition to the butterflies, a selection of large
and showy moths is presented. The Black Witch rep-
resents the Noctuidae. There are 14 saturniids, 13
sphingids, and miscellaneous arctiids, pericopids, la-
siocampids, ctenuchids, ete.—all common and famil-
iar. There are also several pages of photos of represen-
tative (again, showy) larvae.
The text accompanying these photos is somewhat
disappointing. Frequently it consists merely of a verbal
description of what is evident in the photo. (Since the
pictures are natural size, this is hardly necessary.)
There is little biology, and some of what is given is sus-
pect. This is especially true of host plant records,
which are given without attribution and (as Varga told
me) sometimes straight out of Hayward. Opening ran-
domly to page 72, I immediately saw the familiar fiery
Skipper, Hylephila phyleus, which is as much a lawn
and garden “weed” in Buenos Aires as in Los Angeles
or Houston. In B.A. it routinely breeds on (surprise!)
Bermuda Grass, Cynodon dactylon (“Pasto Bermuda,”
“Gramilla Blanca,” “Pato de Perdiz’) and other lawn
grasses and on Paspalum (“Pasto Miel,” “Gramilla
Dulce”) and other weedy species. Varga says: “The
caterpillars eat grasses, including Sugarcane, Achiras,
etc.” “Achira” is the Spanish name for Canna, which is
not a grass but is a skipper host. I do not know if either
Sugarcane or Canna really is a host of H. phyleus, but
they are certainly not the most important ones. On the
same page is the Argentine Least Skipper, Ancy-
loxypha nitidula. No host is specified. In my experi-
ence this species is often intimately associated with the
grass Leersia hexandra; at any rate, it should be easy to
find out what it eats. One hopes later editions and vol-
umes will rely more on information from life and less
on antique and questionable published sources.
The taxonomy appears up-to-date and was vetted by
Profs. Carlos Mielke and André Victor Lucci Freitas of
Brazil and a local Argentine specialist, Joaquin Carreras.
There are too many typographical errors. Many of
the bibliographic citations are incomplete.
Factual errors are rare, but there is one important
bit of confusion that appears on p. 17. Under Euryades
corethrus (Papilionidae) we read: “They are character-
ized by two mighty extremities (harpagones) in the
posterior part of the abdomen, which the females pos-
sess in order to hold on during copulation and prevent
new fertilizations.” This refers, of course, to the
sphragis—a “chastity belt” secreted by the male during
copulation, not an integral part of the female. The
sphragis of Euryades is indeed “mighty.” It is also flam-
boyant, bilobed, and—when fresh—bright green (!).
What are “harpagones?” In the glossary this Spanish
term is given as a synonym of harpes, which are cor-
rectly defined as “periphallic moveable processes . . .
which usually function in subjugating the female dur-
ing copulation.” The lobes of the sphragis no tienen
nada que ver con harpagones.
This book is a vast improvement over what was
available before. It is not cheap. Books have always
been expensive in the Southern Cone, despite the fact
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
that Buenos Aires shares with Mexico City the tradi-
tional intellectual leadership of Latin America. Mari-
posas Argentinas, the set, will however be indispens-
able to any serious student of the butterflies of the
region: the later volumes all the more so. The fauna of
the Province of Buenos Aires is probably the least in-
teresting of the lot, but it has its biogeographical
charms. It is mostly derived from the lowland
Neotropics. A number of such taxa enter the Province
only in the riverine and swamp forests of the Tigre
Delta; from the capital south the Neotropical element
is very dilute. Endemic austral-Patagonian elements
enter in the grasslands. The admixture is well illus-
trated by the Satyridae. Two radiations are repre-
sented. The euptychiines are lowland-Neotropical.
The pronophilines have radiated twice in South
America, with one large radiation in the tropical Andes
(often associated with bamboos) and the other in the
temperate grasslands, extending into alpine steppe and
grassland. It is the latter group that reaches its north-
ern limit in the littoral in the Province of Buenos Aires.
One species, Etcheverrius (=Cosmosatyrus) chilensis,
extends from there to far-southern Patagonia, and also
occurs widely in Chile. Except for some of the skip-
pers, the Buenos Aires fauna is fairly well-known. The
same cannot be said for the fauna of any other region.
Varga’s emphasis on collecting is justified in places
where one can still collect three new species in one
day (as I once did in Patagonia). He asks on p. 6: “How
can we claim to conserve that which we still don’t
know?” A good question, and this book may stimulate
others to get out and learn the Argentine fauna while
it is still in place. Otherwise, he says, we will all too
soon “lament, as we remember this beautiful world
that we had but couldn’t take care of.”
ARTHUR M. SHAPIRO, Center for Population Biology,
University of California, Davis, California 95616, USA.
Date of Issue (Vol. 55, No. 3): 26 February 2002
EDITORIAL STAFF OF THE JOURNAL
Caria M, Penz, Editor
Department of Inyertebrate Zoology
Milwaukee Public Museum
Milwaukee, Wisconsin 53233, USA
flea@mpm.edu
Pui. DeVries, Book Review Editor
. Genter for Biodiversity Studies
Milwaukee Public Museum
Milwaukee, Wisconsin 53233, USA
pjd@mpm.edu
Associate Editors:
~~ Gerarpo Lamas (Peru), Keneum W. Pure (USA), Rosert K. Rospins (USA),
Feux Sperune (Canada), Davin L. Wacner (USA), Curister WikLuND (Sweden)
NOTICE TO iy eee peas eakeuie
: ce Lena Fo Feature ‘Photoeraphs and Chea Illustrations are ao vet in Volume 44(2): 1 and on the ue s web
re at ttp:/Avww.furman.edu/~snyder/snyder/lep/.
ise SE d. rnal submissions to the editor at the above address (or Aectomculy to: flea@mpm.edu). Contributors should feel free to
‘commend one or two reviewers upon submission of their manuscript. Send requests to review books for the Journal and book re-
oe u ripts to ug DeVries at ‘the above address (or omy to: pid@mpm.edu). Short “manuscripts seighad new state
Seirinioally to: philjs@mail.utexas.edu), Before eubmitting pauiaabiiple or electronic files, consult dastructionsin vol:
47-50 or the Society's web site at http://www.furman.edu/~snyder/snyder/lep/.
nuscripts in cp lteate, si ante entirely alae es with wide eas on one side only vA white letter- sized ig
or ‘Maeintosh: based word processing format:
: Include an OE TEE abstract for Articles, Profiles, and Technical Comments.
Up to five key words or terms not in the title should accompany Articles, Profiles, General Notes, and
ok with precision, clarity and economy, sad use the active voice and the first person whenever appropriate. Make title
it, de lescriptive, and as short as possible. The first mention of a plant or animal in the text should include the full scientific name
or, and family. Use metric units for measurements, and a 24-h clock (0930 h, not 9:30 AM) to express time. Underline only
ics are aptended.
ot MED ee Illustrate only half of s Aeuea objects such as adults sith wings spread, unless ole acta is ee
y y i gs Sp
Mount: ee ae and eens on stiff, white backing, areas in the peered format. Bear in mind os Your sie pees will be
seh abel age: 16. 5 em width, 22.5 cm hei sht). Illustrations larger than ia are not acce tie and ae be reduced
‘page P ge § gS P
ph to ee to that size or smaller. On the back of each illustration, print ees NaS name and a numbers as s.cited 1 in the
‘or oun Hae Golor Statens are cao contact Lane for has Peni eaionie bed aot
‘Tables: Number tables consecutively in Arabic numerals, Label them in small caps (e.g., Taste 1.) and use a concise and infor-
heading. ‘Type each table on a separate sheet and place after the Literature Cited section, with the approximate desired po-
ition indicated i in the text. Avoid vertical lines and vertical writing,
Voucher specimens: When appropriate, manuscripts must name a public repository where specimens documenting the iden-
f organisms ¢ can be found. Kinds of reports that require vouchering include descriptions of new taxa, life histories, host associ-
roofs: The edited manuscript : and galley proofs will be mailed:to the author for correction of printer's errors, Excessive author's
= will be charged to authors at the rate of $3.00 per line. A purchase order for reprints will accompany proofs.
a age charges: For authors affiliated with institutions, page charges are $50 per Journal page. For setae ago institutional
page. ‘Authors roable: to pay page charges for any reason should apply to the editor at the time of sobiitsian bi a reditted rate or
ho) free: ea Authors of Book oR se and Obituaries are sty oe from page pera
PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A.
CONTENTS
A NEW SPECIES OF EucosMoMORPHA FROM Nortu America (TortricipaE) William E. Miller ----- 81
AN OVERVIEW OF StrYMOon HtsNer (LyCAENIDAE: THECLINAE: EumAEini) Robert K. Robbins and.
Stanley S. Nicolay ----------------------------------------------------2 2m nnn nnn 85 a
LepmorrerIsTs’ PERCEPTIONS OF A PROPOSED PERMITTING SYSTEM FOR BUTTERFLY COLLECTING ON PUBLIC
LANDS Kristine C. Mazzei and Arthur M. Shapiro ------------------------=----------------—--- 101
LIFE HISTORY AND LABORATORY HOST RANGE TESTS OF PARAPOYNX SEMINEALIS (WALKER) (CRAMBIDAE:
NyMPHULINAE) IN FLortwa, U.S.A. Gary R. Buckingham and Christine A. Bennett ----- a is ha
GENERAL NOTES
NYCTEOLA FRIGIDANA WALKER (NOCTUIDAE: SARROTHRIPINAE) REPORTED AT AN UNORTHODOX BAIT
Timothy L. McCabe ----------------------------------- 2 =n =n nanan nnn nnn Glo tee
HEPIALUS CALIFORNICUS (HEPIALIDAE) OVIPOSITION PREFERENCE ON THE LUPINE LUPINUS ARBOREUS ret
TQTE PP FCUSSO Ro arr re a ee ee ee 119
Status OF PIERIS VIRGINIENSIS (PrERTDAE) IN New York state Edward J. Stanton ------------ 122: i
Book Reviews
FLYING Jewers—Butrerriy Imaces. P. J. DeVeies (124 ee
SESIDAE—CLEARWING Motus. Hanppoox oF PaLakarctic MACROLEPIDOPTERA. VoLuME i
David Adamski. -------------------------------------------=~ ---nn nnn nn nein nn nnn ------ 124
Mariposas ARGENTINAS: GUiA PRACTICA E ILUSTRADA PARA LA IDENTIFICACION DE LAS PRINCIPALES
MARIPOSAS DIURNAS Y NOCTURNAS DE LA PROVINCIA DE Buenos Aires Arthur M. Shapiro -- 126
This paper meets the requiremenis of ANSI/NISO Z39.48-1992 (Permanance of Paper).
Number 4
ISSN 0024-0966
eieds SocreTy
- Published quarterly by’ THE LEPIDOPTERISTS SOCIETY
Publi ei LA SOCIETE DES LEPIDOPTERISTES
“Piblieado por ta SOCIEDAD DE Los LEPIDOPTEROLOGOS
THE LEPIDOPTERISTS’ SOCIETY
EXECUTIVE CoUNCIL
Joun W. Brown, President Susan S. Borin, Vice President
Micnaet J. Smitu, Immediate Past President Mirna M. Casacranpe, Vice President —
Manuen A. Batcazar-Lara, Vice President \ Davin’ C: IrrNer, Treasurer
Ernest H, Winuiams, Secretary
Members at large:
Ronald L, Rutowski M. Deane Bowers George L. Balogh
Felix A. H. Sperling Ron Leuschner Andrew V. Z: Brower
Andrew D. Warren Michael Toliver Brian Scholtens ©
E,piroriaL. BOARD
Rosert K. Rossins (Chairman), Joun W. Brown (Member at large).
Caria M. Penz ( Jowrnal)
WinuiaMm EF. Miniter (Memoirs)
Puituir J; Scuarrert (News)
Honorary Lire MEMBERS OF THE SOCIETY
Cuarues L, Remincton (1966), E. G. Munroe (1973), Ian F. B: Common (1987),
Joun G. Francremont (1988), Linco.n P. Brower (1990), Douctas C. Fercuson (1990),
Hon. Miriam Roruscuip (1991), Craupe Lemaire (1992), Freperick H. Rinpcr ened
The object of The Lepidopterists’ Society, which was formed in May 1947 and formally constituted in December 1950, is “to ae -
mote the science of lepidopterology in all its branches, . . . to issue a periodical and other publications on Lepidoptera, to: facilitate
the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all mea
sures’ directed towards these aims.
Membership in the Society is open to all persons interested in the study of loolaonlenk All members receive the Journal and the
News of The Lepidopterists’ Society. Prospective members should send to the Assistant Treasurer full dues for the current year, to-
gether with their full name, address, and special lepidopterological interests. In alternate years.a list of members of the Bec hs is asa
sued, with addresses and special interests.
Active members——annual dues $45.00 within the U.S., $50.00 outside the U.S. —
Affiliated members—annual dues $10:00 within the U.S., $15.00 outside the U.S. pereass 0
Student members—annual dues $20.00 within the U.S., $25.00 outside the U.S. zg ins ager
Sustaining members—annual dues $60.00 within the U.S., $65.00 outside the U.S, ae
Life members—single sum $1,800.00
Institutional subscriptions—annual $60.00 se
Airmail postage for the News $15.00 aie fear
Send remittances, ches to The Lepidopterists’ Society, to: Kelly M. Richers, Asst. Treasurer, 9417 Carvalho Court, Bakersfield, Bh:
CA 93311; and address changes to: Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057, —
For information about the Society, contact: Ernest H. Williams, Department of Biology, Hamilton College. Clinton, NY 13323, To.
order back issues of the Memoirs, write for availability and prices to Kelly M. Richers, Asst. Treasurer, 9417 Carvalho Court, Bakers- >
field, CA 93311.
The additional cost for members outside the U.S. is to cover mailing costs.
Journal of The eh ese Society (ISSN 0024-0966) is published quarterly by The Lepidopterists’ sanieae Yo Los sages oy
County Museum of Natural History, 900 Exposition Blvd:, Los Angeles, CA 90007-4057. Periodicals postage paid at Los Angeles, CA
and at additional mailing offices. POSTMASTER: Send address changes to The ee Society, > Natural oe Museum,
900 Exposition Blvd., Los Angeles, CA 90007-4057.
Cover illustration: Rhetus arcius caterpillar (Riodinidae), by artist Jennifer Clark. Jn: DeVries, P. J. 1997. The butterflies of Costa Rica
and their natural history, Volume II: Riodinidae. Princeton Univ. Press. é
JOURNAL OF
Toe LeriporTterRistTs’ SOCIETY
Volume 55 2001 Number 4
Journal of the Lepidopterists’ Society
55(4), 2001, 129-139
REVIEW OF REBINEA RAZOWSKI AND ELIACHNA RAZOWSKI (TORTRICIDAE: EULIINI)—
SISTER GROUPS ENDEMIC TO CHILE AND ARGENTINA
JOHN W. BROWN
Systematic Entomology Laboratory, Plant Sciences Institute, Agricultural Research Service, U.S. Department of Agriculture,
c/o National Museum of Natural History, Washington, DC 20560-0168, USA
e-mail: jbrown@sel.barc.usda.gov
AND
TsiTs1 Y. MCPHERSON
c/o Faculty of Natural Sciences or Faculty of Arts, University of Guyana, Turkeyen Campus,
Greater Georgetown, GUYANA
ABSTRACT. Rebinea Razowski and Eliachna Razowski, two formerly monotypic genera known only from males, are redescribed based on
large series of specimens (n = 320) including both sexes. As presently defined, Rebinea is monotypic, with a single variable species, R. erebina
(Butler, 1883), and its synonym, Arotrophora balsamodes Meyrick, 1931. It is possible that two (or more) species are concealed within the vari-
ation, but we were unable to separate them using traditional morphological characters. Eliachna is represented by three species: E. chileana Ra-
zowski, 1999, E. digitana Brown and McPherson, new species, and E. hemicordata Brown and McPherson, new species. Both genera
are restricted to south-central Chile and southwestern Argentina, ranging from coastal lowlands (5 m) to middle elevations (1200-1700 m) at
the southern end of the Andes. A phylogenetic analysis of the four species (plus two out-group species) provides support for the sister relation-
ship of Rebinea and Eliachna based on the following synapomorphies: (1) elongate labial palpi (length 34 times horizontal diameter of the com-
pound eye); (2) a pair of stout, digitate, submedial processes on the dorsum of the transtilla; (3) a deep, rounded excavation near the mid-venter
of the valva; and (4) a pair of semicircular, lateral flanges from the posterior edge of the sterigma.
Additional key words: new species, genitalia, phylogenetics, leafrollers.
The tortricid fauna of Chile and Andean Argentina Rebinea and Eliachna based on new information, de-
is distinct from that of the remainder of South scribe two new species of Eliachna, provide data on
America, comprised primarily of endemic, or nearly the geographic distribution of the included species,
endemic genera (e.g., Accuminulia Brown, Acman- and examine the phylogenetic relationship between
thina Brown, Argentulia Brown, Chapoania Razowski, the two genera.
Chileulia Powell, Eliachna Razowski, Haemateulia Ra-
zowski, Nesochoris Clarke, Proeulia Clarke, Rebinea MATERIALS AND METHODS
Razowski, Recintonia Razowski, Varifula Razowski). We examined 320 pinned adults of Rebinea and Eli-
Although the contributions of Razowski (1995, 1999) achna deposited in the following institutions: Ameri-
and Brown (1998, 2000a, b) have added substantially can Museum of Natural History, New York, New York,
to our knowledge of Euliini of this region, phyloge- U.S.A. (AMNH); The Natural History Museum, Lon-
netic relationships among genera of the tribe are unre- don, England (BMNH); Mississippi Entomological
solved and many species remain undescribed. Prior to Museum, Mississippi State, Mississippi, U.S.A.
this study, Rebinea and Eliachna were considered (MEM); Essig Museum of Entomology, University of
monotypic, known only from a handful of males. The California, Berkeley, California, U.S.A. (UCB); Na-
discovery of additional specimens of the two genera, tional Museum of Natural History, Smithsonian Insti-
including both sexes, reveals a close phylogenetic rela- tution, Washington, D.C., U.S.A. (USNM):; and Zoo-
tionship between them and the presence of two new logical Museum, Copenhagen, Denmark (ZMC).
species. The purposes of this paper are to redescribe Specimens were sorted by forewing pattern, geo-
130
graphic location, and sex. The resulting groups then
were examined for differences in male and female
genitalia, which have been shown to provide the most
reliable morphological features for distinguishing
among related species of Tortricidae. Preparation of
genitalia slides followed the methodology summarized
in Brown and Powell (1991). Because of the pheno-
typic similarity of, and variation within the treated
species, we examined the genitalia of all male speci-
mens. For undissected specimens, we used a fine
camel-hair brush to remove scales from the external
margin of one valva, which provided enough detail to
convincingly assign all males to a species-level taxon.
Specimens were examined using a Wild M3Z dissect-
ing microscope; slide mounted genitalia were studied
using the dissecting microscope and a Zeiss compound
microscope. Illustrations of genitalia were drawn with
the aid of a Ken-A-Vision microprojector (model
X1000-1). Unless indicated otherwise, genitalia illus-
trations are of a single preparation. Text descriptions of
all characters are composite, based on all available
specimens. Measurements of forewing and labial palpi
were made with an ocular micrometer mounted in a
Wild M3Z dissecting microscope under low power
(x10-16). Forewing length was measured in a straight
line from the base to the apex of the wing, including
the fringe. Forewing width was measured at the widest
place perpendicular to the length measurement.
Where available, a minimum of 15 individuals of each
sex were measured. Colors follow Ridgway (1912); ter-
minology for wing venation and genitalia structures
follows Horak (1984). Abbreviations and symbols are
as follows: DC = discal cell; ca. = circa (approxi-
mately); n = number of individuals examined; x =
mean; N, E, S, W = compass points.
Polymorphism or moderate variation in phenotype
is uncommon to rare in most Euliini. However, many
euliine species in Chile and Argentina (e.g., Chileulia,
Proeulia, Haemateulia), including the two genera
treated here, are variable in forewing pattern and mac-
ulation. Consequently, examination of the genitalia is
the only reliable method for accurately identifying
species. Comparison with the illustrations provided is
highly recommended. For males, the profile of a single
valva is adequate; females must be dissected.
SYSTEMATICS
Rebinea Razowski, 1986
Rebinea Razowski, 1986:22; Powell et al., 1995:145;
Razowski, 1999:84.
Type species. Sericoris erebina Butler, 1883:72, by
original designation.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Diagnosis. Adults of Rebinea are superficially and
morphologically most similar to those of Eliachna
among described Euliini genera. The two share a sim-
ilar forewing shape, size, and pattern; extremely elon-
gate labial palpi (3-4 times horizontal diameter of the
compound eye); a pair of stout, digitate, submedial
processes from the dorsum of the transtilla; a deep,
rounded excavation near the mid-venter of the valva;
and a pair of semicircular, lateral flanges at the poste-
rior edge of the sterigma. Superficially, males of many
Rebinea can be distinguished from Eliachna by their
slightly longer forewings and paler ground color. Gen-
italia characters that distinguish Rebinea from Eli-
achna include its broad, straight aedeagus, with a
single, large, compound conutus; the absence of an
elongate, free process at the distal end of the sacculus;
and an extremely short, broad ductus bursae.
Redescription. Head: Antennal cilia approximately 1.5 times
width of flagellomere in male; cilia short, unmodified in female.
Labial palpus porrect, 3-4 times horizontal diameter of eye in both
sexes (i.e., without dimorphism). Vertex with overhanging tuft of
scales. Proboscis present, presumably functional. Ocellus moder-
ately large. Chaetosema present. Thorax: Smooth scaled. Male with-
out foreleg hairpencil. Forewing (Figs. 5, 9): Length 2.3-2.6 times
width; length of DC ca. 0.6 times forewing length; width of DC ca.
0.2 times DC length; CuA, originates 0.6-0.7 along DC length; all
veins separate beyond DC; chorda and M-stem absent; CuP weak,
present only at margin. No upraised scale tufts; male without costal
fold. Hindwing: Sc + R and Rs closely approximate at base; Rs and
M, stalked; M, and CuA, connate or short-stalked; CuP present; M-
stem absent; tuft of hairlike scales at base of 1A + 2A in both sexes.
Abdomen: Smooth scaled; dorsal pits absent; no modified corethrog-
yne scaling in female. Male genitalia (Fig. 10): Uncus short, moder-
ately stout, curved, strongly sclerotized; socius moderately short,
hairy, drooping, slightly expanded distally; gnathos arms slender,
with a delicate terminal plate; transtilla a transverse band with a pair
of stout, digitate, submedial processes on dorsum, sometimes with
tips slightly expanded; valva broad at base, with deep, rounded exca-
vation near mid-venter; sacculus restricted to basal portion of valva,
strongly sclerotized, ending in short, dorsal-projecting hook; pulvi-
nus absent; vinculum well developed, strongly sclerotized; juxta
large, stout arrowhead-shaped. Aedeagus large, straight, with a
single large, compound, capitate cornutus, consisting of several
fused filaments; vesica finely spiculate. Female genitalia (Fig. 14):
Apophyses anteriores and posteriores moderate in length, slender.
Sterigma relatively broad, with narrow, shallowly U-shaped, sclero-
tized band and a pair of semicircular lateral flanges from posterior
edge. Ductus bursae extremely short, broad, with a short membra- _
nous region immediately anterad of antrum; a frail, obovate, acces-
sory bursa from a moderately long ductus originating from dorsum
of corpus bursae in posterior one-third; corpus bursae moderately
large, ovoid, with dense spiculae and a large, irregularly rounded,
sclerotized plate, usually along lateral wall.
Distribution and biology. Rebinea is known from
Chile and Argentina between about 30° and 45°S lati-
tude, ranging from the coastal lowlands (50 m) to the
middle elevations (1400 m) of the southern Andes
(Fig. 1). Collecting localities seem to have little in
common in terms of habitat type, ranging from
VOLUME 55, NUMBER 4
Nothofagus forest (e.g., Alto Tregualemu) to xeric ar-
eas dominated by succulents and leguminous trees
(e.g., Nague, Los Vilos) (see Davis 1986 for descrip-
tions of the habitat at many of the collecting localities
listed below). Adults have been captured primarily in
October (n = 16), November (n = 118), and December
(n = 95), with a few specimens recorded from January
through March. The early stages are unknown.
Remarks. Razowski (1986) included two species in
Rebinea: R. erebina (Butler) and R. balsamodes
(Meyrick). Powell et al. (1995) synonymized the two
without explanation. Although the types of the two
nominal taxa are fairly distinct in forewing size and fa-
cies, they have extremely similar genitalia. The abun-
dance of material now available from Chile and Ar-
gentina suggests that the two probably represent forms
of an extremely variable species. Alternatively, there
may be more than one species concealed within this
variation (see discussion below).
Rebinea erebina (Butler, 1883)
(Figs. 1,5, 9, 10, 14)
Sericoris erebina Butler, 1883:72.
Arotrophora balsamodes Meyrick, 1931:381; Clarke,
1963:8 [illustration of adult and male genitalia].
Rebinea balsamodes: Razowski, 1986:22.
Rebinea erebina: Razowski, 1986:22 [illustration of
male genitalia]; Powell et al., 1995:145; Razowski,
1999:84.
Diagnosis. Rebinea erebina resembles Eliachna
species in general facies; most individuals (especially
females) cannot be separated by forewing length and
pattern. The male genitalia of R. erebina can be distin-
guished easily from those of Eliachna by its broad,
straight aedeagus with a large compound cornutus,
and the absence of the elongate, free process of the
sacculus. In addition, the overall shape of the valva is
distinct, with a short hooklike process at the terminal
end of the sacculus; the latter likely represents an au-
tapomorphy for Rebinea. The female genitalia of R.
erebina are similar to those of Proeulia, Argentulia,
and other related Chilean-Argentinean genera, with an
extremely broad ductus bursa and an irregularly scler-
otized, highly spiculate corpus bursa.
Redescription. Male. Head: Lower frons pale tan to cream; up-
per frons gray brown. Labial palpus light brown. Thorax: Mostly
brown. Forewing (Fig. 5): Length 6.6-8.8 mm (x = 7.9 mm; n = 25);
ground color pale gray to burnt umber, with small, scattered, indis-
tinct patches of orange-brown, orange-red, and cream scales; fre-
quently with faint, parallel bands in distal one-third of wing repre-
sented by series of slightly disjunct, darker scales; often with a
variably developed, diagonal fascia of darker brown from near mid-
costa to dorsum, angled outward near middle of wing. Hindwing:
131
White to pale brown, usually with brownish-gray mottling. Ab-
domen: Pale yellow brown to dark brown. Genitalia (Fig. 10): As de-
scribed for genus (drawn from USNM slide 90383; 15 preparations
examined), Female. Head, thorax, and abdomen: Essentially as de-
scribed for male. Forewing: Length 6.2-8.2 mm (xX = 7.4; n = 15);
ground color burnt umber to cinnamon, with indistinct patches of
orange-brown, orange-red, and cream scales; a variably defined, red-
brown fascia from near mid-costa to dorsum, angled outward near
middle of wing. Genitalia (Fig. 14): As described for genus (drawn
from USNM slide 81239; 10 preparations examined).
Types. Holotype ¢ (erebina), Chile, [Mountains of the hacienda
of Cauquenes, Butler 1883] (BMNH). Lectotype (new designation)
3 (balsamodes), Argentina, Territory Rio Negro, Lake Gutierrez,
3-14.X1.1926 (F. & M. Edwards, BMNH).
Material examined. ARGENTINA: Chubut Province: El Bols6n,
Lago Puelo, 220 m, 2d, 17.X1.1978, 1 d, 2 9, 21.X1.1978 (Mision Ci-
entifica Danesa, ZMC), 1 2°, 22.X.81, 3 9, 23.X.1981 (Nielsen &
Karsholt, ZMC); Esquel, Lago Menéndez, E] Sagrario Puerto, 600 m,
14, 19, 2-4.1.1982 (Nielsen & Karsholt, ZMC). Neuquén Province:
Lago Lacar, Pucaré, 750 m, 4 9, 10.X1.1978, 2 d, 2 2, 25.X1.1978, 5d,
2.2, 1.XII.1978, 3d, 7 9, 2.X11.1978, 5 5, 3 2, 3.X11.1978 (Mision Cien-
tifica Danesa, ZMC); Lago Lacar, Pucaré, 600 m, 21 4, 3 9,
28-29 .X1.1981, 1d, 1 9, 26-27.XII.1981 (Nielsen & Karsholt, ZMC);
Lago Lacar, 5 km E Hua-Hum, 640 m, 2 4, 3 9, 6.X1.1981, 2d, 1 9,
25.X1.1981, 1 2, 26-27.XI1.1981 (Nielsen & Karsholt, ZMC); San
Martin de los Andes, 640 m, 1 d, 14.X.1981, 1d, 1 9, 17-31.X.1981, 4
3, 16.X.1981, 3 4, 2.X1.1981, 1 2, 5-6.X1.1981, 3d, 6 2, 7-15.X1.1981,
1 2, 26.X1.1981 (Nielsen & Karsholt, ZMC); San Martin de los An-
des, Cerro Chapelco, 1400-1600 m, 2 d, 2-19.XII.1981 (Nielsen &
Karsholt, ZMC). Rio Negro Province: San Carlos de Bariloche,
Colonia Suiza, 810 m, 1 2, 6.XI.1978, 1 d, 15.XI.1978, 3 2°,
28.X1.1978, 3 d, 29.X1.1978, 1 9, 4.X1I.1978, 1 2, 5.XII.1978, 3 2,
11.XJ1.1978, 1 d, 12.X11.1978, 1d, 15.XII.1978, 1 2, 9.1.1979 (Mision
Cientifica Danesa, ZMC); San Carlos de Bariloche, Colonia Suiza,
800 m, 1 6, 26.X.1981, 1 6, 31.X.1981, 1 6, 11.X1.1981, 2 d, 1 9,
12-20.X1.1981, 4 3, 21-22.X1.1981, 1 9, 23.X1.1981, 1 d, 24.X1.1981,
14,22, 29-30.X1.1981, 2 6, 3 9, 3.XII.1981, 2 35, 1 9, 5—6.XII.1981, 1
3, 7.X1I.1981, 1 d, 8.X1I.1981, 1 2, 22.X11.1981, 1 2, 5-7.1.1982
(Nielsen & Karsholt, ZMC); San Carlos de Bariloche, Camino de
Tronador, 2 9, 29.X1.1978 (Mision Cientifica Danesa, ZMC); Lago
Nahuel Huapi, Puerto Blest, 770 m, 1 4, 27.XI.1978, 1 9,
18.XII.1978, 1 3, 30.XII.1978 (Mision Cientifica Danesa, ZMC), 1 d,
3.X11.1981 (Nielsen & Karsholt, ZMC). CHILE: Aconcagua
Province: Los Andes, Curimon, 700 m, 1 4, 28.III.1979 (Mision Ci-
entifica Danesa, ZMC). Chiloé Province: Puntra, ca. 30 air km S An-
cud, 50 m, 1 4, 1 9, 21-22.X11.1981 (D. Davis, USNM). Coquimbo
Province: Fray Jorge National Park, ca. 70 km W Ovalle, 4 ¢, 2 9,
6-9.X1.1981 (D. & M. Davis, USNM); Nague, 11 km N Los Vilos, 1
é, 4-5.XL.1981 (D. & M. Davis, USNM); Coquimbo, 1 9°,
1.VII-19.X.1883, “Walker” (BMNH). Llanquihue Province: Casa
Pangue, | 2 (paralectotype of balsamodes), 4—10.X11.1926 (F. & M.
Edwards, USNM); Peulla, 1 2 (paralectotype of balsamodes),
12-13.X11.1926 (F. & M. Edwards, BMNH). Maule Province: Paso
Garcia, ca. 23 km NW Cauquenes, 300 m, 1 2, 29-30.X1.1981 (D. R.
Davis, USNM): Rio Teno, ca. 40 km E Curico, 800 m, 1 4, 1 9,
25-27.X1.1981 (D. R. Davis, USNM). Nuble Province: Alto
Tregualemu, ca. 20 km SE Chovellen, 500 m, 5 2, 1-3.X11.1981 (D.
R. Davis, USNM). Osorno Province: P. N. Puyehue, Ag. Calientes to
3 km W, 600 m, 2 d, 12-20.XI1I.1981 (D. R. Davis, USNM); Parque
Nacional Puyehue, Aguas Calientes, 450 m, 6 d, 1 9, 12.X1.1981, 1 ¢,
22, 13.X1.1981, 6 6,5 2, 10.X11.1981, 4d, 1 2, 11.XIL.1981, 1 d, 6 2,
12.XJ1.1981, 2 2, 13.X1I.1981 (Nielsen & Karsholt, ZMC); Parque
Nacional Puyehue, Anticura, 350 m, 2 4, 2 °, 17.X1.1981, 2 d, 2 9,
132 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
z R. erebina
30} f SS
|
|
40
|
|
50}
“| E. hemicordata ’ | E. chileana
fc : |
PL eee teed
|
40
50
a: 60
Fics. 1-4. Geographic distribution of Rebinea and Eliachna. 1, R. erebina; 2, E. digitana, new species; 3, E. hemicordata, new species; 4,
E. chileana.
oa
4
ge a ee pm
oak
VOLUME 55, NUMBER 4
133
Fics. 5-8. Adults of Rebinea and Eliachna. 5, R. erebina; 6, E. chileana; 7, E. hemicordata, new species; 8, E. digitana, new species.
18.X1.1981, 2 2, 19.XT.1981, 1 2, 15.XII.1981, 3 2, 17.XII.1981, 1 9,
18.X11.1981 (Nielsen & Karsholt, ZMC). Santiago Province: Los
Maitenes, Colorado River, 33°22’S, 70°17’W, 1200-1400 m, 2 4,
16.X.1954 (L. Pefia, USNM); Pilay, Rio Pueco, ca. 45 km S Santiago,
800 m, 1 d, 23-24.X1.1981 (D. R. Davis, USNM). Valdivia Province: 20
km S Valdivia, Rincon de la Piedra, 180 m, 2 °, 14.X1.1981, 1 4, 3 2,
15.X1.1981 (Nielsen & Karsholt, ZMC). Valparaiso Province: Val-
paraiso, | 3, 30.IX— 8.X.1883, “Walker 3074” (BMNH). Unknown
Province: Central Austral, 2 3, I-III.1898 (V. Izquerdo, USNM).
Discussion. Rebinea erebina is either a single,
highly variable species or two (or more) extremely sim-
ilar species that cannot be separated reliably using tra-
ditional morphological characters. At one extreme are
specimens with a large forewing length (7.3-8.8 mm),
a pale (gray to beige) forewing ground color, a poorly
defined forewing pattern, and a pale hindwing, usually
mottled with gray brown (similar to the type of ere-
bina). Although a majority of the specimens of this
phenotype are males, a few females approach this gen-
eral aspect. At the other extreme are specimens with a
shorter forewing length (6.6-7.7 mm), a darker (red
brown to brown) forewing ground color, and a more
uniformly dark hindwing (similar to the type of bal-
samodes). Although this phenotype is typical of fe-
males, some males approach this aspect (see Clarke
1963). Male genitalia are only slightly variable among
all the specimens examined (n > 100), and the varia-
tion is concordant with neither differences in facies,
forewing length, nor geographic distribution. In some
male specimens, the distal end of the venter of the
valva is somewhat pointed and slightly reflexed, while
in others it is somewhat rounded. The width of the
paired processes from the transtilla is somewhat vari-
134
able, as is the development of the hooklike process at
the distal end of the sacculus. Apparent variation in
the latter feature, however, is likely an artifact of slide
mounting of genitalia.
Remarks. In his description of Arotrophora bal-
samodes, Meyrick (1931) indicated that he had six ex-
amples from “Argentina, Territory Rio Negro, Lake
Gutierrez, November; S. Chile, Llanquihue Province,
Casa Pangue and Peulla, December.” Three of these
specimens are in the BMNH, one of which is a female
of Eliachna; one specimen is in USNM. Clarke (1963)
identified the male from Argentina as “type” without
formally designating it as the lectotype. Because the
type series consists of more than one species, we for-
mally designate a lectotype, and we select the speci-
men labeled as such in the BMNH and identified as
such by Clarke (1963). This designation is necessary to
establish the concept of the species and promote
nomenclatural stability.
Eliachna Razowski, 1999
Eliachna Razowski 1999:87.
Type species. Eliachna chileana Razowski 1999:87, by
monotypy.
Diagnosis. Eliachna is most similar to Rebinea in
forewing length, shape, pattern, and venation (see di-
agnosis of Rebinea above for details), and most speci-
mens are difficult to distinguished superficially from
Rebinea. Genitalic differences between the two genera
are conspicuous and are detailed above under Rebinea.
Redescription. Head: Antennal cilia approximately 1.5 times
width of flagellomere in male; cilia short, unmodified in female.
Labial palpus elongate, porrect, length 3-4 times horizontal diame-
ter of compound eye, slightly longer in female. Vertex with over-
hanging tuft of scales. Proboscis present, presumably functional.
Ocellus moderately large. Chaetosema present. Thorax: Smooth
scaled. Legs unmodified, male without foreleg hairpencil. Forewing
(Figs. 6-8): Length ca. 2.4 times width; length of DC ca. 0.6 times
forewing length; width of DC ca. 0.2 times DC length; CuA2 origi-
nates 0.6-0.7 along DC length; all veins separate beyond DC;
chorda and M-stem absent; CuP weak, present only at margin. No
upraised scale tufts; male without costal fold. Hindwing: Sc + R and
Rs closely approximate at base; Rs and M, stalked ca. one-third dis-
tance from DC to margin; M, and CuA, connate or short-stalked;
CuP present; M-stem absent; tuft of hairlike scales at base of 1A +
2A in both sexes. Abdomen: Dorsal pits absent; no modified coreth-
rogyne scaling in females. Male genitalia (Figs. 11-13): Uncus slen-
der, short, simple, strongly sclerotized; socius moderately short,
broad, hairy, slightly expanded distally; gnathos with slender lateral
arms connected to terminal plate by membrane. Transtilla a simple
band, highly sclerotized laterally, weaker medially, with a pair of sub-
medial, digitate processes, slightly rounded apically. Valva broad
basally, with variable excavation near mid-venter; sacculus well de-
fined, with free, elongate-digitate terminal process of variable shape
and length; pulvinus absent; juxta strongly sclerotized, stout arrow-
head-shaped. Aedeagus somewhat elongate, variably curved near
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
middle, usually with distal spine projecting dorsoposteriorly; vesica
with or without minute spinules and/or small patch of tiny cornuti.
Female genitalia (Figs. 15-16): Papillae anales somewhat slipper-
shaped. Apophyses anteriores and posteriores slender, nearly equal
in length. Sterigma usually crescent-shaped, with a pair of rounded,
sclerotized lateral flanges at posterior edge. Ductus bursae moder-
ately short; corpus bursae ovoid, with dense spicules, at least in pos-
terior one-half.
Distribution and biology. Eliachna apparently is
confined to south-central Chile and adjacent Ar-
gentina, ranging from coastal lowlands (5 m) to mon-
tane areas (1700 m). Adults have been collected from
October to April. Nothing is known of the early stages.
Eliachna digitana Brown & McPherson,
new species
(Figs. 2; 8; 11, 15)
Diagnosis. Superficially, E. digitana is difficult to
distinguish from other species in the genus; the
forewing length is usually a little shorter and the
ground color slightly more orange brown rather than
gray brown. The male genitalia can be distinguished
easily from its congeners by the subrectangular distal
portion of the valva; the short, straight, digitate process
at the termination of the sacculus; and the weakly
curved aedeagus. Female genitalia, likewise, are easily
distinguished; the lateral pouches of the sterigma and
the sclerotized, knoblike diverticula of the ductus bur-
sae are unique to this species.
Description. Male. Head: Labial palpus dark brown. Forewing
(Fig. 8): Length 5.9-7.0 mm (x = 6.5; n = 15); ground color and
maculation somewhat variable; ground color usually gold gray to red
brown, with faint, dark brown reticulations, infrequently with ill-de-
fined, darker area in basal one-third; a variably developed median
fascia from costa to dorsal margin, brown to red brown, angled out-
ward near middle of forewing; apex frequently with darker patch.
Hindwing: Brownish gray, infrequently with faint mottling. Ab-
domen: Gold brown to dark brown. Genitalia: As in Fig. 11 (drawn
from USNM slide 90484; 7 preparations examined). Uncus, socius,
gnathos, and transtilla as described for the genus. Valva long, sub-
rectangular, only slightly narrowed near middle and slightly broad-
ened distally; sacculus broad basally, with a free, mostly straight, dig-
itate process distally. Juxta as described for the genus. Aedeagus
weakly undulate; spine at termination of coecum strongly sclero-
tized, slightly disjunct from coecum; vesica densely punctate, espe-
cially in distal three-fourths; a row of minute cornuti near the base.
Female. Essentially as described for male. Forewing length 6.5-7.6
mm (xX = 6.9; n = 15). Genitalia: As in Fig. 15 (drawn from USNM
slide 81228; 6 preparations examined). Sterigma subrectangular,
with a pair of shallow lateral pouches at anterior edge. Ductus bur-
sae short, with a short, stout, knoblike, sclerotized diverticula dor-
sally; corpus bursae oblong, uniformly covered with fine spinules.
Type. Holotype ¢, Chile, Nuble Province, 17.5 km S$ Curanipe,
near coastal stream, 50 m, 25.1.1979 (D. & M. Davis & B. Aker-
bergs, USNM).
Paratypes. ARGENTINA: Chubut Province: El Bolsén, Lago
Puelo, 220 m, 1 d, 18.X1.1978 (Mision Cientifica Danesa, ZMC), | 2,
13.X.1981 (Nielsen & Karsholt, ZMC); Esquel, 550 m, 1 d, 1.1.1982
(Nielsen & Karsholt, ZMC); Sierra Colorada, 800 m, | d, 29.1.1983
VOLUME 55, NUMBER 4
(M. & P. Gentili, USNM). Neuquén Province: San Martin de los An-
des, 640 m, 1 2, 13.X.1981, 1 d, 7-15.X1.1981 (Nielsen & Karsholt,
ZMC); Lago Lucar, Pacara, 650 m, 1 d, 10.X1.1978 (Mision Cientf-
fica Danesa, ZMC), 1 6, 26-27.X11.1981 (Nielsen & Karsholt,
ZMC). Rio Negro Province: Lago Gutierrez, 1 ° (paralectotype of
balsmodes), 3-14.X1.1926 (F. & M. Edwards, BMNH); San Carlos
de Bariloche, Colonia Suiza, 810 m, 1 4, 2.XI1.1981, 1 9,
20.X11.1981, 1 6, 23.X11.1981, 1 3, 5-7.1.1982 (Nielsen & Karsholt,
ZMC), 1 2, 9.XI.1978, 1 6, 19.XI.1978, 1 6, 29.XI.1978, 1 9,
9.XI11.1978, 1 2, 12.XII.1978, 1 9, 9.1.1979, 1 2, 10.1.1979, 1 2,
11.1.1979 (Mision Cientifica Danesa, ZMC). CHILE: Cautin
Province: Fundo Neltume, 2 km N Villarrica, 200 m, 1 4, 1 9,
27.11.1979 (D. & M. Davis & B. Akerbergs, USNM); Fundo el
Coigue, 27 km NE Villarrica, 500 m, 2 4, 28.1— 3.111.1979 (D. & M.
Davis & B. Akerbergs, USNM). Malleco Province: Rio Manzanares,
5 2, 19.X.1979 (Flint & Barria, USNM). Nuble Province: Forel Car-
rizalillo, 250 m, 1 2, 30.1-5.1I.1981 (L. E. Pefia, USNM); Alto
Tregualemu, 500 m, 1 2, 27-28.1.1981 (L. E. Pefia, USNM); Alto
Tregualemu, ca. 20 km SE Chovellen, 500 m, 2 ¢, 26-27.1.1979 (D.
&. M. Davis & B. Akerbergs, UCB), 4 2, 1-3.XII.1981 (D. Davis,
USNM); 17.5 km S Curanipe, near coastal stream, 50 m, 3 d, 2 9,
25.1.1979 (D. & M. Davis & B. Akerbergs, USNM); Pierdra de la
Iglesia, 8 km N Cobquecura, 5 m, 4 9, 25.1.1979 (D. & M. Davis &
B. Akerbergs, USNM), 1 d, 4 9, 24.1.1979 (D. & M. Davis & B.
Akerbergs, USNM); Cachapoal, Cajon de Lisboa, Alhue, 800 m, 1 d,
19-21.XII.1987 (L. E. Pefia, USNM). Llanquihué Province: Llan-
quihué, Petrohue, | 4, 8.111.959, 1 d, 12.111.959 (J. F. G. Clarke,
USNM). Osorno Province: Parque Nacional Puheyue, Anticura, 250
m, 1 d, 17.XII.1981 (Nielsen & Karsholt, ZMC). Santiago Province:
Rinconada Maipt, 450 m, 35°31’S, 70°47’W, 1 4, 14.1V.1966 (W.
Hichins & M. E. Irwin, UCB); Pilay, Rio Peuco, ca. 45 km S$ Santi-
ago, 800 m, 1 3, 23-24.XJ7.1981 (D. Davis, USNM). Valdivia
Province: Valdivia, 1 °, 7.111.1960 (E. Krahmer, ZMC); 20 km S$ Val-
divia, Rincon de Piedra, 180 m, 1 d, 24.X1.1981 (Nielsen & Karsholt,
ZMC).
Distribution and biology. Eliachna digitana oc-
curs from Santiago Province, Chile, to Chubut
Province, Argentina (Fig. 2), ranging from coastal
Nothofagus forests (5 m) to arid uplands (1300 m)
dominated by Fabaceae and Lauraceae. Capture
records are from October (n = 3), November (n = 5),
December (n = 11), January (n = 20), February (n =
4), March (n = 3), and April (n = 1). Nothing is known
of the early stages.
Etymology. The species name refers to the digitate
process that comprises the distal portion of the sacculus.
Remarks. N. Obraztsov probably was the first to
recognize this species as distinct and undescribed; he
labeled the specimen from Llanquihué, Peulla
(USNM) with a manuscript name.
Eliachna hemicordata Brown & McPherson,
new species
(Figs. 7, 13)
Diagnosis. Eliachna hemicordata has a_slighter
greater forewing length and a paler ground color than
other species in the genus. Males can be distinguished
from other Eliachna by the somewhat cordate distal
135
Fics. 9-10. Rebinea erebina. 9, Wing venation; 10, Male geni-
talia, aedeagus removed, valvae spread.
portion of the valva; the elongate, curved, digitate, free
process of the sacculus is similar to that of E. chileana.
The female is unknown.
Description: Male. Head: Labial palpus dark brown. Thorax:
Brown to dark brown. Forewing (Fig. 7): Length 7.8-8.0 mm (x =
7.9; n = 5); ground color pale orange cream, with tiny black specks
throughout; moderately broad median fascia, extending from costa
ca. 0.6-0.7 from base to apex, to dorsum ca. 0.7—-0.8 from base to
tornus, angled outward near middle of forewing Hindwing: Pale
gray brown with variably developed darker mottling. Abdomen:
Gold brown to dark brown. Genitalia: As in Fig. 13 (drawn from
USNM slide 81222; 7 preparations examined). Uncus, socius,
gnathos, transtilla as described for genus. Valva long, distal portion
ovoid, with short hooklike process from venter of apex; sacculus with
elongate, slightly flattened, weakly curved free process. Juxta as de-
scribed for genus. Aedeagus curved dorsad just beyond coecum,
with dorsoposteriorly projecting spine at distal end; vesica without
spicules. Female. Unknown.
136
Zl
BE
AS
EFL:
aT he
Fics. 11-13. Male genitalia of Eliachna, aedeagus removed, val-
vae spread. 11, E. digitana, new species; 12, E. chileana, new
species; 13, E. hemicordata.
Type. Holotype 4, Argentina, Nuequén, Chapelco Lenga, 1700
m, 24.1.1984 (M. & P. Gentili, USNM).
Paratypes. ARGENTINA: Neuquén Province: Chapelco Techos,
1400 m, 1 d, 21.1.1982 (M. & P. Gentili, USNM); Lago Lacar,
Trompul, 1200 m, 1d, 6.1.1983 (M. & P. Gentili, USNM); San Martin
de los Andes, Tr. Kura, 1000 m, 1 d, 29.X1J.1985 (M. & P. Gentili,
USNM). Rio Negro Province: Lago Nahuel Huapi, Puerto Blest, 1 4,
23.XII. 1978 (Mision Cientifica Danesa, ZMC). CHILE: Bio-Bio
Province: Lago El Barco, Guallali, Sta. Barbara, 1200 m, 1 4,
25-28.11.1981 (L. E. Petia, USNM). Cautin Province: [Parque Na-
cional] Conguillio, 1200 m, 1 d, 48.11.1988 (L. E. Pefia, USNM).
Unknown Province: V. Villarica, 16 km S Pucon, | 4, 20.XI1.1982 (R.
Brown, MEM).
Distribution and biology. Eliachna hemicordata
is known from Neuquén and Rio Negro provinces, Ar-
gentina, and Bio-Bio and Cautin provinces, Chile, be-
tween 1000 and 1400 m (Fig. 3). Capture records are
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics, 14-16. Female genitalia of Rebinea and Eliachna. 14, Re-
binea erebina; 15, E. digitana, new species; 16, E. chileana.
from January (n = 3), February (n = 2), and December
(n = 3). Nothing is known of the early stages.
Etymology. The species name refers to the half-
hearted shape of the distal portion of the valva.
Eliachna chileana Razowski, 1999
(Figs. 6, 12, 16)
Eliachna chileana Razowski, 1999:88 [male genitalia
illustrated].
Diagnosis. The male genitalia of E. chileana can be
distinguished from those of other species in the genus
by the greatly expanded distal portion of the valva, ter-
minating in an attenuate, pointed tip. The female gen-
italia can be distinguish by the simple U-shaped
sterigma.
Redescription. Male. Head: Labial palpus light brown. Thorax:
Mostly brown. Forewing (Fig. 6): Length 7.1-7.8 mm (x = 7.5 mm; n
= 4); ground color dull silvery gray, faintly overscaled with red orange
and copper orange; basal one-fourth usually with patch of slightly
darker scales; variably defined, red-brown median fascia from near
mid-costa to dorsum, angled outward near middle of forewing; termi-
nal area with irregular patches of black and orange-red scales, de-
creasing toward apex. Hindwing: Pale olive brown, with faint brown-
VOLUME 55, NUMBER 4
Fic. 17. Hypothesis of phylogenetic relationship among the
taxa. Numbers on the left refer to characters (1-15); numbers on the
right refer to character states (see Table 1).
ish-gray reticulations. Abdomen: Pale yellow brown to dark brown.
Genitalia: As in Fig. 12 (drawn from USNM slide 81223; 5 prepa-
rations examined). Uncus, socius, and gnathos as described for genus.
Valva broad basally, narrowed at middle, greatly expanded distally,
with elongate, curved, beaklike process directed ventrally; sacculus
with elongate, free, slightly flattened, curved, digitate process. Juxta as
described for genus. Aedeagus strongly curved near middle; vesica
with several minute spines and a few short, capitate cormuti. Female.
Essentially as described for male. Forewing: Length 6.7—7.0 mm (x =
6.9; n = 4). Genitalia: As in Fig. 16 (drawn from USNM slides 90493
and 90177; 5 preparations examined). Sterigma weakly U-shaped, uni-
form in thickness. Ductus bursae moderately short, with large ductus
seminalis originating dorsally about midway between ostium and cor-
pus bursae. Corpus bursae as described for the genus.
Type. Holotype 4, Chile, Nuble Province, Las Trancas, Shangri-
la, Chillan area, SE Recinto, 1500 m, 15.XII.1983 (L. Pefia,
AMNBH).
Additional specimens examined. CHILE: Malleco Province:
nr. Los Gringos Camp, Nahuelbuta Nat. Park, 1300 m, 3 d, 2 9,
6-11.1.1982 (D. R. Davis, USNM). Nuble Province: Shangri-la, SW
side Volcan Chillan, 1600 m, 1 3, 19-21.1.1979 (D. &. M. Davis & B.
Akerbergs, USNM); Las Trancas, 21 km E Recinto, near high wa-
terfall, 1300 m, 14,32, 17.1.1979 (D. &. M. Davis & B. Akerbergs,
USNM).
Si
Distribution and biology. Eliachna chileana is
known only from Malleco and Nuble provinces (Fig.
4). With all captures between 1300-1600 m, this
species appears to be restricted to higher elevations
than its congeners. Capture records are from Decem-
ber (n = 1) and January (n = 10). Nothing is known of
the early stages.
EXPLANATION OF CHARACTERS AND
PHYLOGENETIC ANALYSIS
A phylogenetic analysis was performed on the four
species that comprise Rebinea and Eliachna, plus two
out-group species, Proeulia triquetra Obraztsov and
Haemateulia haematitis (Meyrick). The use of Proeu-
lia and Haemateulia as out-groups is somewhat arbi-
trary because sister group relationships within this
clade of Euliini previously have not been demon-
strated. However, the two genera share a variety of
features with Rebinea and Eliachna (e.g., forewing
pattern, polymorphism, broad, short valvae, etc.), and
all appear to belong to a complex of endemic Chilean-
Argentinean genera that is taxonomically isolated
from other Euliini present in South America. The pri-
mary purposes of the analysis were to confirm the sis-
ter relationship of Rebinea and Eliachna, and ensure
that the two in-group genera are monophyletic with
respect to each other. The analysis was based on 15
morphological characters (11 binary and 4 multistate),
including two of the head, one of the thorax, nine of
the male genitalia, and three of the female genitalia.
Character state polarity was determined through the
out-group method and using Horak’s (1984) assess-
ment of characters of Tortricinae. The character state
data were subjected to parsimony analysis using the
“mhennig*” command of Hennig86 (Lipscomb 1994).
Characters used in the analysis are listed and discussed
briefly below; the character matrix is present in Table 1.
1. Labial palpi: (0) upturned, ca. 1.5—-2.0 as long as
horizontal diameter of the compound eye; (1) some-
what porrect, ca. 2.0-3.0 as long as horizontal diame-
ter of the compound eye; (2) porrect, 3-4 times the
horizontal diameter of the compound eye. While elon-
gate labial palpi occur in several groups scattered
throughout Euliini (e.g., Proeulia, Seticosta Razowski,
Anopinella Powell, etc.), none of these taxa have palpi
as long as those of Rebinea and Eliachna.
2. Male antennal cilia: (0) conspicuous, elongate, ca.
1.0-1.5 times the width of the flagellomere; (1) incon-
spicuous, extremely short, ca. 0.5 times the width of
the flagellomere. The value of this character is dimin-
ished somewhat by its variability in the out-group taxa
(e.g., length of male antennal cilia varies among
species of Proeulia).
138
TABLE 1. Character matrix (“P” = missing data).
haematitis 00100 00001 11020
triquetra 11000 00100 00001
erebina 20111 00110 00101
digitana 20111 10211 11111
chileana 20111 01312 11111
hemicordata 20111 00312 11???
3. Male foreleg hairpencil: (0) present; (1) absent.
The presence of a male foreleg hairpencil is assumed to
represent the plesiomorphic condition in Euliini
(Brown 1990). However, because the structure is evo-
lutionarily labile, there is no evidence that its shared
absence is truly a synapomorphy for the taxa that lack it.
4. Transtilla: (0) a simple bridge; (1) with a pair of
stout, digitate, submedial processes on dorsum.
Although digitate structures are present on the dorsum
of the transtilla of Inape Razowski and Ortognathosia
Razowski (see Razowski 1988 for illustrations), few
other features of the male or female genitalia of the
latter two genera indicate a close relationship with Re-
binea and Eliachna. Hence it is suspected that the
structures represent convergent development in In-
ape, Ortognathosia, and Rebinea + Eliachna.
5. Valva: (0) venter uniform; (1) venter with a deeply
excavated portion near middle resulting in a broad
basal portion, a narrow “neck” near the middle, and an
expanded distal portion. The distinctive shape of the
valva is apparently unique to Rebinea and Eliachna,
and is reminiscent of the valva of some Eucosmiini
(Olethreutinae).
6. Valva: (0) distal one-third somewhat club-shaped;
(1) distal portion narrowed, somewhat elongate-
rectangular. The latter character state is considered an
autapomorphy for E. digitana.
7. Valva: (0) distal one-third somewhat club-shaped;
(1) distal portion greatly expanded into an elongate,
curved, beaklike process directed ventrally. The latter
character state is considered an autapomorphy for E.
chileana.
8. Sacculus: (0) weak, lacking free terminal
process; (1) well-defined, with short, free, distal ter-
mination; (2) well-defined, with slender, digitate
process; (3) well-defined, with long, slightly flattened,
curved process. A sacculus lacking a free terminal
process, such as that in the genitalia of Haemateulia,
is considered the plesiomorphic condition. A well-de-
fined sacculus with a short, free, distal termination,
such as that in the genitalia of Proeulia, is considered
derived; and the development of the free tip into an
elongate, digitate process is considered a synapomor-
phy for Eliachna. Its further development into an ex-
tremely long, slightly flattened, curved process is
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
considered a synapomorphy for E. chileana and E.
hemicordata.
9. Juxta: (0) shield-shaped, unmodifed; (1) stout ar-
rowhead-shaped. The presence of a stout, arrowhead-
shaped juxta is not particularly compelling as a synapo-
morphy for Eliachna and Rebinea because other
genera of Chilean-Argentinean Euliini may possess a
similarly shaped juxta. The structure frequently is not
included in illustrations of male genitalia or is dis-
torted by slide mounting.
10. Aedeagus: (0) broad, straight, relatively large; (1)
slightly more slender, weakly curved; (2) conspicuously
more slender, strongly curved. A broad, straight,
relatively large aedeagus, characteristic of Proeulia and
Rebinea, is considered the plesiomorphic condition. A
slightly more slender, weakly curved aedeagus is con-
sidered the first step in a transformation series lead-
ing to a conspicuously more slender, strongly curved
aedeagus.
11. Aedeagus: (0) without external projections; (1)
with a small dorsoposteriorly projecting spine from
near junction of the coecum and the phallobase. The
latter character state appears to represent a synapo-
morphy for Eliachna.
12. Vesica: (0) with one or few large capitate cornuti;
(1) with numerous tiny non-capitate cornuti. The pres-
ence of one or few large cornuti, assumed to represent
the plesiomorphic condition, is typical of Proeulia; the
vesica of Rebinea, likewise, has a single, large, com-
pound cornutus.
13. Sterigma: (0) unmodified; (1) with a pair of
semicircular sclertotized flanges located at the poste-
rior edge. The latter character state appears to repre-
sent a synapomorphy for Rebinea and Eliachna.
14. Ductus bursae: (0) extremely broad, nearly as
broad as the corpus bursae; (1) slightly more narrow,
weakly differentiated from corpus bursae; (2) rela-
tively narrow, clearly differentiated from corpus
bursae. In Proeulia, Argentulia, and related Chilean-
Argentinean genera, the ductus bursae is extremely
broad. This condition, considered the plesiomorphic
state, also is present in Rebinea. The slightly more nar-
row ductus bursae of Eliachna is considered a synapo-
morphy for this genus, and the relatively well defined
ductus bursae of Haemateulia is considered the most
advanced state. Because this character varies through-
out Euliini (sometimes even within a single genus), it
is a less compelling indicator of relationship.
15. Corpus bursae: (0) finely punctate; (1) densely
spiculate. A densely spiculate corpus bursae is shared
by Proeulia, Rebinea, Eliachna, Argentulia, and addi-
tional related Chilean-Argentinean genera. The finely
punctate corpus bursa of Haemateulia, similar to most
— =
VOLUME 55, NUMBER 4
Euliini, is considered the plesiomorphic state for this
character.
The parsimony analysis of the 15 characters de-
scribed above resulted in one most parsimonious tree
with a length of 24, a consistency index of 0.83, and a
retention index of 0.69. The consistency and retention
indices are biased by the fact that the data set includes
characters that are non-informative in a phylogenetic
context (i.e., autapomorphies and characters consis-
tent within the in-group). The cladogram (Fig. 17)
shows strong support for the monophyly of Rebinea +
Eliachna on the basis of the following characters: labial
palpi extremely elongate and porrect (character state
1.2); male foreleg hairpencil absent (character state
3.1); transtilla with a pair of stout, digitate, submedial
processes on the dorsum (character state 4.1); venter
of valva with a deeply excavated mesal portion (char-
acter state 5.1); juxta stout and arrowhead-shaped
(character state 9.1); and sterigma with a pair of semi-
circular sclertotized flanges located at the posterior
edge (character state 13.1). The most convincing of
these are characters 1, 4,5, and 13.
The monophyly of Eliachna is supported by the
presence of a digitate process at the termination of the
sacculus (character state 8.2); a slender, weakly curved
aedeagus (character state 10.1); a dorsoposteriorly pro-
jecting distal spine from near the junction of the coe-
cum and the phallobase (character state 11.1); numer-
ous tiny cornuti in the vesica of the aedeagus
(character state 12.1); and a slightly more narrow duc-
tus bursae (character state 13.1). The most convincing
of these are characters 8, 11, and 12.
While the sister relationship of Eliachna and Re-
binea is well supported in the context of the genera
used in this analysis, their relationship to other euliine
genera endemic to Chile and Argentina remains un-
known. Fortunately, the unique leafroller fauna of this
region is receiving greater attention from systematists
and pest managers worldwide, as many of the native
species have broadened their host ranges to include
economically important crops (Brown 2000b). An an-
ticipated increase in specimens and host information
will undoubtedly shed additional light on phylogenetic
relationships among these genera.
ACKNOWLEDGMENTS
We thank Richard Brown (MEM), Ole Karsholt (ZMC), Jerry
Powell (UCB), Eric Quinter (AMNM), and Kevin Tuck (BMNH)
for allowing us to examine material in their care. Marianne Horak
(CSIRO, Canberra, Australia) arranged for the ZMC specimens to
be examined by us. We thank David Adamski (USDA Systematic
Entomology Laboratory, Washington, D.C.) for preparing slides of
genitalia and wings, for rendering the drawings of wing venation
and genitalia, and for arranging for photographs of the adults.
139
Donald Davis (USNM) provided assistance in mapping collecting
localities. The following provided reviews that enhanced the clar-
ity and quality of the paper: David Smith (USDA Systematic Ento-
mology Laboratory, Washington, D.C.), Ronald Ochoa (USDA
Systematic Entomology Laboratory, Beltsville, Maryland), Josef
Razowski (Polish Academy of Sciences, Krakow, Poland): William
Miller (Department of Entomology, University of Minnesota, St.
Paul, Minnesota): and Carla Penz (Milwaukee Public Museum,
Milwaukee, Wisconsin). The taxonomic portion of this study was
funded in part by a Smithsonian Women’s Committee grant to T.
McPherson in the context of the Smithsonian Research Training
Program (2000).
LITERATURE CITED
Brown, J. W. 1990. Taxonomic distribution and phylogenetic sig-
nificance of the male foreleg hairpencil in the Tortricinae (Lep-
idoptera: Tortricidae). Entomol. News 101:109-116.
. 1998. A new genus of tortricid moths from Chile and Ar-
gentina related to Varifula Razowski (Lepidoptera: Tortricidae).
J. Lepid. Soc. 52:177-181.
. 2000a. A new genus of tortricid moths (Tortricidae: Euli-
ini) injurious to grapes and stone fruits in Chile. J. Lepid. Soc.
53:60-64.
2000b. Acmanthina: a new genus of tortricid moths (Lep-
idoptera: Tortricidae) from Chile and Argentina. J. New York
Entomol. Soc. 108:106—113.
Brown, J. W. & J. A. POWELL. 1991. Systematics of the Chrysoxena
group of genera (Lepidoptera: Tortricidae: Euliini). Univ. Calif.
Publ. Entomol. 111. 87 pp. + figs.
BuTLER, A. G. 1883. Heterocerous Lepidoptera collected in Chili
by Thomas Edmonds, Esq. Trans. Entomol. Soc. London
1883:49-90.
CLARKE, J. F. G. 1963. Catalogue of the type specimens of mi-
crolepidoptera in the British Museum (Natural History) de-
scribed by Edward Meyrick. Vol. 4. Published by the Trustees
of the British Museum, London. 521 pp.
Davis, D. R. 1986. A new family of monotrysian moths from austral
South America (Lepidoptera: Palaephatidae), with a phylo-
genetic review of the Monotrysia. Smithsonian Contrib. Zool.
434.
Horak, M. 1984. Assessment of taxonomically significant struc-
tures in Tortricinae (Lep., Tortricidae). Bull. Soc. Entomol. Suisse
57:3-64.
Liescoms, D. 1994. Cladistic Analysis Using Hennig86, version
1.5. Documentation for Hennig86 computer program. George
Washington University, Washington, D.C. 122 pp.
Meyrick, E. 1931. Micro-Lepidoptera from south Chile and Ar-
gentina. An. Mus. Nac. Hist. Nat., Buenos Aires 36:377-415.
PowELL, J. A., J. Razowski & J. W. Brown. 1995. Tortricidae:
Tortricinae, Chlidanotinae, pp. 138-151. In J. B. Heppner (ed.),
Atlas of Neotropical Lepidoptera, Checklist Part II:
Hyblaeoidea—Pyraloidea—Tortricoidea. Association for Tropi-
cal Lepidoptera, Scientific Publishers, Gainesville, Florida.
RazowskI, J. 1986. Description of new neotropical genera of
Archipini and rectification of the Deltinea problem (Lepi-
doptera: Tortricidae). Sci. Nat Bull. 52:21—25.
. 1988. New genera and species of Neotropical Archipini
(Lepidoptera, Tortricidae). Acta Zool. Cracoy. 31:387-422.
. 1995. Proeulia Clarke, 1962, the western neotropical Tor-
tricidae genus (Lepidoptera), with descriptions of five new
species and two allied genera. Acta Zool. Cracov. 38:271-283.
. 1999. Euliini (Lepidoptera: Tortricidae) of Chile. Polskie
Pismo Entomol. 68:69-90.
Ripeway, R. 1912. Color standards and color nomenclature. Pub-
lished by the author, Washington, D.C. 41 pp. + 53 plates.
Received for publication 31 July 2001; revised and accepted 24 Oc-
tober 2001.
Journal of the Lepidopterists’ Society
55(4), 2001, 140-143
THE LIFE HISTORY OF DASYPYGA ALTERNOSQUAMELLA RAGONOT (PYRALIDAE) FEEDING ON
THE SOUTHWESTERN DWARF MISTLETOE (ARCEUTHOBIUM VAGINATUM) IN COLORADO
KAILEN A. MOONEY
Department of EPO Biology, University of Colorado, Boulder, Boulder, Colorado 80309-0334, USA
kailen.mooney@colorado.edu
ABSTRACT. The immature stages, feeding and oviposition behaviors, patterns of larval abundance, and associated arthropod fauna of
Dasypyga alternosquamella Ragonot (Pyralidae) on Arceuthobium vaginatum susp. cryptopodum (Hawks.), the Southwestern dwarf mistletoe,
are described and illustrated. The study was conducted at the Manitou Experimental Forest, U.S.D.A. Rocky Mountain Research Station,
Woodland Park, Colorado, where the Southwestern dwarf mistletoe parasitizes Pinus ponderosa (Laws.) scopulorum.
Additional key words: _ biological control, herbivory, Phycitinae, Promylea lunigerella, Blue Hairstreak.
Dwarf mistletoes have significant economic and
ecological impacts on coniferous species throughout
the West and have been called “the single most de-
structive pathogen of commercially valuable conifer-
ous timber trees in several regions of Mexico, western
Canada, western United States, and parts of Asia”
(Hawksworth & Wiens 1996). Past work has catalogued
the lepidopteran and other arthropod fauna associated
with dwarf mistletoes (Stevens & Hawksworth 1970),
but there is little or no natural- or life-history informa-
tion is available for many of these species. Dasypyga al-
ternosquamella Ragonot (Pyralidae, Phycitinae), a
common herbivore of dwarf mistletoes can have sig-
nificant effects on dwarf mistletoe standing biomass
(e.g., Reich 1992), and could be an important agent of
biological control for this important conifer parasite.
Dasypyga alternosquamella feed on multiple
species of dwarf mistletoes (Arceuthobium spp. [Vis-
caceae]) throughout western North America (Heinrich
1920). This species was described by Ragonot (1887)
from specimens collected in California. A description
of larva and pupa were first given by Heinrich in 1920,
and relevant information was again summarized by
Heinrich in 1956. This is the first published work to
provide basic natural- and life-history information on
this species. In this paper I describe the immature
stages of D. alternosquamella, larval feeding and
oviposition behaviors, and a brief account of other
dwarf mistletoe-associated arthropods at a field site on
the eastern slope of the Colorado Rockies.
MATERIALS AND METHODS
This work was conducted at the Manitou Experi-
mental Forest, an administrative unit of the U.S. De-
partment of Agriculture Forest Service Rocky Moun-
tain Experiment Station located in Woodland Park,
Colorado. The field portion of the work was in a stand
of pure 50 to 60 year old ponderosa pines (Pinus pon-
derosa var. scopulorum Laws.) growing at an elevation
of 2414 m (39°06’40’N, 105°06”50’W). These trees
are heavily parasitized by Southwestern dwarf mistle-
toe (A. vaginatum subsp. cryptopodum Hawks.).
All dwarf mistletoes (Arceuthobium spp.) are leaf-
less, have highly reduced flowers, and are dioecious.
Plants of A. v. cryptopodum (Fig. 1) reach a maximum
height of approximately 20 cm and a single plant con-
sists of multiple shoots emerging directly from the
bark of host pine trunks and branches. Individual
shoots range from 2—5 mm in diameter. Coloration is
uniform within plants, but highly variable among
plants including yellows, pale greens, and browns, of-
ten with reddish tints.
Southwestern Dwarf Mistletoe plants were col-
lected from the field between 30 June and 1 August
1999 in individual plastic bags and brought into the lab
on eight separate occasions. Individual plants ranged
from 3 to 10 cm in height and in most cases only one
or two plants were taken from any single host-pine.
Both eggs and early instar larva were isolated from
these plants using a dissecting microscope, as well as
other arthropods associated with dwarf mistletoe.
Because dwarf mistletoes are not free-living and
only grow on conifers, D. alternosquamella were not
reared on living host plants, but instead they were
Fic. 1. The Southwestern dwarf mistletoe parasitizing ponderosa
pine. Scale bar 10 cm.
VOLUME 55, NUMBER 4
14]
Fic. 2. Dasypyga alternosquamella adult. Scale bar 1 cm.
reared individually in clear plastic petri dishes lined
with filter paper in a laboratory facility. The larvae
were fed small (2-5 cm) shoots of dwarf mistletoe col-
lected from the same general location as the larvae
themselves, and they were replenished with fresh
plant material approximately every third day. The filter
paper linings of each petri dish were wetted on a daily
basis. The lab building was neither heated nor cooled,
and petri dishes were stored in the open near a win-
dow where they received indirect but not direct sun-
light. Although every precaution was taken to maintain
a “natural” rearing environment within the laboratory
facility the quality of harvested food plants, as well as
other environmental variables, likely differ to some ex-
tent from that of a living plant.
Larval head capsule widths and resting body lengths
were measured using a stereomicroscope with an ocu-
lar micrometer. Head capsule widths were taken daily
while resting body lengths were taken only at the time
of molting. Five larvae were reared from eggs through
pupation, three larvae were collected at second instar
and reared through pupation, and one additional larva
was collected at the third instar and reared through
pupation. All pupae were measured and weighed ap-
proximately two months after pupation. Voucher spec-
imens of adults reared for this work are housed at the
University of Colorado Museum in Boulder, Colorado.
To determine egg hatch-time 25 field-collected eggs
were reared at least through first instar. To document
pupation behavior several late-instar larvae were
reared on dwarf mistletoe plants still attached to
clipped pine branches in a terrarium with several cen-
timeters of soil and needles in the bottom. The pine
branch, dwarf mistletoe, and soil were subsequently
searched for pupae. Larval feeding behaviors both in
the laboratory and in the field were recorded.
Fic. 3. Dasypyga alternosquamella egg on Southwestern dwarf
mistletoe. Scale bar 1 mm.
RESULTS
Oviposition. Eggs and first instar larvae appeared
in the field beginning 30 June 1999 and the last eggs
were found on 3 August 1999. As the egg stage lasts
approximately seven days (see below), adult emer-
gence likely began sometime in the middle of June. In
two years of fieldwork at this site, I have seen an adult
(Fig. 2) only once in the field during the day and have
not observed oviposition and other adult behaviors.
These behaviors may be occurring nocturnally. Eggs
were laid singly, although the frequent presence of
multiple pees of differ ent instars on a single plant
suggests that ovipositing females may not make any ef-
fort to avoid plants on which previous oviposition has
occurred.
Eggs. Eggs are circular, approximately 0.5 mm in diameter, and
slightly domed in shape (Fig. 3
slight white mottling, ail along the margins, and adhere
tightly to the surface of the dwarf mistletoe shoots, thus becoming
readily visible in the field. Eggs were never found on the pine foliage
The dates of oviposition of the 25 eggs collected from
the field are not known, but all hatched within eight days. Specifi-
cally, one hatched eight days after collection, six hatched seven days
after collection, mal the other 18 hatched in less than seven days,
). They are light red in color with
or branches.
suggesting a maximum egg stage of seven to eight days. Without
knowing the actual dates of egg laying minimum egg stage duration
can not be estimated.
Larvae. Dasypyga alternosquamella has six instars. Heinrich
(1920) provides a formal description and illustration of larval char-
acters. The size (head capsule width and larval length) and instar du-
ration of each stage are shown in Table 1. The average duration from
egg hatch to pupation was 47 days (N = 7, SE = 0.76) over which
time larvae grew from a mean length of 1.19 mm (N = 5, SE =
0.048) at hatching to 16.56 mm (N = 9, SE =
(Table 1).
First instar larvae feed on the plant surfaces including terminal
1.034) at pupation
shoots and flowers, presumably because they are unable to penetrate
the harder exterior surface of the dwarf mistletoe shoots. Later instars
(Fig. 4) frequently mine shoots, often entering the shoot at the base
and moving distally. Large aggregations of frass can accumulate at the
142
TABLE 1.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Mean values for head capsule width, pre- and post-molt body lengths of resting larvae, and instar duration for Dasypyga alter-
nosquamella. Sample sizes (N) given in column two, standard errors (SE) follow each measurement. Post-molt body length for instar one is size
at time of hatching.
x head capsule
X post-molt body
X instar duration
X pre-molt body
Instar N width mm (SE) length mm (SE) length mm (SE) days (SE)
1 5 0.15 (0.005) 1.19 (0.048) 1.61 (0.093) 7.33 (0.558)
2 8 0.20 (0.004) 1.62 (0.093) 2.30 (0.088) 6.5 (0.563)
3 9 0.29 (0.010) 2.31 (0.088) 3.25 (0.124) 6.38 (0.263)
4 9 0.43 (0.012) 3.26 (0.124) 5.36 (0.288) 6.33 (0.471)
5 9 0.64 (0.011) 5.37 (0.288) 8.25 (0.310) 7.11 (0.351)
6 9 0.96 (0.111) 8.26 (0.310) 16.56 (1.034) 14.78 (0.760)
entry holes to the shoots, and is not typically found within shoots. Be-
cause D. alternosquamella begins feeding at shoot bases, even small
amounts of feeding result in the death of the entire shoot.
Whether or not the larvae move among separate dwarf mistletoe
plants is unclear. However, when disturbed, larvae of all instars will
either drop from the plant on a line of silk or depart and travel
across pine branches. There are typically many dwarf mistletoe
plants on a single tree at this study site, making larval movement
among plants feasible in at least some instances. Frequently a larva
which has dropped will then ascend the same line of silk to return to
its original location.
Heinrich (1920:84) notes that “the color of the individual larvae
varies in harmony with the color of the individual batches of mistle-
toe on which they feed.” The coloration of larvae is variable, and
generally this range of variation is similar to some host plant colors,
but no close association between larval color and host plant color
was observed. Furthermore, larval coloration did not change as a
function of color of the dwarf mistletoe plants on which they were
fed in the laboratory.
Pupae. Dasypyga alternosquamella has a single generation per
year and over-winter as pupae. Sixth instar larvae drop to the ground
and in the litter and soil they construct a small chamber of frass, soil
and silk where they pupate. This chamber is approximately 10 mm
in length and 4 mm in width and just encloses the pupae with little
excess room. Pupae (Fig. 5) average 9.1 mm in length (N = 8, SE =
0.34) and 2.8 mm in width (N = 8, SE = 0.11). Two months after pu-
pation average pupal weight was 35.4 mg (N = 7, SE = 1.94).
Larval abundance. During the course of this study
284 larvae were collected on 112 plants for an overall av-
erage of 2.5 larvae per plant. Most of these larvae were
used in an unrelated field experiment. Larval abundance
was quite variable in both space and time. For instance,
on 12 July only five of the 10 A. vaginatum (collected
from approximately as many host pines) had larvae, and
the mean was 0.5 larvae per plant. On 3 August all of 16
A. vaginatum (collected from approximately as many
host pines) had larvae, and larval abundance ranges from
1 to 10 per plant with a mean of 3.25 per plant. On this
same date (3 August), an additional 146 larvae were col-
lected from 18 dwarf mistletoe plants all growing on a
single tree for a mean of 8.0 larvae per plant. In general,
larval abundance appeared to increase over the course of
the summer, peaking in mid-August, but larvae were still
common into the middle of September.
Associated fauna. In addition to D. alterno-
squamella, at least three other species of lepidopteran
herbivores were also found feeding on A. vaginatum.
The most abundant of these three, Promylea lu-
nigerella glendella Dyar (Pyralidae, Phycitinae), was
only slightly less abundant than D. alternosquamella.
These two phycitines can be distinguished by the fact
that the head capsule widths of sixth instar P. 1. glen-
della is a mean of 0.75 mm (N = 6, SE = 0.0097;
Mooney in prep.), approximately 20% smaller than the
mean of 0.96 mm (N = 9, SE = 0.111) for D. alter-
nosquamella. Although Heinrich (1920) describes and
Fic. 4.
Sixth instar D. alternosquamella and Southwestern dwarf
mistletoe. Scale bar 2 cm.
Fic.5. Pupae of D. alternosquamella. Scale bar 5 mm.
VOLUME 55, NUMBER 4
illustrates larval D. alternosquamella, the only descrip-
tion of P. 1. glendella is that of the adult (Dyar 1906,
Heinrich 1956). Far less common, but also feeding on
A. vaginatum was Callophorys (Mitoura) spinetorum
Hewitson (Lycaenidae), the Blue Hairstreak. On three
occasions cryptically colored geometrid larvae were
also found feeding on the dwarf mistletoe. Most at-
tempts at rearing C. spinetorum (nine of 10) and all at-
tempts at rearing the geometrids were unsuccessfully
due to parasitoids. In addition to lepidopterans, Neo-
borella tumida (Hemiptera: Miridae) and unidentified
mites (Acari) were common and, especially the latter,
present on most plants. The relatively high abundance
of both N. tumida and the mites, along with the fact
that no predation events were observed, suggests they
are herbivores.
DISCUSSION
The feeding behavior of D. alternosquamella is con-
sistent with the general patterns previously described
for related taxa: Almost all pyralids are concealed feed-
ers, and the Phycitinae in particular are known to feed
within host plants (Neunzig 1987). What may be
somewhat unique is that D. alternosquamella appar-
ently changes feeding modes during their develop-
ment; they are terminal shoot feeders early in their on-
togony and later progress to mining modes. Because
dwarf mistletoes are leafless, it is the shoots that are
mined, and this feeding might more properly be com-
pared to stem boring.
This mining has significant consequences for dwarf
mistletoes. Dasypyga alternosquamella can be abun-
dant, and because it feeds on shoots even low levels of
herbivory results in death of all plant tissue distal to
the site of herbivory. Other, unrelated work has
documented the nearly complete destruction of all
dwarf mistletoe shoots by D. alternosquamella in a
several-hectare area of heavily parasitized ponderosa
pines near Boulder, Colorado during the summer of
1998 (unpubl. data). At the Boulder site, D.
alternosquamella was the only abundant herbivore. At
the Manitou Experimental Forest the lepidopteran as-
143
semblages appeared to be more diverse; P. |. glendella
was nearly equal in abundance to D. alternosquamella.
The nature and consequences of interactions between
D. alternosquamella and other dwarf mistletoe associ-
ated fauna are unknown, but diversity between sites
can vary.
ACKNOWLEDGMENTS
This research was supported in part by funds provided by the
Rocky Mountain Research Station, Forest Service, U.S. Depart-
ment of Agriculture. Brian Geils (USDA Rocky Mountain Re-
search Station) provided valuable advice on dwarf mistletoe-related
matters. M. Deane Bowers (University of Colorado, Boulder) pro-
vided advice on rearing. Yan B. Linhart (University of Colorado,
Boulder) and M. Deane Bowers reviewed drafts of this manuscript.
Herbert Neunzig assisted in the literature review and identified P.
I. glendella. Kathy Thomas provided invaluable assistance in the
field and laboratory. Two reviewers provided insightful comments
that improved this manuscript. Wayne Shepherd and Steve Tapia
(USDA Rocky Mountain Research Station, Manitou Experimental
Forest) provided logistical assistance for this work.
LITERATURE CITED
Dyar, H. G. 1906. Descriptions of new American moths. J. New
York Entomol. Soc. 14:30-31.
Hawkswoktu, F.G. & D. Wiens. 1996. Dwarf mistletoes: biology,
pathology, and systematics. USDA Forest Service Agricultural
Handbook 709. 410 pp.
HEINRICH, C. 1920. On some forest Lepidoptera with descriptions
of new species, larvae, and pupae. Proceedings of the U.S. Na-
tional Museum 57:84.
. 1956. American moths of the Subfamily Phycitinae.
United States National Museum Bulletin 207. Smithsonian In-
stitution, Washington, DC. 581 pp.
NEuNzIG, H. H. 1987. The Pyralids, web worms, waxworms, cereal
worms, dried fruit worms, casebearers, etc. In F. W. Stehr (ed.),
Immature insects. Vol. 1. Kendall/Hunt Publishing. Dubuque,
Iowa. Pp. 462-494.
RAGONOT, N. 1887. Diagnoses of North American Phycitidae and
Galleriidae. Published by the author, Paris. 20 pp.
REICH, R. 1992. Voracious moth larvae feed heavily on lodgepole
pine dwarf mistletoe shoots in Prince George Forest Region.
Pest Management Progress Report 11. Victoria, BC: British Co-
lumbia Ministry of Forests. 22 pp.
STEVENS, R. E. & F. G. Hawkswortu. 1970. Insects and mites as-
sociated with dwarf mistletoe. USDA Forest Service Research
Paper RM-59. 12 pp.
Received for publication 12 June 2001; revised and accepted 3
October 2001.
Journal of the Lepidopterists’ Society
55(4), 2001, 144-149
RECOGNITION OF WESTERN POPULATIONS OF SPEYERIA IDALIA
(NYMPHALIDAE) AS A NEW SUBSPECIES
BARRY E.. WILLIAMS!
Department of Animal Biology, University of Ilinois, 505 S. Goodwin Ave., Urbana, Illinois 61801, USA
ABSTRACT. Western populations of Speyeria idalia are described as separate subspecies, S. idalia occidentalis, new subspecies. East-
ern and western populations can be diagnosed morphologically by differences in the size of spots on the underside of the hindwings. Further-
more, mitochondrially encoded cytochrome oxidase I and II genes reveal five synapomorphies for an extant eastern population in Pennsylvania,
indicating a unique genetic diversity possessed by this population. Recognition of subspecies status for the eastern population may lead to a pe-
tition for an emergency listing under the Endangered Species Act of 1973.
Additional key words: Mitochondrial DNA, morphology, conservation, taxonomy.
Taxonomy within the genus Speyeria Scudder has
long been a troublesome issue among lepidopterists
(dos Passos & Grey 1945, 1947, Moeck 1957, Arnold
1983). No consensus has been reached on the number
of species within the genus and relationships among
subspecies are even less well resolved (Howe 1975,
Arnold 1983, Hammond 1985, Scott 1986). Confusion
among taxa stems from numerous examples of poly-
morphism, sexual dimorphism, convergence and clines
(Hovanitz 1941, 1943, Moeck 1957, Rindge 1987,
Grey 1989, Hammond 1991). Most of the variation
within the genus is found among western North
American species and subspecies, whereas eastern taxa
are perceived as relatively well resolved. However, be-
cause previous research has been focused on the prob-
lematic western species, intraspecific variation in the
eastern fauna may have been overlooked.
Speyeria idalia (Drury) is an example of an eastern
species that has not been an issue of contention (Moeck
1957, Hovanitz 1963). Because S. idalia is so easily
identified among Speyeria species, intraspecific varia-
tion may not have been thoroughly examined. The
original range of S. idalia extended from the plains of
North Dakota and Colorado, east to Virginia and Maine
(Howe 1975, Scott 1986). However, within the last cen-
tury, populations of S. idalia have been extirpated over
most of the species’ range due to habitat destruction
(Hammond & McCorkle 1984, Hammond 1995, Swen-
gel 1997). Only two populations are known to exist east
of Illinois, one in Pennsylvania and the other in Vir-
ginia. Hence, if differences do exist between eastern
(=Pennsylvania and Virginia) and western (=all other)
populations, taxonomic status could have important im-
plications for the conservation of the remaining eastern
populations. The purpose of this research is to examine
mitochondrial DNA (mtDNA) and morphological vari-
ation among populations of S. idalia to determine if: (1)
significant intraspecific variation exists, (2) there is a
‘Current address: Laboratory of Molecular Biology and Howard
Hughes Medical Institute, University of Wisconsin, 1525 Linden
Drive, Madison, Wisconsin 53706, USA.
pattern to the variation, and (3) any patterns of variation
are worthy of taxonomic recognition.
MATERIALS AND METHODS
Morphological variation was examined via measure-
ments of the size of white spots on the underside of the
hindwings. All measurements were taken from museum
specimens at the American Museum of Natural History
(New York, NY). Sample sizes and collecting localities
are provided in Table 1. No specimens from counties
adjacent to the extant Pennsylvania population could be
found in the American Museum collections; therefore,
analyses of eastern populations used specimens from ex-
tinct populations from nearby counties in Pennsylvania,
New York, and New Jersey to represent morphological
variation of the extant Pennsylvania population (Table
1). Traits selected for analyses (Table 2, Fig. 1) were
those determined by a pilot study to be most variable
and therefore most likely to provide information on pat-
terns of intraspecific variation. Maximum diameter of
each spot was measured using a hand held digital mi-
crometer. To account for allometric relationships all
measures were transformed using the equation X’ = log
(X/Y), where Y = size measure. The size measure used
was the length of the second cubital vein. An alternative
measure for size, wingspan, was not incorporated be-
cause not all specimens had both wings and because
some specimens had wings folded for display. A regres-
sion of second cubital vein length on wingspan was sig-
nificant (r? = 0.62, p < 0.001), suggesting that second
cubital length is a valid size measure. Analysis of mor- —
phological data included both a MANOVA, incorporat-
ing all 11 traits as a group, and univariate F-tests, to ex-
amine variation at each trait independently. Both
analyses tested the effects of region (eastern vs. western;
Table 1) and sex on pattems of variation. All analyses
were completed using Systat v. 5.0.
Mitochondrial DNA analysis used samples collected
from extant populations found across the range of the
species. Sample sizes and locales are provided in Table
3. Collection of tissue from those populations of special
VOLUME 55, NUMBER 4
TABLE 1. Sample sizes and collection locales for morphological
measurements from museum specimens.
State County Male Female
Eastern populations
Pennsylvania Montgomery
Alleghany
Butler
Renssalaer
Albany
Columbia
Westchester
New York
Unknown
New Jersey Passiac
Essex
Morrris
Hunterdon
Unknown
=
New York
—
=
ADH OAWWHUDSUC0 WWE
TWnwhOkNDANHHrNHH DL
as |
—
Total eastern
Western populations
Nebraska Pawnee
Otoe
Lancaster
Douglas
Stanton
Dixon
Cedar
Knox
Keyapaha
Cherry
Douglas
Johnson
Riley
Towa Dickinson
Plynouth
St. Clair
Franklin
Polk
Guthrie
Pocahantas
Illinois Cook
Mercer
Iroquois
=
bo
RAW WHAUANWWNWNUERrPOKNW W # UL
NOWUGCDSCODFORFRFOGCONOCORrPWWOWrOCS
—
S
(oy)
Total western
conservation concern (Pennsylvania, Illinois, lowa, and
Wisconsin) consisted of the removal of the anterior leg
on the right side so that specimens could be released
alive. Anecdotal observations of seven captive females
suggested no decrease in survivorship or oviposition
ability following leg removal. These results indicate
SCRE sal SGRICA
os = ~HNDCELL
; —CUA2
Fic. 1. Arrows indicate underside hindwing spots measured for
morphological analysis. Trait abbreviations correspond with Table 1.
TABLE 2. Traits selected for analysis and the corresponding ab-
breviations used in Figs. 1-3. Cell names follow Scott (1986) (Figure
53, p. 146).
Trait Abbreviation
Basal-most spot in cell 1A + 2A LA2A
Basal spot in cell CuA2 CUA2-A
Median spot in cell CuA2 CUA2-B
Basal-most spot in cell CuA1 CUAI
Basal-most spot in cell M3 M3
Basal-most spot in cell M2 M2
Basal-most spot in cell M1 MI
Basal-most spot in cell Rs RS
Basal-most spot in cell Se + R1 SCRI-A
2nd most basal spot in cell Sc + R1 SCRI-B
Basal-most spot in the hindwing cell HCELL
that this method may be useful for future genetic stud-
ies in other insect species of conservation concern, al-
though more thorough, species specific, studies should
be conducted. DNA extractions were carried out by
digesting homogenized tissue at 65°C for 3-12 hours
in: 10 mM Tris-HCl, 10 mM EDTA, 50 mM NaCl, 2%
SDS, 20 yl dithiothreitol, and 0.4 mg Proteinase K.
Nucleic acids were extracted with an equal volume of
phenol, the aqueous phase was transferred to a new
microcentrifuge tube and the organic extraction re-
peated with 24:1 chloroform:isoamyl alcohol. Total nu-
cleic acids were precipitated with the addition of 1/10
volume 3 M sodium acetate and 3 volumes of cold
95% ethanol. Nucleic acids were resuspended in
50-100 ul of sterile water.
Mitochondrial DNA sequences were generated by
direct sequencing of PCR products from partial cy-
tochrome oxidase I and II genes (COI and IT). Primers
used for PCR amplification were: C1-J-2183 (alias
Jerry) 5’-CAACATTTATTTTGATTTTITGG-3’ (Si-
mon et al. 1994) and TK-N-3772 5’-GACCATTA-
CTTGCTTTCAGTCATCT-3’ (listed as TK-N-3782 in
Sperling & Hickey 1995). PCR reactions were carried
out using: 60 ng genomic DNA, 20 mM Tris-HCl, 50
mM KCl, 3 uM MgCl, 0.25 mM of each dNTP, 2 uM
each primer, 1 Unit Taq DNA polymerase, and water
to a final volume of 20 ul. Each PCR reaction was then
subjected to 30 cycles of amplification at the following
conditions: 94°C for 30 seconds, 45°C for 30 seconds,
72°C for 1 minute. PCR reactions were purified using
Qiagen (Valencia, CA) PCR purification columns, fol-
lowing manufacturers recommendations.
Sequencing reactions used two internal primers, se-
quencing in opposite directions, with 30 base pairs of
overlap. Sequencing primers were: TL2-N-3014 (alias
PAT) 5’-TCCAATGCACTAATCTGCCATATTA-3’ (Si-
mon et al. 1994), modified to 5’-TCCATTACATAT-
AATCAGCCATATTA-3’ and C1-J-2983 (alias LANAE)
146
Fic. 2. Comparison of similarly-sized males (upper) and fe-
males (lower) from western (left) and eastern (right) populations.
Western specimens were from Nebraska and eastern specimens
from New York.
5’-TACCTCCTGCTGAACATTCT-3’. Sequencing re-
actions were carried out using the Perkin Elmer (Fos-
ter City, CA) Big Dye cycle sequencing kit, following
manufacturer recommendations. Sequences were vi-
sualized on an ABI 377 automated sequencer at the
University of Illinois Biotechnology Sequencing Cen-
ter. All sequences were edited using EditView V1.0.1
and aligned using Sequencher V3.0.
Phylogenetic hypotheses were constructed using
PAUP* V4.0b3 (Swofford 1998) via maximum parsi-
mony analysis with a heuristic search and tree bisection
and reconnection (TBR) branch swapping. Sequences
generated from Speyeria cybele (Fabricius) and S.
nokomis (Edwards) were used as outgroups to root the
tree. Phylogenetic reconstructions were without
weighting schemes because all substitutions but one
within S. idalia were transitions. Statistical support for
nodes was estimated using 100 bootstrap replicates.
RESULTS
Morphological analyses revealed that all traits exam-
ined were significantly different between eastern and
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
western populations in both the multivariate test
(Wilks’ lambda = 0.381, F = 36.006, df = 11, 244, p<
0.001) and all univariate F-tests (p < 0.001 for all
traits). As an example, Fig. 2 shows a male and female
from an eastern population next to a similarly-sized
male and female from a western population. Only one
of the traits, the basal-most spot in the third median
cell, was significantly different between the sexes (Uni-
variate F = 7.523, df = 1, 254, p = 0.007); however, the
multivariate results suggest that as a group, the traits
did not differ between the sexes (Wilks’ lambda =
0.946, F = 1.254, df = 11, 214, p = 0.252). Figure 3
graphically represents the differences in the trait
means between eastern and western populations for
the non-transformed data.
Results of mtDNA sequence analysis revealed 30
variable sites, 18 of which were parsimony-informative
(Genbank accession number AF295040). Within the
COI gene there were 11 parsimony-informative sites:
three first-position and one second-position non-
synonymous substitutions and seven third-position
synonymous substitutions. Within the COII gene
there were seven parsimony-informative sites, all
third-position synonymous substitutions. Maximum
parsimony analysis resulted in five most-parsimonious
trees of 126 steps. All individuals in the Pennsylvania
population shared the same unique haplotype, which
has five synapomorphies (Fig. 4). All other populations
sampled revealed a total of 22 unique haplotypes with
no apparent geographic associations among them, with
the possible exception of some Wisconsin haplotypes
(Fig. 4). Also, 45 out of 84 individuals sampled from
the 17 western populations shared the same haplotype
(haplotype 1, Fig. 4), suggesting little genetic structur-
ing among those populations.
Speyeria idalia idalia (Drury |1773])
See dos Passos and Grey (1945, 1947) for a descrip-
tion and type specimens of S. i. idalia. Diagnostic char-
acters that separate subspecies are provided below.
Speyeria idalia occidentalis Williams,
new subspecies
Diagnostic characters. Speyeria idalia occidentalis can usually
be diagnosed by eye because the hindwing spots are usually much
larger than in S. i. idalia. The basal-most diffuse white spot (not in-
cluded as a trait used in this study) in the second cubital cell (Figs.
1, 2) is often entirely absent in S. i. idalia but usually present in S i.
occidentalis. While all of the ventral hindwing spots included in this
study are significantly larger in S. i. occidentalis, the most pro-
nounced differences are in cells A1A2, M1 and RS (Figs. 3, 4). In S.
i. occidentalis the spot in A1A2 is usually greater than 10 mm,
whereas S. i. idalia it is usually less than 8 mm (these numbers are
general rules of thumb for quick reference and therefore do not in-
corporate size or sex differences). For M1, S. i. occidentalis is usu-
VOLUME 55, NUMBER 4
14
ZZ) EASTERN
12 - (—] WESTERN
fi
= 1@s
E
eS
fH 8
ma
2
ae 8
i ps
oe)
ae =
Be a
Alle
Fe AVL Al
AIL
7 a as MI ORC Se SN
ser ah a apy WP Wh WH O_o Pind
b ow oo & » Remo ee”
Fic. 3. Means and standard errors of underside hindwing cell
spots for raw data. Trait abbreviations correspond with Table 1 and
Fig. 1.
ally >3 mm, whereas S. i. idalia is usually <3 mm. For RS, S. i. occi-
dentalis is usually >4 mm, whereas S. i. idalia is usually <4 mm.
Description. Male wingspan 63-92 mm (N = 108). Dorsal sur-
face of forewing with orange background coloration and the usual
Speyeria pattern of black spots and bars. Black outer marginal bor-
der with a row of white spots. Hindwing is orange with hints of blue-
black basally and entirely with blue-black background color from the
submedian to outer margin. A postmedian row of orange or fulvous
spots, a submarginal row of white spots and an outer marginal row of
white spots. Ventrally, forewing lighter orange background with sub-
marginal and outer margin rows of white spots. Also, varying num-
bers of diffuse apical and subapical white spots. Hindwing is unique
to Speyeria idalia. Background is entirely yellow to black with white
spots, either single or multiple, in every cell. Female wingspan
71-110 mm (N = 33). Dorsal surface, forewing with larger marginal
black border than in males and a marginal row of white spots. Black
apical and subapical background color with white spots. Hindwing as
in male except with a post median row of white spots, rather than or-
ange. Ventrally, same as the male except for a deeper fulvous back-
ground coloration in the forewing, more black background col-
oration apically and in the margin.
Types. HooryPeE: (male), 1 mile S.E. of Crete, IL, 7 July 1965
(R.R. Inwin). ALLOTYPE: (female), same location data as holotype, 23
August 1965 (R.R. Irwin). PaRATYPES: Two males, one female, all
with same collection and data as holotype. Deposition of specimens:
All specimens at the Illinois Natural History Survey collections.
DISCUSSION
Morphological data suggest that eastern and western
populations were distinct entities; in that specimens
from eastern populations had spots on the underside of
the hindwing that were significantly smaller. The ge-
netic basis for these morphological traits has not been
determined, so the potential exists for differences be-
tween eastern and western populations to be correlated
with as of yet undetermined environmental variables,
rather than indicative of a unique evolutionary history.
However, mtDNA sequence analysis suggests that the
147
Wl-c1
1 WI-c3
WI-b2
WI-b5
1 Wai
Wl-a2
1 WI-a3
Wl-a4
Wl-a5
95 Wl-c2
3 WI-d4
WI-d5
NE-b1
1 NE-b4
SD-5
KS-2
1 IA-a5
WI-b3
1 lA-a4
NE-c1
NE-c3
1 SD-2
MO-a5
1 |A-a2
PA-1
98 PA-2
5 PA-3
PA-4
1 PA-5
|A-a1
NE-b3
IA-b3
MN-5
KS-3
KS-4
IL-b4
IL-b5
IL-a5
MO-b5
NE-c2
HAPLOTYPE 1
WI-b1
Speyeria cybele
Speyeria nokomis
100
— }
47
Fic. 4. Strict consensus of the five most-parsimonious trees re-
sulting from maximum parsimony analysis of mitochondrial DNA se-
quence data from cytochrome oxidase I and II genes (126 steps, con-
sistency index = 0.96). Bootstrap values >60 are shown above
branches and the number of synapomorphies for each clade are
shown below branches. Forty-five specimens that shared a single
haplotype were collapsed together into “haplotype 1” to conserve
space. Sample labels correspond with collection locations in Table 3
and numbers in parentheses indicate which of the five individuals
per sample correspond to a given haplotype.
Pennsylvania population is a distinct evolutionary lin-
eage that can be diagnosed with 5 synapomorphies,
which are unique and fixed in the population. Hence,
mtDNA support, in part, differentiation of the eastern
populations. Based on these data I recommend that the
eastern (=Pennsylvania) and western (=all other) popu-
lations be recognized as separate subspecies, and the
morphological characters be used as a means for exter-
nal diagnosis of each subspecies. Fortunately, the sub-
species of S. idalia named here can be identified on the
basis of easily quantified measures (spot size), which
make diagnosis relatively straightforward when com-
pared with other Speyeria species, in which subspecies
are typically differentiated by slight differences in basal
148
TABLE 3. Sample sizes and collection locales for mitochondrial
DNA tissue collection.
State County Sample Size
Pennsylvania Lebanon 30
Nebraska — a Keyapaha 5
Nebraska — b Greeley 5
Nebraska — c Kearney 5
Kansas Riley 5
Iowa —a Muscatine 5
Iowa —b Plymouth 5
Illinois — a Ogle 5
Illinois — b Mason 5
Illinois — c Cass 4
Wisconsin — a Crawford 5
Wisconsin — b Iowa 5
Wisconsin — c Dane 5
Wisconsin — d Portage 5
Missouri — a St. Clair 5
Missouri — b Vernon 5
Minnesota Lincoln 5
South Dakota Hughes 5
Total sample size 114
coloration of the hindwing discal area (Howe 1975,
Scott 1986). Figure 3 can be used as a guide to distin-
guish among S. idalia subspecies. For example, the
maximum diameter of the A1A2 cell spot used in this
study (Fig. 1) is less than 8 mm in S. i. idalia but
greater than 10 mm in S. i. occidentalis (Fig. 3).
If the relationship between mtDNA, morphology
and longitude were to be generalized, then the Vir-
ginia population would be recognized as the same sub-
species as the Pennsylvania population. However, the
formal status of the Virginia population will remain un-
determined until data from this population can be in-
cluded in the analyses. Assuming that the neotype des-
ignation for S. idalia (New York, NY) of dos Passos and
Grey (1945) applies to the eastern populations in gen-
eral, the name for the Pennsylvania subspecies be-
comes Speyeria idalia idalia. Western populations
therefore fall under the subspecies name Speyeria
idalia occidentalis, new subspecies.
An initial examination of the male genitalia from five
eastern and 25 western specimens did not result in any
distinguishable differences (unpubl. data). However,
other anecdotal evidence of differentiation does exist.
Previous descriptions of habitat use note that eastern
populations are typically found in xeric habitats
whereas western populations are found in mesic habi-
tats (Scudder 1889, Opler & Krizek 1984). Also, Bar-
ton (1996) notes that Viola saggitatta is the preferred
host plant for the Pennsylvania population, whereas
other studies focusing on western populations have
noted V. pedatifida and V. pedata as preferred host
plants (Swengel 1997, Kelly & Debinski 1998).
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Subspecies status for the eastern population has im-
portant conservation implications. The Pennsylvania
population is found on a military installation where
current land use practices threaten to destroy the re-
maining S. i. idalia habitat (Barton 1996). Because this
population resides on federally owned land, formal
protection can only be afforded under federal legisla-
tion, i.e., the Endangered Species Act of 1973 (ESA).
Therefore, the designation of S. i. idalia may result in
a petition for an emergency listing under the ESA.
ACKNOWLEDGMENTS
I would like to thank my advisors Drs. Ken Paige and Jeff Brawn
for assistance and support, and Eric Quinter for his patience and as-
sistance during my visit at the American Museum of Natural His-
tory. Deane Bowers, Diane Debinski, Carla Penz and an anonymous
reviewer provided many helpful comments that greatly improved an
earlier draft of this manuscript. Funding for this work was provided
by a collections study grant from the American Museum of Natural
History and a Cooperative Agreement with the U.S. Fish and
Wildlife Service.
LITERATURE CITED
ARNOLD, R. A. 1983. Speyeria callippe (Lepidoptera: Nymphal-
idae): application of information-theoretical and graph-cluster-
ing techniques to analyses of geographic variation and evalua-
tion of classifications. Ann. Entomol. Soc. Am. 76:929-941.
BarTON, B. 1996. Final report on the regal fritillary 1992-1995:
Fort Indiantown, Annville, Pennsylvania. Report to U.S. De-
partment of Defense.
Dos Passos, C. F. & L. P. Grey. 1945. A genitalic survey of Argyn-
ninae (Lepidoptera, Nymphalidae). Am. Mus. Nov. 1296:1—29.
. 1947. Systematic catalogue of Speyeria (Lepidoptera,
Nymphalidae) with designations of types and fixations of type
localities. Am. Mus. Nov, 1370:1—57.
Grey, L. P. 1989. Sundry Argynnine concepts revisited (Nymphal-
idae). J. Lepid. Soc. 43:1-19.
HAMMOND, P. C. 1985. A rebuttal to the Arnold classification of
Speyeria callippe (Nymphalidae) and defense of the subspecies
concept. J. Res. Lep. 24:197—208.
. 1991. Patterns of geographic variation and evolution in
polytypic butterflies. J. Res. Lep. 29:54—76.
. 1995. Conservation of biodiversity in native prairie com-
munities in the United States. J. Kan. Entomol. Soc. 68:1-6.
HAMMOND, P. C. & D. V. McCorke. 1984. The decline and ex-
tinction of Speyeria populations resulting from human environ-
mental disturbances (Nymphalidae: Argynninae). J. Res. Lep.
99:21 7-224.
Hovanitz, W. 1941. Parallel ecogenotypical color variation in but-
terflies. Ecol. 22:259-284.
. 1943. Geographical variation and racial structure in Ar-
gynnis callippe in California. Am. Nat. 77:400—-425.
. 1963. Geographical distribution and variation of the genus
Argynnis. I. Introduction I. Argynnis idalia. J. Res. Lep.
E23"
Howe, W. H. 1975. The butterflies of North America. Doubleday
& Company, Garden City, New York.
KELLy, L. & D. M. DEBINSKI. 1998. Relationship of host plant den-
sity to size and abundance of the regal fritillary Speyeria idalia
Drury (Nymphalidae). J. Lepid. Soc. 52:262-276.
Moeck, A. H. 1957. Geographic variability in Speyeria: comments,
records and description of a new subspecies (Nymphalidae).
Milw. Entomol. Soc. Spec. Pub.
OpLER, P. A. & G. O. KrizEK. 1984. Butterflies east of the Great
Plains, an illustrated natural history. John Hopkins University
Press, Baltimore, Maryland.
VOLUME 55, NUMBER 4
RINDGE, F. H. 1987. Speyeria collection of Paul Grey to the Ameri-
can Museum of Natural History. J. Lepid. Soc. 41:123.
Scort, J. A. 1986. Butterflies of North America. Stanford Univer-
sity Press, Stanford, California.
SCUDDER, S. 1889. Butterflies of the eastern United States. Cam-
bridge Press, Cambridge, Massachusetts.
SIMON, C., F. Frati, A. BECKENBACH, B. Crespl, H. Liu & P. FLOOK.
1994. Evolution, weighting, and phylogenetic utility of mitochon-
drial gene sequences and a compilation of conserved polymerase
chain reaction primers. Ann. Entomol. Soc. Am. 87:651—701.
SPERLING, F. A. H. & D. A. Hickey. 1995. Amplified mitochondr-
ial DNA as a diagnostic marker for species of conifer-feeding
149
Choristoneura (Lepidoptera: Tortricidae). Can. Entomol.
127:277-288.
SWENGEL, A. B. 1997. Habitat association of sympatric violet-
feeding fritillaries (Euptoieta, Speyeria, Boloria) (Lepidoptera:
Nymphalidae) in tallgrass prairie. Great Lakes Entomol. 3:1-18.
SworrorD, D. 1998. PAUP* 4.0b3. Phylogenetic analysis using
parsimony (and other methods). Sinauer Associates, Sunder-
land, Massachusetts.
Received for publication 11 September 2000; revised and accepted 3
November 2001.
Journal of the Lepidopterists’ Society
55(4), 2001, 150-157
POPULATION STUDIES OF AERIA OLENA AND TITHOREA HARMONIA
(NYMPHALIDAE, ITHOMIINAE) IN SOUTHEASTERN BRAZIL
ANDRE V. L. FREITAS*
Museu de Histéria Natural, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109, 13083-970,
Campinas, Sao Paulo, Brazil
JOAO VASCONCELLOS-NETO, FABIO VANINI, JOSE R. TRIGO
Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109, 13083-970,
Campinas, Sao Paulo, Brazil
KEITH S. BROWN JR.
Museu de Hist6ria Natural and Departamento de Zoologia, Instituto de Biologia,
Universidade Estadual de Campinas, CP 6109, 13083-970, Campinas, Sao Paulo, Brazil
ABSTRACT. Populations of Aeria olena and Tithorea harmonia pseudethra (Lepidoptera: Nymphalidae: Ithomiinae) showed large varia-
tions in abundance along the year in four sites, with peaks at the end of the wet season. The sex ratio of captures in A. olena was male biased. In
A. olena, males showed longer residence times than females. Both species differ from other members of the Ithomiinae community in the re-
gion by feeding on Apocynaceae vines as larvae, not congregating in “ithomiine pockets” and having low population numbers in the dry season.
Additional key words: mark-recapture, Ithomiinae pockets.
The Ithomiinae (Nymphalidae) are an exclusively
Neotropical butterfly group (Fox 1967, Brown & Frei-
tas 1994) except possibly for the Australian genus
Tellervo Kirby (Ackery & Vane-Wright 1984). The sub-
family is distributed from Mexico to Argentina and is
largely restricted to moist forest habitats from sea level
up to 3000 m (Fox 1967, DeVries 1987).
Ithomiinae populations are considered difficult to
study due to low adult recapture rates, even in dense
pockets (Gilbert 1993). Thus there are few published
population studies in this subfamily (Drummond 1976,
Haber 1978, Young & Moffett 1979, Vasconcellos-Neto
1980, 1991, Trigo 1988, Freitas 1993, 1996, Pinto & Motta
1997). Knowledge of population parameters is impor-
tant to the understanding of this family and of the whole
butterfly community (DeVries 1994, Freitas 1996).
The genera Tithorea Doubleday (two species) and
Aeria Hiibner (three species) both belong to basal
branches of the Ithomiinae (Brown & Freitas 1994),
with aposematic “danaoid” larvae feeding on Apocy-
naceae (such as Prestonia acutifolia (Benth.) K.
Schum. and P. coalita (Vell.) Woodson) and bearing
fleshy tubercles (Brown 1987, Brown & Freitas 1994,
Trigo et al. 1996). In Southeastern Brazil, both species
are most common in semi-deciduous forests of the in-
terior, being scarce in the humid forests of the Atlantic
mountain slopes and coastal plain.
*To whom correspondence should be addressed.
This paper describes the population parameters of
three populations of Aeria olena olena Weymer and
one of Tithorea harmonia pseudethra Butler in semi-
deciduous forest fragments in SE Brazil, comparing
them with other populations of Ithomiinae in Brazil.
STubDY SITES AND METHODS
The present study combines four data sets collected
by different researchers from 1974 to 1998 in four dif-
ferent sites in SAo Paulo state, southeastern Brazil. Al-
though the basic method to study all the populations
was mark-recapture (see Freitas 1993, 1996), there
were some differences among data sets, requiring that
methods and results be presented separately. The re-
gional climate in the four sites is markedly seasonal,
with a warm wet season from September to April and
a cold dry season from May to August.
The most recent study area was in the Santa Gene-
bra Forest Reserve (SG, 22°49’S, 47°07’W), a 250 ha
forest fragment in Campinas. The study area is cov-
ered by semideciduous forest, with annual rainfall near
1400 mm and an average annual temperature of
20.6°C (Morellato & Leitéo-Filho 1995). A large part
of the forest is old secondary growth, with a predomi-
nance of forest edge plants and lianas.
In this area, a mark-recapture census of A. olena ex-
tended from January 1997 to June 1998 (AVLF and
FV), along an interior trail 1100 m long, with 103 field
days of about four hours each at intervals of 2 to 15
VOLUME 55, NUMBER 4
Oy,
INDIVIDUALS PRESENT/DAY
3)