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jtmnwi of thv Lejtidojilzriats' Snriety 158(1). 2004, 33-43
THE BUTTERFLIES (PAP1LIONOIDEA, HESPERIOIDEA) OF TIIE GALAPAGOS ISLANDS, ECUADOR: DISTRIBUTION, HOSTPLANTS AND BIOLOGY.
Lazaro Roque-Albelo
Charles Darwin Research Station, Galapagos Islands, EO. 17 01 3891, Quito, Ecuador, email: lazaro@tcdarwin.org. ec. and Biodiversity and Ecological Processes Research Croup, Cardiff School of Binsoiences Cardiff University, P.O. Box 915, Cardiff CF10 3TL, Wales. United Kingdom
ABSTRACT. Tile butterfly fauna of the Galapagos islands is reviewed. Ten species are recorded from the Archipelago. Leptotes parrlia-sioides (Wallengren), \Jrbaim& dorantex galapagensis (WilliamsJ, and Agranlis vtijiillae galapagensis (Holland) are endemic and the others are Neotropical faunal elements, A number of new island and host plant records are given.
Additional key words: flower visi tat ion, p honology. \ cotrop ical region.
The butterfly fauna of the Galapagos Islands was first discussed by Williams (1911). Six species were then recorded from the archipelago. Previous reports of Galapagos butterflies were published by Wallengren (I860, 1861). Butler (1877) and HollandJ(1889). .Subsequent to Williams' revision, little mention was made of the butterflies in the Galapagos until Beebe (1923) and Parkin et al. (1972) added two species of milkweed butterflies, Danaus plextppus (Linnnaeus) and Danaus gilippus (Cramer), to the fauna. A literature-based checklist of the insects of Galapagos by Linsley and Usinger (1966) and Linsley (1977) listed eight species and indicated on which island each species occurred. During the last 33 years only two faunistic notes have been published. Onore and Mielke (1988) reported Calpodes ethlius (Stoll) and Hoque et al. (1997) added Hemiargus ranum (Dognin). Additional information such as flower visits and host plant records were given by Linsley (1966), McMullen (1986, 1990, 1993), McMullen and Viderman (1994) and Roque (1998a, b). Finally, Peck et al. (1998) listed the introduced insect species that occur on the archipelago and included C. ethlius and H. rainon.
This paper reports new information on Galapagos butterflies gathered during 8 years of fieldwork on the Islands. The butterfly fauna of the archipelago includes 10 species and subspecies, three of which are presumed to be endemic.
Materials and Methods
From November 1994 to June 2002 the author conducted daytime observations and collections of butterflies in the Gal&pagos Islands, A total of 387 specimens were sampled and identified, and observations on habitat associations and behavior of each species was noted. 'Die methods included: a) field collections with a light mesh net, b) field observations of adult behavior and visits to food sources, c) collecting and rearing of immature stages.
Immatures were collected and reared in the laboratory. Eggs and larvae were reared in plastic bags at room
temperature under the natural day-night photoperiod regime. Fresh leaves were provides at least every other day. Pupae were kept moist until adults hatched.
The majority of this material was deposited in the Invertebrates Collection of the Charles Damin Research Station (CDRS), Santa Cruz Island, Gal&pagos Islands, Some of the other specimens are in the following collections: Pontificia Universidad Carolica del Ecuador (PUCE), Quito, Ecuador, Museo Ecuatori-ano de Ctencias Naturales (MECN), Quito, Ecuador and California Academy of Sciences (CAS), San Francisco, USA. Additional records of distributions were obtained from the literature and unpublished data from Dr Bernard Landry, whose material is deposited in the Canadian National Collection (CNC), Ottawa, Canada.
The classification adopted here follows Ackery and Vane-Wright (1984) and Ackery et al. (1999). A brief description is given for the immature stages of the endemic species. The nomenclature of the plants follows McMullen (1999).
The following abbreviations were used: FW = fore wings, !)KW = dorsal (ore wings, VFW = ventral lore wings. HW = hind wings, 1)1 IVV = dorsal hind wings, VHW = ventral hind wings. DVV = dorsal side of the wings, VW = ventral side of the wings. The size of die butterflies in mm refers to their fore wing length.
Results
Doubtful Records
Previous authors reported several species whose occurrence in the Galapagos has not been demonstrated convincingly. It is better to exclude these as probable mis identifications or mislabelling as is explained below.
Actinote sp. (Nymphalidae): Daniel Fitter, a naturalist guide with the Galapagos National Park Service, sent a color photograph to the author for examination. The photograph, taken on Media Luna, Santa Cruz Island. was published in Fitter et al. (2001:115) and was
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JOl.HVU. Ol■'THE LEI'IUOITEHISTS" SOCIETY
identified as Actinole sp. (Nvmphalidae). It lias not l>een collected or seen in tlie archipelago since tlie picture was taken. The establishment in the Galapagos ol Ll lis species will be onlv cotilinned or refuted when additional material is found.
Hypolimmnas misipptis (Lhuie). (Nyinphalinae): The only notice of the occurrence of this species in the archipelago is from one specimen deposited in the American Museum of Natural History, New York, The specimen has the following label data: "Conway Bay, Indefatigable Is, Galapagos, March 15 1935, Crocker Exped". It is likely that this is a mislabelling or if the record is valid it was a migrant individual. The species has never been seen or collected since.
Agratilis vanillin: lucina (C, & R. Felder): This species was reported by Moreno et al. (1997). It is likely a niisideiitifieation of the (ialapagos's endemic sob-species Agranlis lanilhicgalapage/isLt (Holland. 1889).
Heteropia sp. (Hesporiidae): One male from Albemarle [Isabcla] was reported by Van Duzee (1933), although the author suggested that the specimen had been mislabeled and collected in Mexico.
Erynnis funeralis (Scudder & Burgess), (Hesperiidae): One specimen (as Thanos persius (Scudder)) deposited in the CAS is said to have been collected in Chatham [San Cristobal], 15 IV 1932 by M Willows Jr. However, Van Duzee (1933) suggested the specimen was mislabelled.
Leptotes trigemmstus (Butler), (Lycaemdae); Larsen (1991) mentioned this species from the Galapagos Islands but it is likely a niisideiitifieation of the sister taxon Leptotes parrhasioides (Wallengren), as noted by Baliut and Johnson (1995).
Phocbis agarithe (Boisduval) (Pieridae); Moreno et al. (1997) reported this species from the archipelago. 'ftiesc authors probably niisidentified Phoehh sennae marcellina (Cramer) as this species which is superficially similar in size and color, P. agarithe has never been collected on the Galapagos Islands.
Species Accounts
JIespekiidae
Pyrgiuae
Urbantis dorantes galapagensis (Williams 1911)
"Galapagos Skipper"
(Fig. 1)
Eudamus galapagensis Williams 1911:3(0; Drauilt 1921:H53; Beubc
1923:55; Van Dutjce- 1933:144. Urbanus fivrantcs fcaiapagensis: Kvans 1952:93; Linslev & Usinger
1966:157; Parkin et al. 1972:103. UrbatiHs galapagen&k Williams & Eiqyward 1944:93.
Diagnosis. FVV = 15-20 mm. This is the only tailed skipper of the archipelago. FW dark brown with
translucent spots and a pattern of lilac and brown bands in VliW, [n some specimens, the hyaline spots are smaller. The lilac bands on (lie \ \\ lend to lie pale with lighter brown or yellowish scales. The sexes are similar, but males have coastal folds.
Geographical distribution. Reported from Bal-tra, Eden, Espahola, Floreana, Geuovcsa, lsabela, Pinzon, San Cristobal, Santa Cruz and Santiago (Lins-ley & Usinger 1966). Additional records include: Fer-nandina, Piuta and Rahida.
Natural history. Adults were common in open areas, roadsides, and forests from sea level to the highest elevations. The flight is erratic and low to the ground. Both sexes were seen visiling flowers. Oviposition was observed on leaves and ffower buds of various Legu-minosae. Eggs greenish laid singly on Phaseoltts atro-pnqxireits DC, P. mollis Hook, Desmodium incanum DC and Galaetia striata (Jacq). Caterpillar green with the head and prothoracic shield black. In the last in star the body is brown. Pupa pale brown speckled with darker brown,
Flowers visited. Acanthaceae: Justicia galapagana Lindau; Asteracoae: Darwiniothamnus tenuifolhts (Hook, f.) Hading, Sealesia cordata Stewart, Scalesia microcephala Robins, Scalesia villosa Stewart, Scalesia retrofiexa Hemsley; Boragiuaeeae; Cordia leitcophli/e-iis Hook, C. lutea Lam.; Convolviilaceac: Meiremia aegyptica (L.) Urban, Lcguminosae: Acacia insulae-lacobi Riley, P. mollis; Malvaceae: Sida rhombifotia L.; Plumbaginaceae: Plumbago scandens L.; Hubiaceae: Diodta radida Cham.& Schlecht; Rutaceae: Citrus limon (I..) Bum, Solauaeeae: Lycoperstcon chesmanii Kilev, Pi11 /salis pitbescens L.; Stereuliaceac: Walt he ha ovata Cav.; Verbenaeeae: Laniana camara L., Lantana pedunmlaris Anderss, and lAppiastrigidosa Mart &Gal.
Flight period. Multivoltiue with a peak in January and February.
Remarks. The holotype is in the California Academy of Sciences, San Francisco, USA (CAS); it bears tlie following lal>el information: Galapagos Is, Chatham I. [San Cristobal], X 15 06, Coll. By F, X. Williams, California Academy of Sciences Type No. 3209. A specimen de posited on CAS has the following label information: Puerto Valkrta, Jalisco, Mexico, VIII 21 32, M Willow Jr coll. K\p hi'52. This could lie a mislabelling. Evans (1952) treated it as a subspecies of Urbanus dorantes (Stoll, 1790) compared to Urhamts dorantes Santiago (Lucas, 1856) from Cuba and U. dorantes dorantes from mainland Ecuador, U. dorantes galapagensis is smaller and has shorter hindwiiig tails. The male valve is much less dentate and the arms of haqie have few spines. The genus Urbanus Ilubner 1807, is used provisionally here until revisionary work is undertaken. According to S.
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FiCS. I-S, GalSpagos bullerflies species. I, Urbamts galapagerais (DWand VW): Z, Catpodes ethlius (DW):3, Mate o[ Leptotes parrha-stohies (DW and VW); 4, Mile of Hemtergia ronton (DW): 5, Male of Hemitirgw, ronton (VW): 6, Female of Hemiargtis ranum (DW): 7, Female cX Phoebis lennae marcetiina (DW): 8, Male of Phoebvi sennae marcetlina (DW),
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Journal of tmf. Lepidopterists' Society
Steinhauser (pers. com.) die species may belong in a separate, mi describe genus.
Material studied. (70d and 16s1}. ECUADOR. Galapagos. ES-I'A.NIOLA: Hood, 15 V 1899, [B. E. Snodgrass, E. Heller] (CAS) (Id). Bahia Maiizanilb, 24 [V 1992, B. L[andry] (CNC), (2d). EER-NANDINA: 1 IX 1966, (CDRS) (Id), Cueva'Norte, Arid zone, 20 VI 1898, L. Rloque] (CDRS) (Id). FLOREANA: Arid zone, 130 m, 17 IV 1996. L. II (CDRS) (Si and 39). Cerra Pajas, ScaUsta forest, 320 m, 18 IV 1996, L. R (CDRS) (6tJ). Cerro Pajas 520 m, 19 IV 1996, L R (CDRS) (9d). Charles, 5 X 1899, [H. E. Snodgrass, E. Heller) (CAS) (&>'). !SABELA:TagnsCove, 111 1906, Alofype, F. X. W[illiams) (CAS) (I*). Cowley Mt, Vlll L906, F. X. W (CAS) (3d). Albemarle, 27 IV 1932. M. Willows (CAS) (1<J); 28 IV 1932, M. Willows (CAS) (1*). Albemarle, 12 VI 1899, [R. E. Snodgrass, E. Heller] (CAS) (IS). Albemarle Iguana Cove, 21 V 1932, M. Willows (CAS) (29). V[olcan] A[lcedo| SIX) m„ 2 IV 1996, I.. K (CDRS) <5d 2?). V Slierra) N[egra], 2 III 1989, S. B. Peek (CNC) (Id). VSN, 6 IV 1996, L. R (CDRS) (Id 19). VSN, Las mcrceditas 120m, 24 IV 1996, L. R (CDRS) (lo* 79). Sierra Negra pampas ferns zone, 24 IV 1996. L. R (CDRS) (19). Santo Touuis, 20 III 1995, P. Dentin (CDRS) (3d); 6 IV 1996. L. R (CDRS) (2<f); 23 IV 1996. L, 11 (CDRS) (Id); 23 IV 1996, (CDKS) (Id). V W[olf], 1200 m. Wrai« forest, 19 IV 1996 (CDRS) (7d 49), Playa Tortuga Negra, 22 VI 1998. L R (CDRS) (Id). PINTA: Abingdon, 24 VI 1899. [R. E, Snodgrass, E, Heller]. (CAS) (Id). PINZON: Plava escondida. 20 IV 2002. L. R & B. Landry (CDRS) (2d 39). RARIDA: Sendero tiirislico, 13 111 1998, L. R, (CDRS) (Id). SANTA CRUZ: Home-man farm 220m, 18 III 1964, (Id); 3 V 1964, (29), D. Q. Cavagnaro, (CAS). Puerto Ayora, 17 I 1989, R. L (CNC) (Id); 8 XI 1995, A. Herrera (CDRS) (Id); 29 X 1995, L R, V. Cm/. (CDRS) (Id), 4 km N Puerto Ayora. 20 1 1989, B I. (MECN) (Id), Tortug;i Reserve W Santa Rosa, 6 V 1989, S. B. Peck (CNC) (Id); 6 11 1989. B. Landry :CDRSHld). Cerro Croeker. 9 11 1996, L. R (CDRS) (Id), Bella-vista, 13 11 1999, reared from leaves D. ctmum, L. R (CDRS) (4d If); Punt a Nunez, 29 VII 2002, R. Boada (CDRS) (19). SAN CRISTOBAL; Chatam, X 1905, F, X. W, (CAS) (3d); 1 X 1906, Holotype, F. X. W (CAS) (Id); 23 V 1899, [R. E. Snodgrass, E. Heller], (CAS) (6d 19); 15 IV 1932. (3d); 17 IV 1932, (Id); 18 IV 1932, (2d). M, Willows, (CAS). Puerto Rh.|U<.™>, 17 II 1989, S. B. Petk (CNC) (Id), Airport 30 in, 6 III 1996, L. H (CDRS) ((>' 29). SANTIAGO: James, 21 IV 1899, [R. E. Snodgrass, E, Heller], (CAS) (29). Central 700 m,9 IV 1992, B. L (CNC) (Id).
Ilesperiinae
Cdpodcs ethlius (Stoll, 1782) "Canna Skipper"
(Fig. 2)
Ftipitiv ntMint Stoll 1782:212.
Cafyodes pf Wilis; Onore & Mielke 1988:619; Peek el al 1998:227.
Diagnosis. FW = 19.5-24 mm. The Canna Skipper is a large brown species with translucent spots on the; wings. Diagnostic features include a linear row of four spots on the HWand the absence of a tail. Specimens collected in the Galapagos are similar in shape and size to mainland specimens.
Geographical distribution. This species is widely distributed from Florida to Argentina including the West Indies. In the Galapagos Islands it has been reported previously from Isabel a (Onore & Mielke 1988) and here from Santa Cruz Island
Natural history. Calpndea ethlius is a powerful river that is most active at dawn and dusk, but is also seen at other times of the day. The species is most
abundant in urban areas and gardens but it has the ability to find isolated hosts inside the forests. Eggs are pale green and laid singly on the leaves of Canna edulis Kerr and Canna lutea Mill (Cannaecac), which have been introduced lo the Archipelago.
Flowers visited. Verbenaeeae: Cleradendrum molle HBK.
Flight period. Specimens have been collected in [■Yin-nary, April, May and July.
Remarks. Peck et al. (1998) listed this large skipper as recently introduced in the archipelago, and it is a known migrant (Ferguson et al. 1991).
Material studied. (5d and 19). ECUADOR, Cahipagos. IS-ABEI.A: VSN, 1 km W Santo Toinas, 12 V 1996, ex larvae, L. R (CDRS) (Id). SANTA CRUZ: Road to Baltra, 11 VI 1981, V. Lilian, (CDRS) (Id). Puerto Ayora, 24 VII 1996. (CDRS) (3d); 18 11 1998, (CDRS) (19); 22 IV 1998, L, R (CDRS) (Id)
Pier i dae
Coliadinae
Phoebis sennae marccHina (Cramer, 1779)
"Sulfur Butterlly"
(Figs. 7, S)
Pnpilio mtinTlliiia Cramer 1779; 103.
Callidryas eubele: Holland 18S9:19^; Williams 1911:296; Beebe
1923 "ill. Catopsila euhele; Van Duzee 1933:140.
fhtu-his \rntim-: I .iiisl<\ l\ l;siiiL;ri I IJMi. I oh. l'.u'kin <■( .il.
1972:103. Phoebis sennae mnrcellina: Moreno Ctal. 1997:146,
Diagnosis. FW = 24-36 nun. Easily recognized, it is the only yellow butterfly on the Galapagos Islands. Male: bright yellow. Female; yellow with black spots along the outer margins of the wings and a black spot with a pale center at the end of the forewing cell above. Galapagos populations are very variable in color pattern and size, but are identical with mainland specimens in genitalic features.
Geographical distribution. Pluwbis sennae marcel Una flies from Mexico to Uruguay (D'Ahrem 1981). In the Galapagos Islands the species has been previously reported from Baltra, Daphne, Eden, Femand-ina, Floreana, [sabela, Pinta, Santiago, San Cristobal, and Santa Cruz (Linsley & Usinger 196fi) and is here reported from Marcheua.
Natural history. One of the most common species on the Galapagos Islands. Adults occur in open habitats such as beaches, dry forests and road sides from sea level to 900 m. Both sexes have been observed vis-iiing flowers and puddles, The flight is fast and erratic. Beebe (1924) reported a southeast migration of this species from Santiago and Isabela to Santa (a u/„ and individuals have been observed commonly flying between islands. Williams (1911) reported Senna picta G. Don as a host plant in Galapagos. Females have
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been observed laying eggs in Senna tora L., Senna bi-capsularis L., and Senna occidentals L. (Fabaceae).
Flowers visited. Acanthaceae: J. galapagana; Apocynaceae: Catharanthus ruseus (L.) G. Don; Aster-aceae: Bidens pilosa L., Boraginaceae: C. leucophlyctis, C. hite.a, Heltotroptum angiospermum Murr., Tourne-fortia rufosericea Hook, f.; Cactaeeae: Opunlia insu-laris Stewart; Convolve! laceae: Ipomoea pes-caprae (L.) R. Br., Ipomoea nil ([..) Roth; Leguminosae: S. picta; Malvaceae: Gossypium barbadense van dartvinii (Watt.) Hutch, Hibiscus rosa-sinensis L., S. rfiomhifo-lia; Nyctagiuaceae: Bougainvillea spectabilis VVilld, Flumbaginaceae: P. scandens; Rubiaeeae: D. radula; Verbenaceae: L. carnara, Verbena Utoralix HBK.
Flight period. Multivoltine. Specimens have been seen throughout the year.
Material studied. (32d and I OS) ECUADOR. Galapagos. FERNAND1NA: Cabo Douglas, 15 IV 1999, P. Polo (CDRS) (1.1). ISABELA; VA, Pega-pega, 6 IV 1999, L. R[oque] (CDRS) (Id). Puerto Villamil, 2 III 1989, B. L[andryJ (CNQ(ld), Arid Zone, 7 III-1989, S. D. Peck (CNC) (Id). VSN, 6 IV 1996, L. R (CDRS) (So' and 2!). VSN, Santo Tomds 300 m, 26 IV 1996, L, R (CDRS) (8d). VSN, Pampas, 24 IV 1996, L. R (CDRS) (Id and 3?). VSN, La Bo-eanilla 1000 m, 28 [V 1996, L. R (CDRS) (2d). VSN, Alemania 350 in, 29 IV 1896,1. R (CDRS) (26). PlayaTortuga Negra. 22 V] 199S, L. R (CDRS) (36). MARCHENA: Playa Negra, 30 IV 2000, L. R, (CDRS) (Id). SAN CR1ST6BAL; Puerto Baquerizo, 17 II 1989, B. L (CDRS) (2d); El Potvetiir, 15 111 1996, L. R (CDRS) (Id1 and 59). SANTA CRUZ: Santa Cruz, 8 VIII 1990, C.K McMiillcn (CDRS) (Id). SANTUGO: Los Jaboncillos, XII 1974, (CDRS) (ld); Es-pmnilla, 17 III 1998, L. R (CDRS) (Id).
L.YCAENIDAE
Polymnmatinae
Leptotes parrhasioides (Wallengren 1860)
"Galapagos Blue Butterfly"
(Fig. 3)
Luceena parrhasioides Wallengren 1860:37:1861:355.
Cupula parrhasioides; Williams 1911:300, Beebe 1923:55; Van
Duzee 1933:143; Moreno etal. 1997:158, Leptoten parrhasioides; Bridges 1988:1 268, Linsley & Usinger
1966:157; Balinl & Jonhson 1995:9,
Diagnosis. This species and the following are the only lycaenid butterflies in the Galapagos Islands. Both arc similar, but the wing pattern and genitalia are distinctive. FW = 8-12.5 mm. Male: DFW and DHW violet blue with two or three black anal spots. VW ground color grayish with a pattern of white markings. VHW with three or four marginal spots, black in the middle, and filled with metallic blue. Female: similar to male but DW ground brown or blue.
Geographical distribution. Endemic to the Galapagos Islands, and reported from the islands of Baltra, EdeVi, Fspafiola, Fernandina, Floreana, Isabela, Pinzoii, San Cristobal, Santa Cmz, Santiago (Linsley & Usinger 1966). It is reported here from Marchena, Pinta and Rdbida.
Natural history. Common in the arid zone of the islands. The flight is slow and close to the ground, but may also he rapid and erratic. Both sexes have heen seen visiting (lowers and were often observed on muddy puddles early in the morning or after rainfall. The pale green eggs are laid singly on the flowers, young fruits, or young leaves of host plants. Williams (1911) reported Cardiospermum halieacabum L. as a host plant and indicated Cardiospermum gala-pageium B. L. Rob. & Greenm (Sapindaccae) as another probable food plant. New hostplants records include: P. seasides, Crotalaria incana L. and Rhijn-chosia minima (L.) DC, (Leguminosae). The larvae are variable, either green or red with a pattern of dark dorsal marks, depending on the host. The body is covered by short white hairs. Pupae are greenish with brown markings.
Flowers visited. Acanthaceae: /. galapagana; Asteraceae: Darwiniothamnus lancifolius (Hook, f.) Hailing; Boraginaceae: hi. cura.ssavicum, Tournefortia psiloslachya HBK, T. rufo-sericeac; Cueurbitaceae: Momordica charantia L.; Cyperaceae: Cyperus ligu-laris L.; Leguminosae: Vigna lute.ola (Jacq.) Benth.; I.inaceae: Linum cratericola Eliass; Lythraceae; Cuphea racemosa (L. f.) Spreng.; Malvaceae: S. rhombifolia, Plumhaginaceae; P. scandens; Nyctagi-naceae: Commicarpus tuberosits (Lam.) Standi.,-Solanaceac: L. cheesmanii; Zygopbyliaceae: Tribulus cistoides L
Hight period. The species is multivoltine.
Remarks. Wallengren (1860) described this endemic species from at least one male and one female [locality not specified) collected in May 1852 on the Galapagos. However, in 1861 Wallengren redescribed the species with additional material labelled "Puna" [gulf of Guayaquil] and "San Jose island" [Panama], Balint and Jonhson (1995) studied a specimen labeled "type" in the Naturhistoriska Riksmuseet, Stockholm, Sweden (NRS). They stated that this specimen belongs to the sister species Leptotes trigemtnatus (Butler) and that the specimens involved in the redescrip tion (Wallengren 1861) were incorrectly associated with the original scries from the Galapagos. The real type of L. parrhasioides was not examined, although it is probably located in NRS.
Material studied. (52d and 129). ECUADOR Galapagos. ES-PANOLA: Bahfa Maiiranillo, 25 IV 1992, B. 1. (CNC) (Id) FERNANDINA; NaiWough, 16 IV 1906, F X Wfllliamsj, (CDRS) (IS). Near to summit 4800ft. 5 111964. D. Q Cavagnaiu, (CAS) (3d). Cabo Douglas, 21 VI 1998, L. R[ocnie] and C. C[austonJ, (CDRS) (3d and 2?). Campamento Perez South Punta Espinoza, 18 VI 1998, L. R and C. C, (CDRS) (7d and 49). Cueva norte, 20 VI 1998, L. R and C. C, (CDRS) (44 and 29). North Side 300in, 12 I 2002, L. R and C. C, (CDRS) (19). Zona de vegetacion. 20 VI1996, L R, (CDRS) (4d and 39). FLOREANA: Charles, 10 X 1905, F X W. (CAS) (29); 23-31 V
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Journal of the Lepidopterists' Society
1906, F X W, (CAS) (IV); 1^ VI 1906, FXW, (CAS) (2d). Zona Costera, 2 IV 199.5, L R and \' Cruz, (CDRS) (Id). Scalesta forest, 4 III 1 995, I.. R ami V. Cruz (CDRS) (13dand49). Arid zone 130m, 4 III 1995, in //. ruraxsaviciiiji flowers {36): in C. mt)We flowers (Id), L. R and V. Cruz (CDRS). Road to the highland, 4 IK 1995, L, R and V. Cruz (CDRS) (3d*). Parte alta, 4 III 1995, L. R and V. Cmz (CDRS) (W). Ccrro Alien 340 m, 21 XII 1999, in Ltaum aratericola flowrs, L R, (CDRS) (Id). ISABELA: Albemarle Hanks Bay, 16 X 1906, F X W, (CAS) (2d). Albemarle Tagus Cove, 22 111 1906, F X W, (CAS) (16). Vjolcan] S|ierraJ \[egra] 1000 in, I Hi 19S9, H. I. (CNC) (I1?), V[olcan] W[olf], Scaksia forest 1700 m, 19 V 1996, L. R (CDRS) (W). VW, Upper arid imne, 1250 m, 20 iV 1996, 1, R (CDRS) (I3<f). Plays Tortuga Negra, 22 VI 1998, L. R, (CDRS) (8d and7J). P1NTA; Pkya Ibetson, 14 III 1992. B. L (CNC) (Id and 19); reared From leaves R minima, 14 III 1992, B. L (CNC) (HI 400 m, 17 111 1992, B. L (CNC) (2d). PINTA: Abingdon green zone, 10 IV 1906, FXW. (CAS) (IV). PINZON: Summit & upper ealdera areas, 7 II 1964, D. Q. Cavagnaro. (CAS) (Id). SANTIAGO: James bav low alt it ud. Vlll 1932, M Willows, (CAS) (1"). SANTA CRUZ: indefatigable, X VI 1932, M. Willows, (CAS) (4d and 19). Academy Hay CDRS, 5 11 1964, R. D. Schuster. (CAS) (Id); 25 II1964, D. Q. Cavagnaro and R. O, Sehuster, (CAS) (Id). Arid Zone, asotiated to Casiela galapn^eia. 19 I 19H9.B. L (CNC) (2W). Transition zone, 20 I 1989, B.L(CNC) (Id). SAN CRISTOBAL: Cliatam Saphocove, 9 I! I MOD. LWV ( \S)il ",.
Herniaryia ramon (Dognin, 1887)
"Ramon's Blue Butterlly"
(Figs. 4-6)
lAjcaena ramon Dognin I8N7:23.
Hmuierffix i union; lioque et al 1997:29; Peet et al. 1998:227
Diagnosis, FW = 8.2-11.6 mm. Resembles L. par-liiasiaidcs, with which it Hies, but H. ruwim lias two black dots in the VHW inner costal margin while L. pfirrlutsioidrs has none. Mali-: DW and 1)1 i\\ violet bine, the outer margins thinly bordered with white. Female: DVV and DHW dark with only a touch of blue at the wing base,
Geographical distribution. Distributed on the western slopes of the Andes from northern Chile to the south of Ecuador. This species was listed by Feck et al. (1998) as introduced in the Archipelago It has been reported from the Galapagos Islands of Baltra, Fspafiola, Floreana, Isahela, Sari Cristobal, Santa Cruz and Santiago (Roque et al. 1998) and is here reported from Fernandina, Pinzt'm, Santa Fc, Seymour Norte and Rabida.
Natural history. Common from the low arid zones to the highest elevations. Roque et al, (1997) reported seven leguminous plants as host of the species [Acacia macracantha Willd, S. tora, Neptunia plena Reoth, P. atrapurpiucns, Prosopis julijtora D.C., Tephrosta de-cumbens (Benth), and one species of Cucurbitaoeae: Cucumis dipsacetts Her. and one species of Oxali-daecae is reported here: OxaUs dombeyi A. St-Hil.
Flowers visited. Asteraceae: Scalesia gordilloi Ilamann & Wium Anders., ,S\ pedunculate, Boragi-naccac: T. mfo-sericeae; Leguminosae: T. decumbens.
Flight period. Observed during all mouths of the
year, but is most common during the rainy season (January to April).
Material studied. (42d and 129). ECUADOR Calapagos ESFANOLA: Hahi'a Ylaiy/amlb, 24 IV 1992. (Id); 25 IV 1999 (Id), 6. Landry. (CNC). FERNANDINA: Campainento Perez south Punta Espinoza, IS VI 1998, L. R[oque] and C. Cfauston] (CDRS) (Id and 23); Calx) Douglas. 21 VI 1998, L, R and C. C, (CDRS) (Id). Zona de vegetaci6n, 20 VI 1998, L. R and C. C, (CDRS) (19), FLOREANA; Zona Costera, 2 IV 1995. L. R and V CJruz] (CDRS) <8d and 5v). High arid zone 150in, 4 111 1995, I,. R and V. C (CDRS) (5d). Road to Highland, 4 III 1995, L. R and V. C. (CDRS) (lOd and 39). Parte media, 4 II) 1995, in He-lioiropium curassavicum flowers, L. R and V. C, (CDRS) (56). ISABELA: V[olean] S[ierraj N[«gra], 24 IV 1996. L. R (CDRS) (2d and 2S). V S N, lais Mercedttas I20m, 24 (V 199fi, L. R (CDRS) (19). V S N, 2 km W Puerto VUlamil, ,1 VI 200O, C. Covell (CDRS) (fid and 29), PINZON: Playa estondida. 20 VI 2002, L. K and B. L, (CDRS) (3d and 35). SANTA CRUZ: Arid Zone, S IX 1999, L It and C. Covell (CDRS) (29 and 4d); 19 I 1989. B. L, (CNC) (3d and 39).
Nymph am dae
Danainae
Danaus g&tppus tkersippus (Bates 1863)
"Queen Butterfly"
(Fig. 11)
Pajhlio tfiersippas Rates \HH3.
O/maus gilippm: Parkin et al. 1972:103; Liiusley 1977:29.
Danaus gilipptts fhcrsippus; Ackery & Vane-Wright 1984:208.
Diagnosis. FW - 3fi—41 mm. Fasily distinguished from D. plcxippus L. by not having the black inner border, and the black veins on the DFW, Males have a black androconial patch on the DHW. Females are similar to males but without an androconial patch.
Geographical distribution. Danaus "ilippus ther-sippun is distributed from the Southern United States to Panama and is native to the Galapagos Islands. The species has been reported from San Cristobal (Parkin et al, 1972), and is here reported from Baltra, Flore-ana, Isabela, Santa Cruz, Santiago, Seymour Norte and Rabida,
Natural history. The queen butterfly is primarily an inhabitant of the arid zones from sea level to 400 m, but its distribution is limited by the occurrence of its loodplants. Adults have been observed flying slowly and frequently visiting flowers in open areas The pale green eggs are laid singly on young and old leaves of the endemic vine Sarcoititnma angusthsima R, W. Holm. The other milkweed that occurs on the islands, Asclcpias curassavica L., apparently is used as an alternative food plant.
Flowers visited. Asclepiadaceae: A. curassavica, S. angustissima; Asteraceae: D. tenuifoliw; Boragi-naeeae: H. angiospermum; Solanaceae: P. pubescens; Vcrbenaceae: L. camara.
Flight period. All available records are From the rainy season.
Volume 58, Number 1
:!')
10
14
16
Fi«. 9-16. 9, Agraulis vanillae galapagensis (DW); 10, Agrtuilis vanitlae gatapagemis (VW); 11, Danau.i gilippu.1 thersippus (DW); 13, Panaus plexipptts megahppe (DW); 13, Vanessa virgfniensis (DW); 14, Vanessa virginienxis (VW); 15, Vanessa carye (DW); 16, Vanessa carye (VW).
40
Journal or the Lehdofterists' Society
Remarks. Ackery and Vane-Wright (1984) suggested that this species could have been introduced from California to the islands in recent times. Based on its current distribution and the well-established dispersal potential, and taking into consideration the dates of discovery of the butterfly and its host plants, it is reasonable to assume that D, g. thersippus arrived in the Archipelago through natural dispersal mechanisms and was not introduced by humans.
Material studied. (23d and 99) ECUADOR, Galapagos, FEK-NANDINA:, Cucva Norte. L. R(oque], (CDKS) (2d and 19). Cam-pamento Perez south Punta Espinnza, 18 VI 1998, L. R and C Causton, (CDRS) (l<t). FLOREANA: Cerro Pajas, 2t) IV 1996, L. R (CDRS) (19). Arid zone, near to Black beach, 22 IV 1996, L. R (CDRS) (2d). C. Cruz farm, 24 VI 1996, [.. R (CDRS) (46*). Asilo de la Paz 342m, 23IV 1996, (.. R (CDRS) (49 and <k<) 1SAHELA: Playa Tortuga Negra, 22 IV 1998, L R (CURS) (8d and 39). Vblean Siena Negra Alemanla 359 in. 29 IV 1996, L. R. (CDRS) (LOd and 69). Vol-can Sierra Negra pampas. 24 IV 1996, L. R, (CDRS) (19). SAN CRISTOBAL: Puerto Baijuerizo, 17 II 1989, B. L, (CDRS) (Id); (MECN)(ld).
DarutttM plexippits tnt:galippi' {Hiibner, [ 1826])
"The Monarch"
(Fig. 12)
Anosia megalippv H[ibner \ 1826J:2, pi. 7, fig. 2. Dan&isptextppus; Beebe 1923:55.
Dimmix /itnijijiiK, ljnslt*y& Usingcr 1966:157; Roque 1998a:9, b:10. Danaus pkxif)fiii.i megfilqipe; Ackery & Vane-Wright 198.1:201.
Din gnosis. F\V = 3,5-50 mm. Tins species is easily distinguished from D. gillppus by having the DFW apex boldly black, a black inner border in DFW and prominent black veins. Males have a black androconial patch on the DFW. females are similar to males but much darker,
Geographical distribution. The Monarch is native in the Galapagos Islands and reported from Flore-ana, San Cristobal and Santa Cruz. The status of this species has been discussed in some detail by Koqne (J99Sa). Dawn is plexlppits megallppe is distributed from Nicaragua to Peru including the North east of Brazil, Venezuela, French Guiana, Surinam, Guyana, Trinidad, Tobago and some islands of the West Indies (Ackery & Vane-Wright 1984).
Natural history. Common in open areas such as fields, roadsides and disturbed sites. In general, the localities where the species occurs are rnesic, but there are a few records in arid 7ones. Galapagos populations do not migrate. The adults fly slowly and visit flowers, females have been observed ovipositing on the milkweeds A. curassavica and S. angwstiaxhiw, but the latter is used as an alternative host plant.
Flowers visited. Asclepiadacea: A. curassavica; Malvaceae: H rosa-sinensis; Verbeuaeeae: /,. camura, V. li to rails.
Flight period. Adults commonly occur only in ihe
rainy season, but are observed sometimes in the dry season.
Materia] studied. (13d and 59). ECUADOR, Galapagos. FLO-RI'ANA Cerro Pajas, 20 IV 1996,1.. H'oque], (CDRS) (Id and K>. Arid zone, near to Black beach, 22 IV 1996. L. K, (CDRS) (4d and 49). Finca de C. Cruz, 24 VI1996, L. K (CDRS) (Id). Asilo de la Paz .342 in, 23 [V 1996, L. R, (CDRS) (**). SAN CRISTOBAL: Puerto BaqueriTO, 17 II 1889. R. Landry (CDRS) (Id); (CNC) (19): (MECN) (19). SANTA CRUZ: El cascajo, II 1999, L. R, (CDRS) (19). Salasaca, 22 VIII 2(KH), R. Oqnendo, (CDRS) (Id).
Helicon iinae
Agratitis vanlllae galapagensis (Holland 1889)
"Galapagos Fritillary"
(Figs. 9, 10)
Agrmdis iximllae gtifapagensis Holland 1889:194; Williams 1911:298; Bcebe 1923:55: Van Diraee 1933:141.
Diotie wmillae galtipagensis; Stichel 1938:92, Michener 1942:4; Linslev& Usinger 1966:156.
Diagnosis. FW = 19-29 mm. This species is distinguished by its silver markings on the VW The females are paler than males.
Geographical distribution. Endemic to the Galapagos islands. It has been collected on Baltra, Eden, Femandina, Floreana, Isabela, Pinta, San Cristobal, Santa Cruz, and Santiago (Linsley 6r I'singer 1966).
Natural history. This is a common species on the larger Galapagos islands. Adults have been observed in open areas and lorests from sea love! to the highlands. Beebe (1924) cited an interesting migration along the West Coast of Isabela Island, Adults usually fly slow, low to the ground and frequently have been seen visiting (lowers Williams (1911) mentioned species of the genus Passifiora I,, as food plants of this fritillary in the Archipelago, but he did not mention which. Ovi position was observed on Passifiora suberosa L. and Passifiora trtdacttjlites Hook, (f'assi-floraeeae). The eggs were laid singly on young leaves. The larvae are black with green longitudinal stripes. The head and body are covered with long black spines.
Flowers visited. Acanthaceae: Blechum brownci Juss, f, /. galapagmia- Asclepiadaceae: S. angustis-sima; Boraginaeeae: Crotort scouleri Hook, f.; Can-naceae: Commelina diffusa Burdm. (.,- Malvaceae: S, hederifolia Gav., S. panictdata L., S. sahiifolia Presl.; Rubiaeeae: D. radula; Rutaceae: C. limnti; Sterculi-aceae: Waltlwria ocata Gav.; Verbenaeeae: L camara, Stachytarpheta cayennensis (Rich.) M.Vahl.
Flight period. All specimens were collected in the rainy season hut individuals were observed during all months of the year except August and September.
Remarks. This species has been treated by several authors as a subspecies of the widely distributed American species Agraulls vanillas. Holland (1889) and Williams (1911) stated that the Galapagos taxon
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11
should be raised to specific status based on several differences in the wing pattern. Michener (1942) recognized the differences in the wing maculation and size, and mentioned some variation in the male geni-talic valvae. Specimens studied here were collected on several localities of the Archipelago.The male genitalia are generally similar to those of mainland specimens of A. vanillae but differ consistently in the valva. The apex of the valva is more slender and with fewer denticules in A. v. galapagewsis. The female genitalia are similar in both forms. Agraulis vanillae galapa-gensis can be separated from the other described subspecies by the smaller size, by the darker and more fuscous color of the basal half of the wings dorsally, by the stronger black markings on the wings, and by the absence of white dots in the black spots of the DFW discal cell. The holotype, collected on Chatham [San Crist6bal], is deposited in the National Museum of Natural History, Washington, DC (USNM). The identity of this species will remain questionable until a revision of the genus makes it more definite.
Material studied. (14d and 519). ECUADOR. CalApagos. FERNANDINA: Campamento Perez south Punta Espinoza, 18 VI IH9S, L. R[oque] tradC. Causton, (CJDRS) (Id). ELOREANA: Arid zone 130m, 17 IV 1996, L. R, (CDRS) (Id), Cerro Pajas, Scalesia forest, IS IV 1996, (yd and 59), 19 !V 1996. L. R, (19); 22 IV 19%. L. R (CDRS) (7d and 29). ISABELA: V[okan] S[ierra] N[egra], SantoToinas, 6 IV1996. (Id); 26IV 1996, L. R, (CDRS) (Id). V S N, bosqnedelosnino5,20II[ 1995, L R, (CDRS) (Id) VS \, pampas 1000m, 1 III 1989. B. L[andrv] (CNC) (19). V S N, la bocanilla 1000m, 28 IV 1996, L. R, (CDRS) (2d and 19). V S N, Las mercedi-las,3(lIV1996.1.. R, (CDRS) (2d and 49), Volcan Wolf, upper arid zone 1700 m, 19 V 1996, L. R, (CDRS) (13d and 4?). P1NTA. 400 m, 17 III 1992, B. L, (CNC) (4rf), SANTIAGO: 1'laya Espumiila, 4 IV 1992, B. L, (CNC) (Id). SANTA CRUZ; CDRS, 22 XI 1963, (CDRS)(Id). Tortugas reserve west of Santa Rosa, 6II 1989, B. L, (CDRS)(ld);(MECN)(ld).
Nymphalinae Vanessa oirginiensis (Drury 1773) "Painted Lady" (Figs. 13,14) Nymphatis cardui virginicmis Dnny 1773:1. Pyrantels fciinfem; Williams 1911:299. Vanessa vtrgmensis; Linsley & Usinger 1966:156.
Diagnosis. FW = 21.7-29 mm. Differs from V carye by the presence of two blue-centered ocular markings on the VHW. Males and females arc similar.
Geographical distribution. This cosmopolitan species was previously reported from Isabela (Volcanoes Darwin and Sierra Negra) (Williams 1911). On Isabela, the species was found at those localities as well as on volcanoes Alcedo, Cerro Azul and Wolf, Other new records include Santa Cruz and Santiago and it probably also inhabits other large islands. The species is distributed from southern Canada to the mountainous region of Colombia, including the West
Indies, Hawaii, Azores, Madeira, the Canary Islands, and occasionally Lurope (DeVries 1987),
Natural history. This species has <ml\ been observed in the open grassy areas of the highlands. All specimens were collected above 900 m in the pampa zone. Although it flies usually under bright and sunny conditions, we have seen it under overcast conditions and even during light rain (Volcan Alcedo, 23.X.1998). Adults fly fast and rest on the ground, flying short distances when disturbed. Eggs, larvae and pupae were found on the leaves of Cnaphalium purjnireum L. (Asteraceae) on the islands of Isabela and Santiago. The plant can be found on more islands (Lawesson et al. 1987). Vanessa virginietms has resident populations in the Galapagos Islands.
Flowers visited. Kubiaceae: Borreria laevis (Lam.) Griseb.
Flight period. Specimens have been collected or observed from January to June.
Material studied. (15d and 49). ISABELA: Albemarle, 20-31 VIII 1906, EX Williams, (CAS) (Id). Vfolean] S[ierra] N|egral, rim northwest side, 4 HI 1989, D. L[andiy], (CDRS) (Id); (MECN) (Id).
V S N, 1000 m, 4 111 1989, S. B. Peel: (CNC) (19). V S N, La Bo-canilk 1000 m, 28 IV 1996, L. Rloquej, (CDRS) (Id and 49). V S N, Las Mcrceditas, 6 IV 199b. L. H, (CDKS) (Id). Volcan Wolf, high arid zone. 1250 m, 21 V 1996, L. R, (CDRS) (4d), Volcan Alcedo, top 1100 m, 24 IV 1998, L. R (2d and 49). SANTA CRUZ: CDRS, 10
VI 1965, (CDRS) (Id). SANTIAGO; Los Jaboncilbs 900 m, 9 IV 1992, B. L. (CNC) (19); 16 111 1998, L. R, (CDRS) (Id), Los Agua-cates, 16 111 1998, L R (CDRS) (Id). NW slope 600 m, 30 V 1964, D. Q. Cavagnaro, (CAS) (19).
Vanessa carye Hiibner [1812]
"Western Painted Lady"
(Figs. 15, lb')
Hamadryas carye Hiibner [1812]:pl. 45, Jigs. 1, 2. Vyrameis caryae; Williams 1911:300. Vanessa caryae; Linsley & Usinger 1966:156.
Diagnosis. FW = 21-28 mm. Differs from V vir-giniensis by having four small blue-centered ocular markings on the VHW.
Geographical distribution. This species has been collected on San Cristobal, Fernandina, and recently ohserved by the author in Floreana. Vanessa carye is distributed in the Andes of Colombia, Ecuador, Peru, Bolivia, Chile, southwestern brazil, Paraguay, Uruguay, and Argentina. It also occurs on Juan Fernandez island, on Easter island, and the Tuainotu Archipelago.
Natural history. Three out of four specimens collected or observed were from the low arid zone at sea level. Two fresh adults were observed visiting flowers on Floreana (131 1995 and 6 II 1995, Puerto Velazco Ibarra) but were not collected. Their fresh condition suggested thai they were from a population extant on the island, mllier (liau miijranls. The husi daul and the immature stages are unknown in the Archipelago.
42
Journal of the Leptdoftemsts' Society
Members of Asteraeeae, Geraniaoeae, Malvaceae, and Urtieaceae arc reported as host plants for the species (Field 1971).
Flowers visited. Verhenaceae: C, moMe.
Flight period. Adults have been collected or ob-served in January and February.
Remarks. The only two specimens collected on the Galapagos were deposited iti the California Academy of Sciences.
Material studied. (Id and 1«). FERN AN DIN A: near to summit 4800 ft. 5 II 1984, D. Q. Cavagnaro, (CAS) (15). SAN CRISTOBAL: Chat am. Wreck Ray. 22 II J 906. (CAS) (M).
Ack N O W i ,!■; i x; vi e nts
I would like to thank the authorities of the Galapagos National Park and the Charles Darwin Research Station for financial and logistical support in field. The Charles Darwin Research Station and < iauacoll foundation pnivided financial suppnrl lot my investiga lions at tlie Canadian National Collection of Insects in 1999.1 thank the curators Cenriania Esteve* (early curator MECN, Quito, Ecuador). Giovanni Onore and Francisco Pinas (PUCE), Bernard Landry (MHNC, Geneva, Switzerland) for their permission to study the specimens deposited in their collections. I am indebted to Bernard Landry and Charles V Covell Jr. (University of I,ouiNvi]]e. Kentucky, USA) for generosity with their time and knowledge of l,epidoptera. and Stewart Peek (Carlcton University, Ottawa, Canada) for his knowledge of Galapagos insects. I am also greatly indebted to Charlotte Causton, Alan Tye, Heidi Sncll and Sol Rojas (CDRS), Olaf Mielke and Mima Casagrande (Univcrsidad Federal de Parana, Parana, Brazil), Philip J. DeVries and Carla M. Pen?. (Milwaukee Public Museum, Milwaukee, USA) and Keith Wilhnoll (NHM London) for their critical review of versions of this manuscript. To Iveui Aldaz (CDRS) for plant identifications, and '/.soil Halint (Hungarian Natural History Museum, Budapest, Hungary) for Lycaenidae literature, 'lb Malts Wedin (CDRS) for producing the plates. Finally I thank to all my field assistants during the years, especially Valentina Cruz.
This research wras partially support cd bv Uncsco project #894976.9: Ecological monitoring in the GalApagos Archipelago, Galapagos Conservation Fund, GEF-Natiira Project: Monitoring the Galapagos Islands and Darwin Initiative for the survival of species project: Terrestrial Invertebrate Biodiversity in Galapagos (Ecuador); Training and collection rehabilitation
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Received for publication 19 June 2002; revised and accepted 7 January 2004