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Redalyc.Is Cycasin in Eumaeus Minyas (Lepidoptera: Lycaenidae) A Interciencia ISSN: 0378-1844 [email protected] Asociación Interciencia Venezuela Castillo Guevara, Citlalli; Rico Gray, Víctor Is cycasin in eumaeus minyas (lepidoptera: lycaenidae) a predator deterrent? Interciencia, vol. 27, núm. 9, septiembre, 2002, pp. 465-470 Asociación Interciencia Caracas, Venezuela Available in: http://www.redalyc.org/articulo.oa?id=33907204 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative IS CYCASIN IN Eumaeus minyas (LEPIDOPTERA: LYCAENIDAE) A PREDATOR DETERRENT? CITLALLI CASTILLO-GUEVARA and VÍCTOR RICO-GRAY arvae exhibit a wide quences for the biology and ecology of and Clark, 1991; Nash et al., 1992; range of defensive strate- these species. Many “chemically de- DeVries, 1994). However, this has not gies to avoid being eaten, fended” Lepidoptera are aposematic, and been tested and the life cycle of E. minyas e.g., mimetic coloration, shelter construc- store plant compounds that are known remains undescribed. We present the re- tion, unpalatability due to urticating hairs, vertebrate toxins, such as cardenolides sults of laboratory and field experiments spines and defensive glands, regurgitation, (Brower, 1984), alkaloids (Rothschild et to evaluate the protective function of cyca- chemicals sequestered from host plants, se- al., 1979; Boppre, 1990; Montllor et al., sin in the aposematic butterfly E. minyas. cretion of volatiles, and noise production 1990), and cyanogens (Jones et al., In particular, the following questions are (see Brower, 1984; Bowers, 1993). Not 1962). Chemical defense of aposematic addressed: In which stages of its life cycle only do they use their bad taste or unpleas- insects has also been shown to be effec- does E. minyas contain cycasin? Does ant odor as a defense, they also announce it tive against invertebrate predators, which pure cycasin repel potential predators of to potential predators by means of a con- can learn to subsequently avoid similar E. minyas? Does cycasin turn eggs, larvae spicuous coloration, and gregarious and prey (Montllor and Bernays, 1993). and adults unpalatable? Is cycasin efficient sedentary behavior (Bowers, 1993). Seques- Cycasin is a secondary in protecting eggs and larvae from their tration of defense compounds from larval metabolite present in cycads (Cycadales), natural predators under natural conditions? host plants may require particular physi- belonging to the azoxyglycosid group This research is part of a wider study on ological adaptations by larvae to ingest, ac- (Matsumoto and Strong, 1963; Whiting, the interactions between E. minyas and E. cumulate, and store those compounds (Brat- 1963; Kobayashi and Matsumoto, 1965; debora and their host plants Z. loddigesii tsten, 1986; Bowers, 1992). Defense com- Morgan and Hoffman, 1983; Norstog and and D. edule. pounds are used for various purposes, par- Nicholls, 1997; Jones, 2000). Species of ticularly against predators, (Bowers, 1990; the American genera Zamia, Ceratozamia Materials and Methods Duffey, 1980; Blum, 1983; Brower, 1984). and Dioon (Cycadales: Zamiaceae) are Research on acquisition of chemical de- hosts to aposematic butterflies of the ge- Study site fenses by insects, particularly in Lepi- nus Eumaeus (Lepidoptera: Lycaenidae). It doptera, has been done using adult indi- has been demonstrated that E. atala se- Field work was accom- viduals. However, it is usually during the questers cycasin from Z. floridana, which plished in an oak forest near Chavarri- larval stage that chemical defenses, seques- is later used as a defense against both ver- llo, in central Veracruz, México (19º24’N, tered from host plants, are ingested, pro- tebrate (Bowers and Farley, 1990) and in- 96º48’W; 1000m altitude), characterized by cessed and stored (Bowers, 1993). Ex- vertebrate (Rothschild et al., 1986; Bowers a calcareous-derived soil and abundant rock amples of chemical defense of Lepidoptera and Larin, 1989) predators. These results outcrops. The climate is temperate-humid, larvae are well known (Brower, 1984; Bow- were based on laboratory experiments, but mean annual temperature is 24.5C°, total ers, 1990; Witz, 1990). no field research has confirmed them. It annual precipitation is ca. 1110mm, with a In aposematic species has also been suggested that cycasin rainy season between June and September unpalatability is coupled with a warning works in E. minyas as a defense mecha- and an extended six-month dry season coloration which can have many conse- nism, providing chemical protection (Clark (García, 1964; Soto et al., 1996). The veg- KEYWORDS / Aposematic Butterfly / Cycads / Deterrent Effect of Cycasin / Lycaenidae / Predation / Received: 03/01/2002. Modified: 07/30/2002. Accepted: 08/08/2002 Citlalli Castillo-Guevara. Ph.D. in Ecology and Management of Natural Resources. Address: Departamento de Ecología Vegetal, Instituto de Ecología, A.C., Apdo. 63, Xalapa, Veracruz 91070, México. e-mail: [email protected] Víctor Rico-Gray. Ph.D., Tulane University. Senior Research Scientist, Departamento de Ecología Vegetal, Instituto de Ecología, A.C. e-mail: [email protected] SEP 2002, VOL. 27 Nº 9 0378-1844/02/09/465-06 $ 3.00/0 465 etation is characterized (Flores, 1995) by a dae, Fabricius 1804) as predators. This 0.075, p= 0.7877) were found in the num- mixture of oak forest [Quercus oleoides, Q. ant inhabits the study site and was ob- ber of visits between solutions (control, laurina, Q. peduncularis (Fagaceae), Nec- served on Z. loddigesii. Before the ex- X= 84.375 +5.123, N= 675; experimental, tandra sanguinea (Lauraceae), Bursera si- periments, ants were collected from a X= 86.875 +6.465, N= 695). maruba (Burseraceae)] and palm groves of colony in Francisco Javier Clavijero Bo- To test the defensive Brahea dulcis (Arecaceae). Epiphytes are tanical Garden (Xalapa, Veracruz, Méxi- function of cycasin under natural condi- present in the Araceae, Bromeliaceae, Or- co; 19º30’N, 96º57’W; 1280m altitude). tions, a predator exclusion experiment was chidaceae and Cactaceae (Castillo, 1985). The sample included workers, soldiers conducted at the study site. Egg clusters of The main shurbs and herbs are (Flores, and larvae in order to ensure that ant E. minyas were located on individuals of Z. 1995) Acacia cornigera, A. pennatula (Mi- behaviour was the least affected by sam- loddigesii and three predator exclusion mosaceae), Zamia loddigesii, and Dioon pling, (Jorge Valenzuela-González, per- treatments were applied: 1) fronds covered edule (Zamiaceae). sonal communication). Ants were placed with mesh, 2) fronds with a band of in 25 x 13cm plastic containers covered Tanglefoot® at the base, and 3) fronds with Species with mesh, and were deprived of food Tanglefoot® and mesh; the control were 48h before the experiments. The contain- fronds with egg clusters under natural con- Zamia loddigesii (Zami- ers were then uncovered and placed ditions. Different treatments were used to aceae, Miquel 1843) (voucher, L. M. White- within a square (0.26m2) of Tanglefoot® determine differential effects of the differ- lock 11/26/1963, XAL) is a small plant (up (Tanglefoot Co., Grand Rapids, MI, ent groups of predators (mainly for birds to 1m tall) with 1 to 6 fronds, inhabiting USA), where different food solutions and ants, but other flying and crawling in- tropical dry and deciduous forests and sec- were offered to the ants (see below). sects as well). Eighty-three eggs (total for ondary vegetation (Vovides et al., 1983). It To test if pure cycasin the study site at the time) grouped in egg is distributed along the coast of the Gulf of deterred S. geminata individuals, ants clusters located on 15 Z. loddigesii indi- Mexico up to Guatemala, from 0 to 1000m were offered two solutions, 1) control viduals were used, 32 on control plants and in elevation (Vovides et al., 1983; Jones, (0.75g sucrose + 0.30ml water), and 2) 51 on treatment plants (16 with mesh, 19 2000). Z. loddigesii is protected under the experimental (0.75 g sucrose + 0.30ml of with Tanglefoot®, 16 with Tanglefoot® and status of threatened species both interna- 1mg/ml cycasin in water). Cycasin was mesh). Over 15 days (the time for egg tionally (CITES Appendix II) and nation- the same as that used for chemical analy- hatching), the plants were visited every day ally (Annonymous, 1994). sis. The number of ant visits to each so- and the number of eggs preyed upon, Eumaeus minyas (Lycae- lution was recorded during 90 min. When hatched and unhatched was registered. The nidae, Hübner 1809) is distributed from an ant lowered its head and touched a so- exclusion experiment was continued apply- México to Costa Rica, where the larvae lution, it was considered a visit. The ex- ing the same treatments to the recently have been reported to consume fronds and periment was replicated eight times with hatched larvae, plus 15 more first-instar lar- female reproductive cones from Z. fur- different ants in the same conditions vae collected elsewhere in the study site. A furacea, Z. skinnery and Z. loddigesii (De- mentioned above. total of 70 larvae, 26 on control plants and Vries, 1976; 1983a; Clark and Clark, To test whether cycasin 44 on treatment plants (19 with mesh, 15 1991). E. minyas is considered aposematic, in eggs, larvae and adults rendered them with Tanglefoot®, 10 with Tanglefoot® and exhibiting a flashy warning coloration unpalatable and deterred S. geminata in- mesh) were observed. The number of lar- (DeVries, 1977; Clark and Clark, 1991; dividuals, individuals of E. minyas were vae that had disappeared, died or still sur- Nash et al., 1992). collected in the study site: 33 eggs vived was counted daily. To establish the (0.02g), 15 larvae of the four instars larval instar, larval length every third day Chemical analyses (0.70g), and 2 adults (0.20g). Extracts until pupation (ca. 16 days) was measured.
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