Athis delecta ( Schaus, 1911 )

García-Díaz, José De Jesús, Ramírez-Ponce, Andrés, López-Godínez, Bernardo & Torres-Yañez, Elier, 2025, Flying in the canopy: taxonomic and ecological analysis with geometric morphometrics and distributional notes on Athis delecta and Athis jaliscana (Lepidoptera: Castniidae), Zootaxa 5666 (1), pp. 74-92 : 78-82

Athis delecta ( Schaus, 1911)

Biogeographical and distributional comments. After Escalantiana escalantei (Miller, 1976) , Athis delecta and Athis inca orizabensis ( Strand, 1913) , is one of the Castniidae species with the northernmost distribution in the Americas, occurring in Cañón de la Peregrina, Victoria , Tamaulipas, Mexico ( Niño-Maldonado et al. 2013). Athis delecta appears to have a broad distribution, with records from Mexico, Guatemala, Honduras, and Costa Rica ( Schaus 1911; González & Hernández-Baz 2012; Morales-Morales et al. 2015; iNaturalist 2025; González 2024; García-Díaz et al. 2024), while its presence remains unconfirmed in Belize, El Salvador, Nicaragua, Panama, and South America. Based on the examined material, the species inhabits evergreen and semi-evergreen forests, as well as montane cloud forests, at elevations ranging from 0 to 2,300 m a.s.l., with most records occurring between 200 and 900 m a.s.l. Adults have been recorded from March to September.

González & Hernández-Baz (2012) reported that the Arthropod Collection of the Universidad del Valle de Guatemala ( UVGC) contained a right hindwing with a frenulum from a male Castniidae specimen collected in Izabal, Livingston, Biotopo Chocón Machacas (25-III-2001, 24 m a.s.l.), which they tentatively identified as Athis delecta , pending further revision. However, Jiichiro Yoshimoto (pers. comm., 2024) informed us that this wing is no longer present in the UVGC, preventing us from verifying that identification.

Athis delecta has been recorded from the following localities: COSTA RICA: Esperanza del Guarco, San José, Turrialba; GUATEMALA: Livingston; HONDURAS: Lago de Yojoa , Santa Rosa de Copán; MEXICO: Campeche: Calakmul ; Chiapas: El Aguacero, Emilio Rabasa ; Guerrero: Acahuizotla, Tierra Colorada ; Oaxaca: San José Chiltepec; Puebla: San Lorenzo, Santiago Yancuitlalpan, Telolotla ; Tamaulipas: Balneario La Florida, Cañón La Peregrina , San José; Veracruz: Camarillo, Cerro El Vigía, Córdoba, Fortín de las Flores, Peñuela, Presidio, Tapalapan, Tuxpan ( Fig. 5 ). According to the biogeographic provinces of the Neotropical region proposed by Morrone et al. (2022), these localities belong to the Chiapas Highlands, Sierra Madre Oriental and Sierra Madre del Sur provinces of the Mexican Transition Zone, and also to the Mosquito, Pacific Lowlands , Yucatan Peninsula and Veracruzan provinces of the Mesoamerican dominion, and to the Guatuso-Talamanca and Puntarenas-Chiriqui provinces of the Pacific dominion, both in the Brazilian subregion .

Ecology and behavior. Depending on the locality, Athis delecta can be sympatric with Athis inca inca (Walker, 1854) , Athis inca orizabensis , Divana diva diva (Butler, 1870) , Telchin atymnius futilis (Walker, 1856) , and/or Telchin evalthe viryi (Boisduval, [1875]) along the Gulf of Mexico slope and in Central America. However, there are four atypical records of this species from the Pacific slope of Mexico, specifically in Guerrero (see discussion below) ( Fig. 5 ).

According to the information collected by ETY in San Lorenzo, Xicotepec (Sierra Norte de Puebla) ( Figs. 2B, 2C ) since 2016, the emergence of Athis delecta adults is influenced by rainfall. The first individuals are consistently observed during the last two weeks of June, following two or three rainy days. The species is closely associated with woody trees such as Spanish cedar ( Cedrela odorata L.: Meliaceae ), pink cedar ( Acrocarpus fraxinifolius Arn. : Fabaceae ), kapok tree ( Ceiba pentandra (L.) Gaertn.: Malvaceae ), and gumbo-limbo ( Bursera simaruba (L.) Sarg.: Burseraceae ). Females only oviposit on individuals of their host plants, Tillandsia polystachia (L.) L. and Tillandsia schiedeana Steud. ( Bromeliaceae ) ( Figs. 2E, 2F ), which grow on these trees. Unlike most Mexican Castniidae , Athis delecta inhabits a distinct vertical stratum: males fly between 4 and 10 m above the ground, with an average height of 5 m, while females fly between 4 and 15 m, with an average height of 7 m above the ground. Both males and females fly on sunny or partly cloudy days; they do not fly on cloudy, rainy, or drizzly days.

Males exhibit fast, straight flight, but when the temperature drops and the day is partly cloudy, they fly in an erratic zigzag pattern, moving up and down. Their activity begins around midday when the temperature exceeds 32°C, reaching up to 37°C. They perch on dry branches of Spanish cedar, pink cedar, kapok tree, or gumbo-limbo at a height of 6 to 8 m above the ground, resembling a dry leaf ( Fig. 2D ). Like other Mexican Athis , when perched, they cover a large portion (or the entirety) of their hindwings in a stegopterous position ( Miller 1986; Ríos & González 2011; Vinciguerra et al. 2011; García-Díaz et al. 2020; González et al. 2021; García-Díaz 2022a, 2022b, 2023; García-Díaz & Turrent-Carriles 2022). Their flight territory spans approximately 8 m ², and they are observed more frequently than females, with an average of three males and one female seen per day (females are often not observed). Males begin to fly when another male invades their territory or when an orange butterfly of the Nymphalidae family, such as Marpesia petreus (Cramer, 1776) or Temenis laothoe (Cramer, 1777) , enters their territory. When this happens, males of Athis delecta approach these butterflies to determine if they are a female. However, once the identification becomes evident, they move away from the butterflies without chasing or fighting. They then perform one or two patrol laps within their flight zone before returning to the same branch (or a nearby one) where they were originally perched, landing 20 to 30 cm from the tip. Afterward, they slowly walk toward the tip, with their middle pair of legs moving rapidly while their wings slowly flap up and down, resembling a dry leaf shaken by the wind. Once they reach the tip and are not disturbed, they stop moving their wings and legs, remaining perched for a while. When one male invades another’s territory, both engage in confrontations, ascending in circles up to 20 meters above the ground. They repeat this behavior several times until one male succeeds in expelling the other from the area. On average, three to four similar confrontations are observed per hour. Males of Athis delecta have not been observed engaging in territorial fights with butterflies of any family. Males can remain perched in their roosting zone for up to an hour if no Marpesia petreus , Temenis laothoe , or other male invades their territory, as they do not take flight if another butterfly species approaches their perch. On very rare occasions, males fly up to 2 to 3 m. Male activity ceases around 14:30 when the temperature ranges between 34 and 37°C.

Females begin flying at midday, usually around 13:30, when the temperature exceeds 33°C. Their flight is slow, heavy, and erratic, moving up and down. Females have never been observed perching, as they are continuously searching for suitable Tillandsia polystachia and T. schiedeana plants for oviposition. They can descend as low as 2 m or ascend up to 15 m above the ground if a potential host plant is present. When a female selects a Tillandsia , she hovers around the plant for approximately 15 seconds to determine the optimal oviposition site, usually at the base of the bromeliad. Upon landing, the female turns around and walks backward before laying a single egg per Tillandsia . After oviposition, she walks to the tip of the leaf and takes flight in search of another bromeliad. Female activity ceases around 15:30.

There are no confirmed predators of Athis delecta , however, some bird species, such as the Great Kiskadee ( Pitangus sulphuratus (Linnaeus, 1766) : Tyrannidae ) and the Social Flycatcher ( Myiozetetes similis (Spix, 1825) : Tyrannidae ), might be potential predators as they are known to attack other lepidopterans.

There are no records of males or females of this species feeding on nectar, mud or fermented fruit. On one occasion, ETY observed a male A. delecta perching on a flower at a height of 2 m, but it was not possible to determine whether it was feeding. No courtship or copulation behaviors have been documented for this species.

Type material. COSTA RICA: Cartago: 1♂, Syntype, Esperanza, May, Type No. 12166 U.S. N.M., Catalog Number USNMENT-1244444 ( NMNH) ; MEXICO: Veracruz: 1♀, Syntype, Castnia delecta Type ♀ Schaus, Cordoba Mex., May, Collection W m Schaus, USNMENT01244445 ( NMNH) .

Additional material examined. Data from 39 specimens (16♂♂, 23♀♀) were recorded: COSTA RICA: Cartago : 1♂, Costa Rica, Cartago, Turrialba , 24-V-1972 H.L. King, Castnia delecta ♂, UF FLMNH MGCL 1138228 ( MGCL) ; San José: 1♂, San José, Museo Nacional de C. R., 4 de mayo 2001, Joaquín Sanchez, MNCR- E 55746 , MNCR-A5055746 ( MNCR) ; 1♀, San José, Museo Nacional de C. R., 8 de mayo 2001, Ernesto Carman, MNCR-E 55745, MNCR-A5055745 ( MNCR) ; MEXICO: 1♀, FSCA/ Florida State Collection /of Arthropods , PROJECT PHOTO/ J. B. Heppner /5997, MGCL/FLMNH/specimen no./48605 ( MGCL) ; 1♀, Mexico, no data ( MB) ; Campeche: 1♀, Calakmul, Camino a Calakmul , 24/III/1997, LN-00344 (ECO-CH-LN); Chiapas : 1♂, Ocozocoautla , El Aguacero, 20-VI-1992, leg. M. G. López Vásquez ( UNACH) ; 1♀, Emilio Rabasa , El Ocote, 02-VI-2021, leg. J. de la Maza E. ( CDM) ; Guerrero: 1♂, Acahuizotla Gro. , 15-V-1975, leg. A. Ibarra, CNIN LEP (México) 6460 ( CNIN) ; 1♂, MEXICO: GUERRERO /nr. Acahuizotla / viii.1983 / T. Escalante, MGCL/FLMNH/ specimen no./43342 ( MGCL) ; 1♂, MEXICO: GUERRERO / Tierra Colorado / viii.1983 /T. Escalante, Allyn Museum Photo / No. 850 827-11/12, MGCL/FLMNH/specimen no./43339 ( MGCL) ; 1♀, MEXICO: GUERRERO / Tierra Colorado / ix.1983 /T. Escalante, Allyn Museum / Acc. 1984-2, MGCL/FLMNH/specimen no./43341 ( MGCL) ; Oaxaca: 1♀, Chiltepec Oax. , VII-1970, leg. A. Díaz F., Ex-Colección Saldaña, CNIN LEP (México) 6463 ( CNIN) ; 1♀, Chiltepec Oax. , VII-1970, leg. A. Díaz F., Ex-Colección Saldaña, CNIN LEP (México) 6466 ( CNIN) ; 1♀, Chiltepec Oax. , VIII-1978, leg. A. Díaz F., Ex-Colección Saldaña, CNIN LEP (México) 6464 ( CNIN) ; 1♀, Sierra de Juárez OAX, 600m, V-2003, Coll. Thierry Porion ( DC) ; Puebla: 1♂, Zihuateutla, Telolotla, El Cajón, 23-VII-2022, leg. E. Yañez, José de Jesús García-Díaz genitalia dissection No. JJGD0006 ( JJGD) ; 1♂, Xicotepec, San Lorenzo , 21-VII-2023, leg. E. Yañez ( JJGD) ; 1♂, Xicotepec, San Lorenzo , 10-VII-2023, leg. E. Yañez ( CFT) ; 1♂, Xicotepec, San Lorenzo , 19-VI-2023, leg. E. Yañez ( RW) ; 1♀, Cuetzalan del Progreso, Santiago Yancuitlalpan , 02- V-2001, leg. F. Mora (CFV) ; 1♀, Athis ahala, Mexico-Puebla, Villa de Juarez, Escalantongo (MS) ; Tamaulipas : 1♀, “La Florida”, Gómez Farías, Tamaulipas, abril de 2003, Colector: Jesús García Jiménez ( RW) ; 1♀, “San José”, Gómez Farías , Tamaulipas, 22 de Agosto de 2009 ( RW) ; Veracruz: 1♀, Fortín de las Flores , Puente de Metlac, 05-VI-1968, leg. R. F. de la Maza R. ( JJGD) ; 1♀, Fortín de las Flores , Puente de Metlac, 22-VII-1967, leg. R. F. de la Maza R. ( CDM) ; 1♀, Fortín de las Flores , Puente de Metlac, 24-V-1996, leg. F. G. Haghenbeck F. ( CFH) ; 1♂, Santiago Tuxtla , El Vigía, 03-VI-2002, leg. F. G. Haghenbeck F. ( CFH) ; 1♀, Santiago Tuxtla , El Vigía, 15-IX-1998, leg. F. G. Haghenbeck F. ( CFH) ; 1♂, Mexico, Veracruz, Camarillo , 24 km SW Conejos, VII-13 16-[19]74, leg. J. A. Chemsak & J. Powell, UC Berkeley EMEC1330794 ( EMEC) ; 1♂, Tapalapa [n] Ver. , VIII-1974, leg. A. Díaz F., Ex-Colección Saldaña, CNIN LEP (México) 6461 ( CNIN) ; 1♂, Tapalapa [n] Ver. , VIII-1975, leg. A. Díaz F., Ex-Colección Saldaña, CNIN LEP (México) 6459 ( CNIN) ; 1♀, Peñuela Ver. , VII-1964, leg. A. Díaz F., Ex-Colección Saldaña, CNIN LEP (México) 6462 ( CNIN) ; 1♀, Sierra Santa Martha, Los Tuxtlas Ver. , 20-VI-1982, leg. A. Díaz F., CNIN LEP (México) 6455 ( CNIN) ; 1♀, Santiago Tuxtla , El Vigía, 06-VI-2022 ( BLG) ; 1♂, T. Escalante /PRESIDIO/ VER./V-42, A. C. Allyn / Acc. 1973-48, Slide No. M-2716 / ♂ Castnia delecta Schaus / Jacqueline Y. Miller , MGCL/ FLMNH/specimen no./43337 ( MGCL) ; 1♀, Castnia ahala [sic], T. Escalante / Presidio / Ver. /V-51, A. C. Allyn / Acc. 1973-48, Slide No. M-2717 / ♀ Castnia / Jacqueline Y. Miller , MGCL/FLMNH/specimen no./43340 ( MGCL) ; 1♀, COL. PEREZ H./ Santiago Tuxtla / Ver. 9-52, A. C. Allyn / Acc. 1973-48, Slide No. M-6887/legs, Ant / Jacqueline Y. Miller, MGCL/FLMNH/specimen no./43359 ( MGCL) ; 1♂, Veracruz, Presidio , Sept.1949, Ex: Escalante ( MS) .

Male genitalia. One specimen dissected ( Fig. 6A ). Tegumen broad, subtriangular in lateral view and subtrapezoidal in dorsal view, slightly narrower in its posterior portion. Uncus simple, short, fused into a pointed apex, and curved ventrally. Gnathos excavate posteriorly and fused anteriorly, with the ventral arms short and more sclerotized than the dorsal arms. Valva broad, oval, rounded, strongly sclerotized, and elongated anterodistally; costal margin straight, ventral margin straight in the basal portion, apex rounded and slightly curved inward. Sacculus well-developed, arising from the inner surface of the valva. Saccus well-developed, narrow, and curved anteriorly. Aedeagus strongly sclerotized, elongated, and recurved anteroventrally, with the distal portion narrower than the basal portion; distal lateral carina forming a flap without spines; coecum highly developed, swollen, and longer than the wider diameter of the ejaculatory bulb foramen.

Variability and sexual dimorphism. This species exhibits minimal variation in both dorsal and ventral views for both sexes. In the forewings, the shape and size of the discal spot, which includes the accessory radial cell, are variable (circular or oval), as is the thickness of the diagonal band extending from the subapical region to the anal margin. In the hindwings, the size of the postdiscal spots is variable, as is the length of the marginal extensions toward the postdiscal band ( Fig. 1 ).

Females are characterized by having forewings slightly more rounded than males, with a distinctly curved apical region. In the hindwings, the anal margin is also rounded in females. Regarding the wing pattern, dorsally, males generally have a darker base coloration in both the forewings and hindwings. The discal spot on the forewings is more elongated and oval in females. In the hindwings, the spots forming the postdiscal band are approximately twice the size in females compared to males. Additionally, both the inner margin and the extensions toward the postdiscal band are black in females and orange in males. Most of these differences are also noticeable in the ventral view ( Fig. 1 ). Overall, females are larger than males.

Figures

FIGURE 1. Dorsal (left wings) and ventral (right wings) variation of Athis delecta males (A–D) and females (E–H) from Mexico. A: ♂, Guerrero, Acahuizotla, 15-V-1975, leg. A. Ibarra, CNIN LEP (México) 6460 (CNIN); B: ♂, Puebla, Xicotepec, San Lorenzo, 21-VII-2023, leg. E. Yañez (JJGD); C: ♂, Veracruz, Tapalapan, VIII-1975, leg. A. Díaz F., CNIN LEP (México) 6459 (CNIN); D: ♂, Veracruz, Tapalapan, VIII-1974, leg. A. Díaz F., CNIN LEP (México) 6461 (CNIN); E: ♀, Oaxaca, Chiltepec, VII-1970, leg.A. Díaz F., CNIN LEP (México) 6463 (CNIN); F:) ♀, Veracruz, Fortín de las Flores, Puente de Metlac, 05-VI-1968, leg. R. F. de la Maza R. (JJGD); G: ♀, Oaxaca, Chiltepec, VIII-1978, leg. A. Díaz F., CNIN LEP (México) 6464 (CNIN); H: ♀, Veracruz, Sierra Santa Martha, Los Tuxtlas, 20-VI-1982, leg. A. Díaz F., CNIN LEP (México) 6455 (CNIN). Scale bar = 2 cm.

FIGURE 2. Aspects of Athis delecta ecology. A: Female perching in stegopterous position in a butterfly house of the Azul Meámber National Park, Honduras (24-VII-2022, photo: Karla Daniela Fiallos Yanez); B–C: general view of San Lorenzo, Xicotepec, Puebla (07-VII-2024, photo: ETY); D: male perching in stegopterous position in a tree at 7 m height in a ravine of San Lorenzo, Xicotepec, Puebla (05-VI-2025, photo: ETY); E: Tillandsia polystachia, host plant of A. delecta in San Lorenzo, Xicotepec, Puebla (21-IV-2024, photo: ETY); F: Tillandsia schiedeana, host plant of A. delecta in San Lorenzo, Xicotepec, Puebla (05-VI-2025, photo: ETY).

FIGURE 5. Geographic distribution of Athis delecta (red triangles) and A. jaliscana (blue circles).

FIGURE 6. Male genitalia of Athis delecta (A) and A. jaliscana (B–D). A: ♂, Zihuateutla, Telolotla, El Cajón, 23-VII-2022, leg. E.Yañez, José de Jesús García-Díaz genitalia dissection No. JJGD0006 (JJGD); B: ♂, Tuxcacuesco, Tuxcacuesco, 16-VII-2018, leg. B. López G., José de Jesús García-Díaz genitalia dissection No. JJGD0007 (JJGD); C: ♂, Tuxcacuesco, Tuxcacuesco, 03- VII-2022, leg. B. López G. (BLG), José de Jesús García-Díaz genitalia dissection No. JJGD0008 (JJGD); D: ♂, Tuxcacuesco, Tuxcacuesco, 03-VII-2022, leg. B. López G. (BLG), José de Jesús García-Díaz genitalia dissection No. JJGD0009 (JJGD). Left: genital capsule; middle: dorsal view of uncus and tegumen; right: aedeagus. Scale bar = 0.5mm.