Flying in the canopy: taxonomic and ecological analysis with geometric morphometrics and distributional notes on Athis delecta and Athis jaliscana (Lepidoptera: Castniidae) Author García-Díaz, José De Jesús 0000-0002-3559-8204 Posgrado en Ciencias, Instituto de Ecología A. C., Red de Biodiversidad y Sistemática, Carretera antigua a Coatepec 351, El Haya, 91073 Xalapa, Veracruz, Mexico; Sociedad Mexicana de Lepidopterología A. C. mexicastnia@hotmail.com Author Ramírez-Ponce, Andrés 0000-0003-4742-7397 Instituto de Ecología A. C., Red de Biodiversidad y Sistemática, Carretera antigua a Coatepec 351, El Haya, 91073 Xalapa, Veracruz, Mexico andres.ramirez@inecol.mx Author López-Godínez, Bernardo 0000-0001-9832-3921 Sociedad Mexicana de Lepidopterología A. C. berlopezg@yahoo.com.mx Author Torres-Yañez, Elier 0009-0005-5424-8508 Adolfo López Mateos, 222 B, San Isidro, 73080 Xicotepec, Puebla, Mexico torreselier161@gmail.com text Zootaxa 2025 2025-07-25 5666 1 74 92 http://dx.doi.org/10.11646/zootaxa.5360.3.8 journal article 311597 10.11646/zootaxa.5666.1.3 1f49e4eb-3262-4cab-8cea-2d76946524b2 1175-5326 16609960 80227F02-3BAA-4C84-A664-212517783753 Athis jaliscana López-Godínez & Porion, 2012 Biogeographical and distributional comments. As Athis miastagma (Dyar, 1925) , A. flavimaculata (Miller, 1972) , Escalantiana escalantei and Mexicastnia estherae (Miller, 1976) , this species is restricted to the Pacific slope of Mexico in localities with the presence of tropical deciduous forest, between 0 and 1265 m a.s.l. It has been recorded in Colima, Guerrero, Jalisco and Michoacán ( García-Díaz et al. 2019), between April and August. To date, the northernmost locality for the species is Tuxcacuesco , Jalisco . The following is a list of localities where Athis jaliscana has been sighted and collected: Colima : Comala , Tamala ; Guerrero : San Roque ; Jalisco : Chamela , Tolimán , Tuxcacuesco ; Michoacán : Playa Azul (García-Díaz et al. 2019) ( Fig. 5 ). These localities belong to the Sierra Madre del Sur province of the Mexican Transition Zone, and to the Pacific Lowlands province of the Mesoamerican dominion in the Brazilian subregion ( Morrone et al. 2022 ) . FIGURE 5. Geographic distribution of Athis delecta (red triangles) and A. jaliscana (blue circles). Ecology and behavior. This species is sympatric mainly with A. flavimaculata and infrequently with A. miastagma gonzalezi López-Godínez, García-Díaz & Turrent-Carriles, 2022 or E. escalantei in places with an abundance of Tillandsia spp. ( Bromeliaceae ). Its usual flight period extends from mid-June to early August; however, there is an April 18 record from Comala, Colima . The adults’ activity is closely linked to the rainy season as the first individuals eclose 4-7 days after a week of rains. Males eclose some 5-8 days before females. In Jalisco , the first two weeks of July is when the greatest number of individuals could be observed per day (between 2- 4 males , and 1 female at most). The habits of A. jaliscana and A. delecta are unique and differ greatly from those exhibited by species of the inca and hechtiae groups, as they fly in the forest canopy. This could explain their scant representation in collections. Athis jaliscana often flies on warm sunny days; it is not active on cloudy, windy or cold days, nor in rain or drizzle. On a few occasions it has been seen flying on cloudy days; however, on those days the clouds were high, and conditions were warm and with high luminosity. Like A. flavimaculata , it flies in undisturbed places, mainly along trails and dirt roads on flat terrain with high Tillandsia grossispicata Espejo, López-Ferrari & W. Till, 2008 ( Bromeliaceae ) density, which is its host plant in Jalisco and possibly in Colima ( Fig. 4D ). The taxon´s host plant in the other states where it has also been recorded is unknown. Without doubt, males of A. jaliscana are among the fastest flying castniids in Mexico . Their flight is straight and not erratic, since they do not exhibit zigzagging behavior. Males begin to fly around 11:30, when the temperature fluctuates between 26–28°C; they rarely fly before 11:00. They usually fly at a height ranging from 5–8 m above the ground (average flight at 6 m above the ground). During the first 30–40 minutes of flight activity, they are slightly slower and calmer. Beginning at 12:00, when the day is warmer, they become more active and fly in a manner similar to or faster than the local skippers. Normally, they can be found on the highest leafless branches of trees of the species Guaiacum coulteri A. Gray, 1855 ( Zygophyllaceae ) measuring between 6–8 m in height, and rarely on trees shorter than 4–5 m ( Fig. 4F ). They perch in a stegopterous position, like the other Athis species known from Mexico ( Miller 1986 ; Ríos & González 2011 ; Vinciguerra et al. 2011 ; García-Díaz et al. 2020 ; García-Díaz 2022a , 2022b ); unlike them, however, they join their antennae together in front of their head, thus resembling the tip of a twig, while their wings look like dry leaves, resulting in perfect crypsis ( Figs. 4A, 4B, 4C ). No predators have been recorded attacking them yet. This is possibly due to their small size, fast flying speed, and their camouflage when perching. Males are highly territorial; they generally choose one or two different perching sites 4 to 6 m apart in their zone. They are usually found waiting for a female and frequently patrol their territory ( 400 m ²). On average, for every three minutes of flight they perch for one minute, so it is a very active species. When one or more males enter the territory of another, the resident male will pursue them for several minutes in a circular trajectory with the purpose of driving them away; generally, the intruders lose and withdraw from the disputed zone. These pursuing flights occur several times a day. Likewise, male A. jaliscana often chase and drive away skippers that fly close to them. However, when the intruder is a medium to large orange Nymphalidae butterfly species (such as Dione incarnata incarnata , Dryas alcionea moderata , Marpesia petreus , Euptoieta hegesia meridiania or Euptoieta claudia claudia , which often fly 5 m off the ground), the male A. jaliscana will fly close to it to investigate whether it is a female to be courted. But when the identification is evident, the male returns to its perch without bothering, chasing or fighting with it. When a male´s perching zone is situated near a wide dirt road, it will fly up to 50 m alongside it to investigate if a female is present and then return to its initial perching site. If the zone is in a dense and narrow trail, the males´ patrol flights are shorter (up to 20 m ). Males cease flying around 13:30, when temperature reaches 35–38°C. Females, on the other hand, begin to fly around 12:20, staying about 6 m off the ground or higher only, among branches of the tree species Guaiacum coulteri , searching for bromeliads on which to oviposit; consequently, they remain most of the time in the shade. Curiously, in Jalisco , it has been observed that they oviposit only on T. grossispicata that live on G. coulteri . They have never been seen perching on tree branches or leaves, suggesting that they are always active and flying. Their flight is slow, heavy and very calm, similar to that of E. hegesia meridiania and D. incarnata incarnata . When a female chooses a plant for oviposition, it will whirl around it to select the optimal site on which to leave an egg (in most cases they choose the base of the bromeliad). It then lands on the plant and slowly walks toward the center (the most protected part). Once on the middle of the leaf, it turns around, walks backwards and deposits an egg (only one per Tillandsia ). When it is done, it flies off in search of another plant for oviposition. No copulations were recorded; they are likely similar to those described by García-Díaz et al. (2020) and García-Díaz (2022b) , but at a minimum height of 6 m . The eggs, larvae and pupae of this species are unknown. Athis jaliscana adults have never been observed feeding on flowers, mud or decomposing fruit. Type material. Jalisco : 1♂ , Holotype , Tolimán , 20-VII-2011 , leg. B. López G. ( CNIN ) ; 1♂ , Paratype , Tolimán , 20-VII-2011 , leg. B. López G. ( BLG ) ; 1♂ , Paratype , Tolimán , 20-VII-2011 , leg. B. López G. ( RW ) . Additional material examined. Data from 35 specimens ( 30♂♂ , 5♀♀ ) were recorded: Colima : 1♂ , Aquiles Serdán ( Tamala ), VI-1973 , leg. A. Díaz F., Ex-Colección Saldaña ( CNIN ) ; 1♂ , Comala , 640m , 18-IV-1967 , leg. R. G. Wind , YPM ENT 901368 ( YPM ) ; Guerrero : 1♂ , Guerrero , mayo, Col. R. Müller, No. 3517 ( MHNCM ) [We think this specimen was possibly collected in Acahuizotla or in the vicinity of Chilpancingo de los Bravo]; Jalisco : 1♂ , La Huerta , Chamela , Estación de Biología Chamela , 2-V-1976 , leg. C. Beutelspacher B., ( CNIN ) ; 1♂ , Tolimán 2-VII-2015 , leg. B. López G. ( BLG ) ; 1♂ , Tolimán 28-VI-2016 , leg. B. López G. ( BLG ) ; 1♂ , Tolimán 8-VII-2017 , leg. B. López G. ( BLG ) ; 1♂ , Tuxcacuesco , Tuxcacuesco , 01-VII-2017 , leg. B. López G. ( CFT ) ; 1♂ , Tuxcacuesco , Tuxcacuesco , 18-VIII-2021 , leg. B. López G. ( CFT ) ; 1♂ , Tuxcacuesco , Tuxcacuesco , 16-VII-2018 , leg. B. López G., José de Jesús García-Díaz genitalia dissection No. JJGD0007 ( JJGD ) ; 1♀ , Tuxcacuesco , Tuxcacuesco , 20-VI-2015 , leg. B. López G. ( BLG ) ; 1♀ , Tuxcacuesco , Tuxcacuesco , 19-VII-2020 , leg. B. López G. ( JJGD ) ; 1♀ , Tuxcacuesco , Tuxcacuesco , 13-VII-2021 , leg. B. López G. ( JJGD ) ; 1♀ , Tuxcacuesco , Tuxcacuesco , 01-VIII-2021 , leg. B. López G. ( BLG ) ; 2♂♂ , Tuxcacuesco , Tuxcacuesco , 22-VI-2019 , leg. B. López G. ( JJGD ) ; 2♂♂ , Tuxcacuesco , Tuxcacuesco , 11-VII-2020 , leg. B. López G. ( JJGD ) ; 2♂♂ , Tuxcacuesco , Tuxcacuesco , 20- VII-2021 , leg. B. López G. ( JJGD ) ; 1♂ , Jalisco , Mexico , 700m , 5/Jul/2017 , leg. B. Lopez ( RW ) ; 2♂♂ , Jalisco , Mexico , 700m , 6/Jun/2018 , leg. B. Lopez ( RW ) ; 1♂ , Tuxcacuesco , Jalisco prov. Mexico , 29-VI-2018 , leg. B. López G. ( DC ) ; 1♂ , Tuxcacuesco , Jalisco , Mexico , 06-VII-2017 , leg. B. López G. ( DC ) ; 1♂ , Tuxcacuesco , Tuxcacuesco , 03-VII-2022 , leg. B. López G. ( BLG ) ; 1♂ , Tuxcacuesco , Tuxcacuesco , 03-VII-2022 , leg. B. López G. ( BLG ) , José de Jesús García-Díaz genitalia dissection No. JJGD0008 ( JJGD ) ; 1♂ , Tuxcacuesco , Tuxcacuesco , 03-VII-2022 , leg. B. López G. ( BLG ) , José de Jesús García-Díaz genitalia dissection No. JJGD0009 ( JJGD ) ; 1♂ , Tuxcacuesco , Tuxcacuesco , 11-VII-2022 , leg. B. López G. ( BLG ) ; 3♂♂ , Tuxcacuesco , Tuxcacuesco , 10-VII-2022 , leg. B. López G. ( BLG ) ; 1♂ , Tuxcacuesco , Tuxcacuesco , 07-VII-2023 , leg. B. López G. ( BLG ) ; 1♂ , Jalisco-Tuxcacuesco , 8 July 1972 [sic] ( MS ) ; Michoacán : 1♀ , Lázaro Cárdenas , Playa Azul , Arenal , V-1974 , leg. A. Díaz F., Ex-Colección Saldaña ( CNIN ) . 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. FIGURE 7. Overview of male wing morphology and interspecific wing shape comparison between the species. A: ♂ Athis jaliscana venation with vein names according to Miller (1986) ; B: ♂ Athis delecta venation with representation of morphological landmarks on fore- and hindwings; C: mean shape comparison between the species (♂ A. jaliscana shape with red lines and ♂ A. delecta shape with black lines). Male genitalia. Three specimens dissected ( Figs. 6B, 6C, 6D ). Tegumen subtriangular in lateral view and subtrapezoidal in dorsal view, slightly narrower in its posterior portion, with two small subtriangular lateral extensions in its distal 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 and sharper than the dorsal arms. Valva broad and almost rectangular; costal margin straight, ventral margin straight in the basal portion. Sacculus well-developed, arising from the inner surface of the valva. Saccus well-developed, narrow, and slightly 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, like A. delecta , shows low variability on both surfaces in both males and females. In the forewings, in some specimens, the discal spot that covers the accessory radial cell merges with the diagonal band. The size of the spots in the apical region is variable, and the spot within the diagonal band rarely disappears. The diagonal band, running from the apex to the anal margin, can be wider in some specimens. The size of the dark maculation between the outer margin and the diagonal band, between Cu 1b - 1A, is variable. In the hindwings, the spots forming the postdiscal band may be larger and more noticeable in some specimens.Additionally, the thickness of the margin and the length of its extensions toward the postdiscal band vary between specimens ( Fig. 3 ). Regarding the wing pattern, Athis jaliscana exhibits little sexual dimorphism. Like most Athis species , females are larger than males, with forewings that have a more rounded external margin and a greater curvature in the apical region. In terms of the wing pattern, the diagonal band is wider in females, and the discal spot is also larger and more elongated. Additionally, the basal and postbasal regions are darker in females. In the hindwings, the postdiscal band is wider in females and, unlike in males, is not separated by the orange coloration of the veins. The margin is darker and more blackish in females, and the margin extensions can merge with the postdiscal band, which rarely occurs in males. Most of the previously mentioned differences between sexes are also visible on the ventral side, with the postbasal band on the hindwings typically being wider and more prominent in females ( Fig. 3 ).