WARREN: New species of Atrytonopsis WARREN & DE LA MAZA: New species of Dalla TROP. LEPID. RES., 21(1): 7-11, 2011 7 A NEW SPECIES OF DALLA FROM CHIAPAS, MEXICO (LEPIDOPTERA, HESPERIIDAE, HETEROPTERINAE) Andrew D. Warren1 and Roberto G. De la Maza E.2 1McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural, History, University of Florida, SW 34th Street and Hull Road, P. O. Box 112710, Gainesville, Florida 32611-2710 U.S.A., e-mail: [email protected]; 2Sociedad Mexicana de Lepidopterología and Amigos de Kolijke A.C., Londres 71 Col. del Carmen, Coyoacán 04100, México, D. F., e-mail: [email protected] Abstract - A new species of Dalla is described from montane cloud forest habitats in southern Chiapas, Mexico. Based on male genitalia and characters of the wings and body, the new species appears to be closest to Dalla steinhauseri, a taxon known from montane cloud forest in eastern Oaxaca, Mexico, although it has been confused with Dalla freemani in the literature. Key words: Endemism, genitalia, Mesoamerica, morphology, skipper butterfly. Heteropterine skippers (Hesperiidae: Heteropterinae) quadrate; basal part of CuA1-CuA2, similarly sized, triangular; are the basal group of monocot-feeding hesperiids, being finally, roughly hourglass-shaped (roughly heart-shaped in distributed from the Himalayas to Japan and Europe, Africa paratype) spanning width of discal cell at proximal section and Madagascar, and North and South America (Warren et al. of distal fourth of cell, centered between the two macules in 2009). Seven genera of Heteropterinae fly in the New World, CuA1-CuA2, similar in size to them. Wing fringe brown. and by far the most diverse of these is Dalla Mabille, 1904, Dorsal hindwing same ground color as forewing; prominent with 96 currently recognized species (Mielke 2005, Viloria et overscaling of semi-irridescent olive setiform scales over basal al. 2008). 2/3 of wing, somewhat paler yellow-olive in discal cell and M3 Most Dalla species fly in the Andes Mountains of South to CuA2, most concentrated in basal half of discal cell and along America, although eighteen described species are known from inner margin to tornus. Pale ochreous, semi-opaque macules Central America and Mexico (Warren et al. 2011), eight of as follows: M1-M3, large, semi-rectangular, continuous across these being endemic to Mexico (Warren in prep.). Most species M2 with longest part in M1-M2, occupying distal end of discal of Dalla occur in highly limited geographic ranges, generally cell and basal half of cells M1 and M2; basal end M3-CuA1, at elevations above 900m, and often above 2500m (Viloria very small, roughly triangular, mostly overlapping macule in et al. 2008). Dalla males are most frequently encountered at M1-M3 though distal end produced somewhat distad; CuA1- damp ground, whereas females of Dalla are very rarely seen CuA2, centered just past mid-cell, larger, roughly half the (Steinhauser 2002, pers. obs.), and remain unknown for a large size of macule in M1-M3, roughly rectangular, overlapping percentage of species. Larval foodplants have been confirmed basal part of macules in M3-CuA1 and M1-M3; paratype with only for a few South American Dalla species, as noted by additional tiny macule at base of Rs-M1, over M1 and macule in Viloria et al. (2008), which are Chusquea Kunth (Poaceae) M1-M3; finally, small but prominent, rounded macule in discal bamboos. cell just proximad of midpoint. Wing fringe brown proximad, Below, we describe a distinctive new species of Dalla from pale ochreous distad, fringes somewhat darkened at vein ends. montane cloud forests of southern Chiapas, Mexico, based on Ventral forewing with macules repeated from the dorsal two male specimens collected by the junior author on two surface, each outlined with a narrow band of golden scales, different volcanoes. which also extend as a broad dash from M1, across Sc-R1 and R1-R2 to just before the costa, and, as somewhat paler golden Dalla austini A. Warren & R. G. De la Maza, sp. nov. scales, below the macules in CuA2-1A+2A, forming a smudge (Figs. 1a-d, 2a-e). spanning the central 2/3 of the cell, with pale golden scales densest (and richest in color) immediately below the macules; Description. Male (Fig. 1a-d)- forewing length = 14.6 mm ground color dark brown, with dense, cinnamon-brown (holotype), 16.4 mm (paratype); forewing apex fairly pointed, overscaling concentrated along the costa (caudad to M1) and termen evenly convex though nearly straight between CuA2 apex (to about M3), and sparsely distributed in adjacent cells; and 1A+2A, no stigma or brand; hindwing slightly produced at few pale creamy cells at ends of cells M1-M2 and M2-M3; base apex, termen convex to CuA1, then concave to tornus at 1A+2A, of discal cell and cells CuA2 and 1A+2A shiny grayish; wing inner margin convex. Dorsal forewing brown, scattered pale fringes dark at vein ends, areas between vein ends pale creamy golden setiform scales over most of wing, densest on basal proximad, especially from R5 to CuA1 (most pronounced on half of wing, especially along costa, inner margin and CuA2- paratype), darker distad, brown on holotype, cinnamon brown 1A+2A. Opaque pale ochreous macules as follows: subapical on paratype. in R3-R4 to R5-M1, quadrate, completely overlapping, forming Ventral hindwing with semi-opaque macules repeated from a nearly straight line perpendicular to the costa; posemedial in dorsal surface; additional pale ochreous (not semi-opaque) mid-M3-CuA1, larger, roughly quadrate with margin along macules in mid-Rs-M1, roughly triangular, and CuA2-1A+2A, CuA1 produced distad; mid-CuA1-CuA2, larger, roughly roughly circular, both arranged in a straight line with respect 8 TROP. LEPID. RES., 21(1): 7-11, 2011 WARREN & DE LA MAZA: New species of Dalla DallaChiapas1818.jpg 1818×2095 pixels 2/17/11 7:33 AM Fig. 1. Adults of Dalla austini, D. freemani and D. steinhauseri; a) dorsal and b) ventral views of holotype male of D. austini from Chiapas, Mexico, complete data in text; c) dorsal and d) ventral views of paratype male of D. austini from Chiapas, Mexico, complete data in text; e) dorsal and f) ventral views of holotype male of D. freemani from GUATEMALA: Dept. Sololá: Olas de Moka, 3000, September 1808 [probably September 1908], Geo. P. Englehardt, Genitalia Vial #96-2, Andrew D. Warren [AMNH]; g) dorsal and hht)t pv:/e/nwtwrawl. bvuitetewrfsli eosof fahmoelorictya.pcoem m/saelcere ot/fD Dall.a Cshteiaipnahsa18u1s8e.rjpig from MEXICO, OAXACA, Sierra Juárez, km. 95, La Esperanza-Cerro PPaeglóe n1, o f 1 7000’, 28 May 1990, John Kemner, Genitalia Vial SRS-3737 [MGCL]. to macules in M1-M3 and CuA1-CuA2; Sc+R1-Rs, roughly Dorsal head black with scattered dark brown, tan, and olive quadrate, borders somewhat indistinct; middle 1/5 of costal setiform scales, paler below eyes; dorsal labial palpus with cell, roughly quadrate (distal border ill-defined on paratype); mix of brown and pale golden setiform scales, paler laterally paratype with streak of pale scales basad of macule in M1-M3, and ventrally, inner surface dark brown; third segment dark along M1, overlapping pale macule in Sc+R1-Rs; scattered pale brown, porrect, barely extending beyond distal scales of second creamy scales at base of costal cell, Sc+R1-Rs, discal cell, and segment. Antennal shaft and club black on dorsum, mostly pale CuA2-1A+2A, as well as in margin, especially between M1 and golden on venter with some black between segments; nudum CuA1, and on paratype at tornus in CuA2 to 3A; wing fringes brown, 14 (holotype) or 13 (paratype) segments. Dorsal thorax dark at vein ends, much paler between vein ends, creamy black with semi-iridescent, olive, setiform scales, ventral proximad (paler on paratype), pale cinnamon distad. thorax with beige setiform scales, continuing onto ventral edge WARREN & DE LA MAZA: New species of Dalla WARREN & DE LA MAZA: New species of Dalla TROP. LEPID. RES., 21(1): 7-11, 2011 9 b a d c e Fig. 2. Male genitalia of Dalla austini holotype, genitalic vial GTA-14147, complete data in text; a) left lateral view of uncus, gnathos, tegumen, saccus, right valva, aedeagus and juxta; b) dorsal view of tegumen and uncus (hair tuft not shown); c) left lateral (exterior) view of left valva; d) left lateral view of juxta; e) ventral view of saccus. Scale = 1.0 mm. of femur; fore-tibia not spined, brownish epiphysis relatively about 3x width, costa shallowly concave on dorsal edge toward short, extending to distal end of tibia; mid-tibia with longitudinal caudal end, ampulla elongate, roughly length of costa, relatively row of 6 spines on inner surface, pair of spurs distad, outer spur broad (length about 2x length), angled slightly dorsad, caudal about 2/3 length of inner; hind-tibia with longitudinal row of end generally rounded but caudal end appearing somewhat about 6 spines, two pairs of spurs, in each pair, outer spurs squared off in lateral view, setose on both surfaces, especially about 4/5 length of inner; tarsus with three longitudinal rows caudad, harpe curved dorsad to blunt caudal end, exceeding and of short spines. Dorsal abdomen dark brown, with long, semi- not overlapping caudal extent of ampulla, dorsal edge weakly iridescent olive and pale golden setiform scales, somewhat grooved; juxta-transtilla (Fig. 2a, d) prominent with pointed paler caudad; ventral abdomen paler, with beige and yellowish dorso-caudal edge and spine pointed caudad in lateral view, setiform scales. subquadrate in ventral view; aedeagus (Fig. 2a) unadorned, Male genitalia (Fig. 2a-e). Uncus in lateral view (Fig. shallowly sinuate in lateral view, straight in dorsal view, caudal 2a) narrow, concave and slightly uplifted caudad of middle, end slightly expanded, vesica with single, sclerotized, roughly caudal end tapering to rounded point, slightly downturned; in rounded cornutus. dorsal view (Fig. 2b) caudal end narrow, rounded point, then broadening gradually cephalad towards junction with tegumen, Specimens examined. Holotype male with the following then narrowing slightly cephalad to overlap large portion labels: white, printed: / CHIAPAS / VN. HUITEPEC / NE. of tegumen as oval plate possessing dense pale brown hair B. MESOFILO / 2700M. 2 AGO 87 / R. DE LA MAZA E. /; tuft (removed in Fig. 2b); gnathos (Fig. 2a) slightly shorter white, printed and handprinted: / Genitalic Vial / GTA-14147 /; than uncus, narrow in lateral view, tapering slightly caudad red, printed: / HOLOTYPE / Dalla austini / A. Warren & R. G. to narrow somewhat rounded caudal end; in ventral view De la Maza /. One male paratype from MEXICO: CHIAPAS: relatively broad proximad, narrowing gradually to narrow Volcán Tacaná, 2800m, 18 April 2000, R. De la Maza E., rounded end; tegumen in dorsal view (Fig. 2b) broadly curved Genitalic Vial GTA-14148. The holotype is deposited at the cephalad, broadening slightly to junction with uncus; combined McGuire Center for Lepidoptera and Biodiversity, Florida ventral arm of tegumen and dorsal arm of saccus fairly evenly Museum of Natural History, University of Florida, Gainesville curved (Fig. 2a), somewhat swollen cephalad near midpoint; (MGCL); the paratype is deposited in the De la Maza Family cephalic arm of saccus (Fig. 2a, e) very long, nearly the length Collection, Mexico City, Mexico. of valva, thin, roughly cylindrical, angled about 20 degrees above horizontal in lateral view, cephalic end slightly swollen Type locality. MEXICO: CHIAPAS: Volcán Huitepec, NE and rounded in lateral and dorsal views; valvae (Figs. 2a, c) Bosque Mesófilo, 2700m. This site is comprised of dense symmetrical, long (1.4x length of tegumen and uncus), length montane cloud forest, dominated by Quercus acatenanguensis 10 TROP. LEPID. RES., 21(1): 7-11, 2011 WARREN & DE LA MAZA: New species of Dalla Trel., Bomarea acutifolia (Link & Otto) Herb., Crataegus The genitalia of the two specimens of D. austini, however, pubescens C. Presl., Salvia chiapensis Fernald, Ternstroemia are remarkably similar, differing slightly only in size, with tepezapote Schltdl. & Cham., Viburnum jucundum Morton, overall proportions of the genitalia of the paratype slightly larger Xylosoma flexciosum (Kunth) Hemsl., Litsea glaucescens than those of the holotype, apparently corresponding with the Kunth, Oreopanax xalapensis (Kunth) Decne. & Planch., slightly longer wing length of the paratype. Given the nearly Ostrya virginiana (P. Mill) K. Koch., Sambucus mexicana identical genitalia, similar habitats and altitudes of origin, and Presl., and some Alsophila R. Br. and Cyathea J. E. Smith considering the range of variation seen in many other species arborescent ferns. Along the rocky creeks in the area, scattered of Dalla (pers. obs.), these two specimens most likely represent colonies of the bamboo Chusquea foliosa L. G. Clark are found the same species. Hopefully, future field studies will be able to (Cortés 2005). This locality is strongly influenced by polar air clarify the range of phenotypic variation in D. austini. masses (nortes) during the winter, and is exceptional for its At first glance, wing markings on D. austini appear closest general scarcity of butterfly species throughout the year, even to those on D. freemani A. Warren, 1997, a species which to date during periods of favorable weather. Other butterfly taxa found is known only from the unique holotype male from Guatemala in the habitat include Colias philodice guatemalena Röber, (Figs. 1e-f). However, several differences between D. austini 1909, Iophanus phirrias (Godman & Salvin, 1887) and possibly and D. freemani exist. Dalla austini is larger (forewing length new subspecies of Oxeoschistus hilaria (H. Bates, 1865) and 14.6, 16.4mm) than D. freemani (forewing length 12.7mm), Pedaliodes napaea (H. Bates, 1865); remarkably, no species of with considerably paler dorsal hindwing spots. The small, Cyllopsis R. Felder, 1869 has yet been found here to date. rounded, deep ochreous dorsal forewing spot in CuA2-1A+2A Volcán Tacaná, where the paratype was collected, is on D. freemani (also shared with D. bubobon as shown by ecologically distinct from Volcán Huitepec in facing the Warren and González 1996 and Warren et al. 2011) is absent Pacific Ocean, and is therefore protected from cold polar air on D. austini, while the small, roughly triangular, semi-hyaline masses during the winter. While overall similar, the vegetation macule in M3-CuA1 on the hindwing of D. austini is absent on Volcán Tacaná includes additional tree species, such as on D. freemani. In addition, forewing macules, especially the Chiranthodendron pentadactylon Larreat., and a greater three apical spots and the macule in M3-CuA1, are considerably diversity of bamboos in the genera Chusquea and Rhipidocladum larger on D. austini than on D. freemani. McClure. A more impressive diversity of butterflies occur on The male genitalia of D. austini differ from those of D. Volcán Tacaná than on Volcán Huitepec, including Eretris freemani in several ways, as illustrated by Warren (1997). maria (Schaus, 1920), Drucina championi Godman & Salvin, The uncus tapers to a much finer point at its caudal end in 1881, three species of Pedaliodes Butler, 1867, four species of D. austini than in D. freemani, while the proximal end of the Cyllopsis, and many others, including Iophanus phirrias. tegumen (in dorsal view) in D. austini is noticeably broader than that of D. freemani. The valva of D. austini is longer and Etymology. Dalla austini is named in honor of the late George proportionally narrower than that of D. freemani; the harpe and T. Austin, Lepidoptera systematist, in recognition of his myriad ampulla narrowly overlap in lateral view in D. freemani, but contributions to our science. do not overlap, and are separated by a small gap in D. austini. The ventral arm of the saccus is proportionally longer in D. Distribution and phenology. To date, Dalla austini is known austini than in D. freemani. Finally (since the caudal end of the only from cloud forest habitats on Volcán Huitepec and Volcán aedeagus is missing from the holotype of D. freemani), while Tacaná, in the southern part of the Mexican state of Chiapas. overall rather similar, the shape of the juxta-transtilla is more It seems likely that D. austini occurs in similar habitats in irregular in D. austini than in D. freemani. Guatemala, and it is hoped that future fieldwork can confirm In essentially all genitalic characters, D. austini is most this hypothesis. Given that D. austini is apparently replaced similar to D. steinhauseri H. Freeman, 1991, a taxon known in similar habitats in Oaxaca by D. steinhauseri, the former is from cloud forest habitat in eastern Oaxaca, Mexico (Figs. not likely to range north of Chiapas. Confirmation of the larval 1g-h). As illustrated by Steinhauser in Freeman (1991), foodplant, likely a bamboo as noted above, will enable targeted the genitalia of the holotype male of D. steinhauseri are searches for D. austini in additional sites. remarkably similar to those of D. austini, including the shape of the long, relatively narrow valva, long saccus, similar Diagnosis and discussion. The two known specimens of Dalla tegumen and very similar juxta and transtilla; the aedeagus of austini show considerable individual variation, which might the two taxa is virtually identical, and no obvious differences in part be seasonal, with the holotype from August and the were noted in the cornutus. The upturned gnathos illustrated paratype from April. Forewing and hindwing macules on the for D. steinhauseri (Freeman 1991) represents an artifact of the holotype are slightly paler than those on the paratype, above and dissection technique- the gnathos on that taxon is actually very below, and the ventral forewing of the holotype has increased similar in shape, length, and orientation to that of D. austini cinnamon overscaling compared to the paratype. On the (Fig. 2a). There are, however, a couple subtle differences hindwing, the paratype has an extra, very small, semi-hyaline between the genitalia of the two taxa. The most prominent macule in Rs-M1, and in ventral view has an extra dash of pale difference is in the valva, where the harpe and ampulla overlap scales basad of macule in M1-M3, along M1, and overlapping slightly in lateral view in D. steinhauseri, whereas there is no pale macule in Sc+R1-Rs. The forewings of the paratype are overlap in D. austini, which shows a small gap between the about 1.8 mm longer than those of the holotype. harpe and ampulla in lateral view. In addition, in dorsal view, WARREN & DE LA MAZA: New species of Dalla WARREN & DE LA MAZA: New species of Dalla TROP. LEPID. RES., 21(1): 7-11, 2011 11 the caudal end of the tegumen is produced laterally to a greater REFERENCES CITED extent in D. austini than in D. steinhauseri. Despite the overall darker ventral ground color of D. steinhauseri, and its paler Cortés, R. G. 2005. El bambú en Chiapas. Bio Bambú, Revista Electrónica. www. forewing and hindwing macules, compared to D. austini, the Bambumex. org. p. 1. size and position of macules on the two taxa is very similar, Freeman, H. A. especially on the forewing, where the spot patterns are virtually 1991. A new species of Dalla from Mexico (Lepidoptera: Hesperiidae). identical. Hindwing spots on the two species differ mainly in Tropical Lepidoptera 2(1):65-67. Glassberg, J. the absence of the macules in the discal cell and in M3-CuA1 2007. A Swift Guide to the Butterflies of Mexico and Central America. on D. steinhauseri, above and below, and in minor details of Sunstreak Books, Inc., Morristown, New Jersey. [iii] + 266pp. ventral pattern and coloration. Thus considering all characters Glassberg, J. of the three taxa, including size (holotype of D. steinhauseri 2008. Skipperlings love geraniums. American Butterflies 16(3):30-38. Mielke, O. H. H. has a forewing length of 14.0 mm), D. austini appears perhaps 2005. Catalogue of the American Hesperioidea: Hesperiidae slightly closer to D. steinhauseri than to D. freemani, although (Lepidoptera). Sociedade Brasileira de Zoologia, Curitiba, Paraná, the three taxa are clearly closely related. Brazil, xiii + 1536 pp. While Dalla austini has remained undescribed until now, Steinhauser, S. R. 2002. Five new species of Dalla from Colombia and Ecuador it has been reported in the literature as D. freemani. Glassberg (Hesperiidae). Journal of the Lepidopterists’ Society 56(2):53-61. (2007, 2008) figured dorsal and ventral views of a live male of Viloria, Á. L., A. D. Warren & G. T. Austin D. austini from Chiapas (as D. freemani), but did not provide 2008. A spectacular new Dalla Mabille, 1904 from Venezuela-Colombia detailed locality information for the images. The live individual (Hesperiidae: Heteropterinae). Bulletin of the Allyn Museum 156:1- 12. is virtually identical to the holotype of D. austini in dorsal and Warren, A. D. ventral views. Thus, reports by Glassberg (2007, 2008) of D. 1997. A new species of Dalla from Guatemala (Lepidoptera: Hesperiidae). freemani from Mexico are in error, and represent D. austini. Tropical Lepidoptera 8(1): 35-37. Warren, A. D. The distribution of Hesperiidae in Mexico and adjacent regions. In ACKNOWLEDGMENTS preparation. Warren, A. D., K. J. Davis, N. V. Grishin, J. P. Pelham & E. M. Stangeland We sincerely thank Nick V. Grishin for discussions, 2011. Interactive Listing of American Butterflies. technical help and providing images of the holotype of Dalla [ 2 9 - I - 1 1 ] freemani, Andrei Sourakov for finalizing Figure 2, Isabel http://www.butterfliesofamerica.com/ Vargas-Fernández, Armando Luis-Martínez and Jorge Llorente- Warren, A. D. & L. C. González Bousquets for countless favors during post-doctoral studies in 1996. Rediscovery of Dalla bubobon in Michoacán, Mexico (Lepidoptera: Mexico in 2007-2009, when the specimens of Dalla austini Hesperiidae: Heteropterinae). Tropical Lepidoptera 7(1): 68-70. Warren, A. D., J. R. Ogawa & A. V. Z. Brower were first examined in the De la Maza Family Collection, Javier 2009. Revised classification of the family Hesperiidae (Lepidoptera: De la Maza for discussions and logistical help, and Thomas C. Hesperioidea) based on combined molecular and morphological Emmel and Jacqueline Y. Miller for providing an atmosphere data. Systematic Entomology 34:467-523. that promotes and facilitates research at the McGuire Center for Lepidoptera and Biodiversity. Two anonymous reviewers carefully reviewed this manuscript, and greatly improved it through their thoughtful suggestions. We also thank the late George T. Austin for help in initiating this project, discussions, genitalia dissections, and many years of friendship.