SYSTEMATICS AND BIOLOGY OF ASCIA (GANYRA) POPULATIONS IN THE SONORAN DESERT (JOSEPHINA, HOWARTH, ATAMISQUEA)
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Bailowitz, Richard Alan
SYSTEMATICS AND BIOLOGY OF ASCIA (GANYRA) POPULATIONS IN THE SONORAN DESERT
The University of Arizona M.S. 1985
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University Microfilms International
SYSTEMATICS AND BIOLOGY
OF ASCIA (GANYRA) POPULATIONS
IN THE SONORAN DESERT
by
Richard Alan Bailowitz
A Thesis Submitted to the Faculty of the
DEPARTMENT OF ENTOMOLOGY
In Partial Fulfillment of the Requirements For the Degree of
MASTER OF SCIENCE
In the Graduate College
THE UNIVERSITY OF ARIZONA
19 8 5 STATEMENT BY AUTHOR
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APPROVAL BY THESIS DIRECTOR
This thesis has been approved on the date shown below:
£ I wish to express my thanks to Steve Prchal, John Palting, and Elaine Greenapple for the photographs, Ken Kingsley for guidance and wisdom, Renee Vitali for the typing, and the members of my committee for general assistance. iii TABLE OF CONTENTS Page LIST OF ILLUSTRATIONS V LIST OF TABLES vii ABSTRACT viii INTRODUCTION 1 METHODS AND MATERIALS 3 RESULTS 5 Immature Stages 6 Egg . 7 Larva . 11 Pupa 16 Adult 17 Male . 17 Female 26 Host-Plant 26 Systematics 28 Josephina 28 Josepha 28 Howarthi 31 kuschei 31 Sevata, 31 DISCUSSION 33 Josepha and Howarthi Phenotypes. 35 Josepha and Howarthi Androconial Patterns 42 Josepha and Howarthi Genitalia 42 Josepha and Howarthi Distribution 43 Taxonomy: A Change in Status 45 APPENDIX A: COMPLETE DATA FOR MAINLAND ASCIA HOWARTHI POPULATIONS 47 APPENDIX B; COMPLETE DATA FOR WEST MEXICAN ASCIA JOSEPHINA JOSEPHA POPULATIONS 48 REFERENCES 49 iv LIST OF ILLUSTRATIONS Page Figure 1. Ascia howarthi egg on Atamisquea emarginata 9 2. Schematic drawing of A. howarthi 5th instar larva 10 3. Scanning electron micrograph of A. howarthi egg shell 12 4. First instar larva of A. howarthi on A. emarginata leaf 13 5. Second and fourth instar larvae of A. howarthi on A. emarginata 14 6. Fourth instar larva of A. howarthi on A. emarginata 15 7. Fifth instar larva of A. howarthi on A. emarginata branch 18 8. Pupa of Ascia howarthi 19 9. Venation patterns of Ascia howarthi and josepha.. 20 10. Ventral maculation of adult Ascia howarthi and josepha 21 11. Male genital capsule of A. howarthi 23 12. Scanning electron micrograph of the aedeagus of male A. howarthi 24 13. Androconial patterns of A. howarthi and josepha.. 25 14. Atamisquea emarginata at OPCNM... 29 15. Distribution of Ascia howarthi's larval host-plant 30 v vi Figure Page 16. Distribution of Ascia howarthi in the Sonoran Desert 36 17. Size and phenotypic comparison between A. howarthi and josepha 39 18. Phenotypic variation within A. howarthi and josepha 40 19. Distribution of A. howarthi and josepha 41 in Mexico and southern U.S. LIST OP TABLES Page Table 1. Time sequence for complete life cycle of Ascia howarthi 8 2. Size comparison of Ascia howarthi and Ascia josepha 37 3. Genitalic comparison between A. howarthi and A. josepha 44 vii ABSTRACT A breeding population of Ascia (Ganyra) (Lepidoptera: Pieridae), in the josephina complex, is confirmed for Arizona. The egg, various larval instars, and the pupa are discussed and figured for the first time. The larval host-plant is found to be Atamisquea emarginata Miers., a plant confined to the Sonoran Desert. This population, while closely related to Ascia josephina josepha, is distinct in maculation, androconial pattern, genitalia, and biology. Its taxonamic status is discussed. Full species status is given to the insect as Ascia howarthi (Dixey). viii INTRODUCTION During the course of a study for the National Park Service on arthropods of the Quitobaquito Management Area in Organ Pipe Cactus National Monument (OPCNM) Pima county, Arizona, Smith, Kingsley, & Bailowitz (in preparation) disclosed an apparent population of a pierid butterfly in the Ascia josephina (Godart) complex. The discovery is noteworthy in that records of this species complex are scarce in Arizona, in that the insect appeared to be breeding in the area, and in the odd phenotypes of the population. The genus Ascia is divided into two subgenera, each one represented by a single, Nearctic species or species complex (Howe, 1975). An additional taxon, A. sevata (c. & R. Felder), is recorded in Mexico. The subgenus Ascia includes the species monuste (Linnaeus) with its subspecies phileta (Fabricius), raza Klots, and cleones (Boisduval & Leconte). The subgenus Ganyra Billberg includes the species (or species complex) josephina (Godart) with its subspecies josepha (Salvin & Godman), howarthi (Dixey), and kuschei (Schaus). A. sevata is unplaced within the genus. 1 2 Most literature on the subgenus Ganyra in North American cites only josephina josepha as a member of this fauna. Howe (1975) stated that josepha is a resident along the Mexican border in southern Texas and that it strays occasionally into southern Arizona. Ehrlich & Ehrlich (1961) cited the species josephina from "southern Texas, straying northward to Kansas." Klots (1.951) considered it a breeding resident of the Rio Grande Valley, Texas. McGuire and Rickard (1974) cited it as migratory from Mexico in the lower Rio Grande valley, Texas. And Pyle (1981) termed it a "Resident in S. Texas, wandering into Arizona." It is not known from Florida (Kimball, 1965). The recent studies at Quitobaquito, OPCNM, confirmed the presence of an Arizona population of the A.josephina complex. Those studies also suggested the possibility that the population is resident and cast doubt on its previous taxonomic assignment. This thesis examines the dimensions of the Arizona Ganyra population and its relationships with other populations in the A. josephina complex. Also, this thesis describes the immature stages of the Sonoran Desert Ganyra population for the first time, links the population to a specific larval food-plant, and alters the taxonomy of the species complex. METHODS AND MATERIALS A series of both sexes of adults of Sonoran Desert Ganyra were collected from several wild populations. Those were augmented by specimens from the San Diego Natural History Museum (SDNHM), the California Insect Survey (CIS) collection, and the private collection of Kilian Roever. A total of 40 specimens was used. For comparative purposes a small series of A. josepha from southern and eastern Mexico as well as southern Texas was also obtained. Those 29 specimens were obtained from the SDNHM and CIS collections as well as from the Texas A & I collection and from the private collections of J. Brock, D. Mullins, and P. Hubbell, all of Tucson, Arizona. Field work included seven observation and collecting trips to OPCNM as well as two trips through Sonora, Mexico. Hastings, Turner, & Warren (1972) furnished plant distributions in Sonora and Baja California, Mexico. Collections were made throughout the year in order to circumvent seasonal dimorphism. Dates include October, 1983; January, February, April, September, November, and December, 1984; and January, March, and April, 1985. With additional specimens from the other sources, as mentioned above, records include all months of the year. 3 4 For the life history studies three females of the Sonora Ganyra were confined in a cylindrical container (diameter - 17 cm., height - 17 cm.) with a screened top. Sprigs of two Capparidaceous plants - Atamisquea emarginata Miers. and capparis spinosa L. - were placed in water containers within the cylindrical cage. Five eggs were obtained; one died, one was preserved, and three were raised to adulthood. Fresh sprigs of A. emarginata were added periodically. An attempt was made to raise A. emarginata plants from cuttings at several nurseries in Tucson without success. Attempts at hand pairing of the reared and wild-caught specimens to obtain additional larvae for chaeotaxy and crochet mapping were unsuccessful. No other wild-caught females could be induced to oviposit. Steve Prchal of the Arizona-Sonora Desert Museum (ASDM) photographed all stages of the life history. Measurements for consistency and variation of phenotypes and genitalia were done on a Lassico Ocular Filar. In addition, the author used the ISI-DS-130 scanning electron microscope at the University of Arizona campus for two micrographs. RESULTS Until 1983 investigators assumed that all Arizona Ascia records, specifically those in the subgenus Ganyra, represented seasonal influxes. However, regular collections of Ganyra in OPCNM beginning in 1983 suggested more than influx status. Freshness of many of the specimens, and a restricted center of abundance, even suggested breeding activity. Other species in the subgenus are known to utilize various members of the family Capparidaceae as larval food-plants (Jordan, 1981). Therefore, I consulted work of Bowers (1980) for members of that plant family with OPCNM, specifically in the vicinity of Quitobaquito. Two capers occur in OPCNM. One, Wislizenia refracta Engelm., occurs commonly and generally; the other, Atamisguea emarginata Miers., occurs very locally in the vicinity of Quitobaquito. For A. emarginata that represents its only known locale within the United States. Because W. refracta is widespread and only semi-perennial, and because adults of the Ascia were on the wing through the winter, this plant seemed the less likely of the two to be a host-plant. 5 6 On 18 November 1984 I collected three females of the Ascia at the OPCNM site. One died later that evening. I confined both remaining adults in a cage, along with fresh sprigs of A. emarginata and Capparis spinosa Linnaeus. (The latter plant was also chosen because it is a congener of a known food-plant for A. josepha josephina in Tamaulipas, Mexico) (Jordan, 1981). Female #2 died on 19 November 1984, with still no eggs evident. In an effort to duplicate the hot dry conditions of OPCNM at that time of year incandescent lights were used as a light and heat source. Activity of the remaining female seemed to increase with the temperature, so she was placed, in a cage, in the front seat of a Mazda truck 20 November 1984. This female laid 3 eggs on the gauze at the cage top just before noon. Two additional eggs were laid on 22 November 1984, one on the underside of an A. emarginata leaf. The female died 23 November 1984. Of the five eggs, one died, one was preserved in alcohol, and three hatched and were raised to adulthood (Table 1). Immature Stages A single egg was laid on the underside of the food-plant leaf (Fig. 1). The female laid three other eggs on the underside of the screen meshing at the cage top. At least three of the eggs were laid before noon, under very warm temperatures (inside my truck cab). Eggs hatched in approximately five days. 7 The larvae fed off and on throughout the day. They remained on the upper surface of the leaves, except when moving to a new feeding site, at which time they used the twigs. During feedings they positioned themselves with the head toward the tip of the leaf and the body along the mid-rib. The leaf was kept in an arrow shape by the feeding, with the mid-rib eaten last. Between feedings the larvae reversed their positions, lying along the leaf mid-rib with their heads just above the leaf petiole. This position was assumed without regard to the absolute positioning of the leaf. The total larval stage lasted approximately 36 days (Fig. 2). Pupation occurred on the branches with a silken girdle attaching the thorax to the twig,, The silk girdle failed on one of the pupae and a support was provided to simulate the original position. When disturbed, the abdomen of this specimen could be gyrated fairly vigorously in all directions. The pupal stage averaged 14 days. At eclosion, a large drop of brick-red fluid appeared beneath the pupal case on the floor of the cage. Egg The eggs are laid singly. They are nearly ovoid in shape with an everted, rimmed micropylar region. There are approximately 20 longitudinal ridges, some extending from the base to the micropyle, others ending some distance from 8 Table 1. Time sequence for complete life cycle of Ascia howarthi 1— B & c 20 Nov 20 Nov 22 Nov li.67 1st instar 25 Nov 25 Nov 26 Nov 6.67 2nd instar 1 Dec 1 Doc li Dec 7.33 3rd instar 6 Dec 7 Dec 15 Dec «.» lith instar 11 Dec 13 Dec 23 Dec 7.33 5th instar 19 Dec 20 Dec 30 Dec 8.67 Pupa 27 Dec 29 Dec 8 Jan lli.00 Adult 9 Jan 12 Jan 23 Jan — Ibtal 55.oo Fig. 1. Ascia howarthi egg on Atamisquea emarqinata 10 A6 SS Ail A3 J. Collar Fig. 2. Schematic drawing of A. howarthi 5th instar larva. Body segments and rudimentary pattern are shown. 11 the apex. The color is pale yellow. The height of the egg is 1.4 mm and diameter is 0.8 mm (Fig. 3). Larva The first instar has the head and body uniform green or yellowish green. There are several rows of setae evident through the entire length of the body. The length is 3-4 mm (Fig. 4). By the second instar, the head and body are still bright green, but T1 is slightly more yellow and somewhat prominent. The setae are larger and primary setae on the dorsal and lateral surfaces are tipped with fluid. Lengths average 5-6 mm (Fig. 5). The third instar is very similar to the second. The setae are still tipped with fluid and the body length has increased to 7-9 mm. In the fourth instar, the dorsum is still shiny green, but a whitish lateral stripe has formed on Tl, T2, and T3, tapering out in Al. Under magnification, this stripe is turquoise, white, and yellow-orange. The thoracic spiracle is very evident. The setae are still tipped with fluid. An additional fringe of small setae, low laterally on the body and nearly touching the substrate, is also apparent. The length is now 10-15 mm (Fig. 6). During the fifth instar, the head is heavily setaceous and the collar is dark with turquoise spots laterally and dorsally. The body is green and the Fig. 3. Scanning electron micrograph of A. howarthi egg shell First instar larva of A. howarthi 14 Fig. 5. Second and fourth instar larva of A. howarthi on A. emarginata The 4th instar larva is showing the typical resting non-feeding posture, with head at the leaf base and body along the leaf mid-rib. Fig. 6. Fourth instar larva of A. howarthi on A. emarginata The light thoracic lateral stripe is visible. 16 longitudinal stripe is evident, being pale turquoise over pale yellow, and extending from T1 through the length of the body. Also, several small yellow spots extend from A1 to the posterior end, on a line just dorsad to the base of the prolegs. Each body segment is divided into six subsegments. The ventral surface is rough, greenish white, closely matching the leaf underside. No fluid is evident at the setal tips. The length is 17-26 mm (Fig. 7). During the pre-pupal stage, the body is condensed. The anal prolegs are not attached to the substrate. The turquoise, white and yellow stripe is lost from segments T1, T2, and T3. The entire length is now approximately 21 mm. Pupa In color, the pupa is pale brown with minute black dots over the entire surface. There is a white projection, approximately 2 mm long, at the anterior end of the pupa. At mid-thorax there is a noticeable hump, behind which a line of several small spines runs down the mid-line to near the end of the abdomen. Short orange dashes mark the spaces between those spines. There are also two striking, large black metathoracic spines, projecting at an angle laterally, approximately 20° above the horizontal. Each extends two mm in length. The wing casings become transparent approximately two days prior to eplosion while the remainder of the pupa darkens. The pupal length is 19-22 mm (Fig. 8). 17 Adult Dixey (1915) described the adult phenotypes of both sexes from Baja California. Parameters of the phenotype are expanded somewhat here to include mainland specimens as well. Male Forewing length 28.14 mm, range 23-33 mm, S=2.82, n=27. The general ground color is white to greenish white. The dorsal forewing has a large black spot at the cell end, averaging slightly smaller on Baja California specimens. While occasionally immaculate, there is usually some dark brown to black marginal scaling at the apex and along veins Ml, M2, M3, Cul, and Cu2 for 1-5 mm. Also, there is often a suffused dark submarginal patch straddling vein M3, especially on Sonora specimens. The dorsal hindwing is usually immaculate white, but occasionally possesses a narrow black dash on the cross-vein at the base of cell M2 (Figs. 9, 10). The ventral forewing is similar to the dorsal surface, but with the dark markings slightly more prominent and with an additional dark patch in cell Cu2, halfway between its base and the wing margin. The ventral hindwing has a prominent small dark spot at the anterior end of the cross-vein at the base of cell M2 as well as two suffused spots distad of the cross-vein near the base of cells M2 and M3. On Baja California specimens, much of the ventral Fig. 7. Fifth instar larva of A. howarthi on A. emarqinata branch f, . 19 A B Fig. 8. pupa of Ascia howarthi a) lateral view b) dorsal view i u? Fig. 9 Venation patterns of Ascia howarthi and josepha a) howarthi b) josepha KS 21 Fig. 10. Ventral maculation of adult Ascia howarthi and josepha a) howarthi b) josepha .j 22 hindwing is overlaid with a fine network of pale, gray-brown reticulations. This phenotype is figured by Dixey (1915). The male genitalia are symmetrical (Fig. 11). The entire genital capsule averages 4.02 mm in length with the saccus occupying approximately l/7th of that length. The valvae are broad, with the posterior projection curving slightly inward. The uncus is wide at its base and has noticeable shoulders before tapering to a point. The aedeagus averages 2.26 mm in length. At approximately l/5th the length from the base, there arises a projection, referred to here as the "elbow" (Fig. 12). Dixey (1915) figured androconial scent scales of the male. However, the scent scales actually occur in two distinct sizes - a major and a minor. The major scales agree closely with Dixey's representation, averaging 0.04 mm in width by 0.47 mm in length. The minor scales are noticeably smaller, averaging 0.03 mm by 0.38 mm. They are placed in thick mats over confined areas of the forewings (Fig. 13). In reflected light, those patches appear exceedingly white, but in transmitted light they appear as dark blotches. The androconia are confined to that portion of the wing outside the discal cell in cells M3, Cul, Cu2, and 2A. The actual pattern within those cells is quite distinct, appearing as elongated C's in M3 and Cul, as chevrons in Cu2, and as a parallelogram in 2A. Male genital capsule of A. howarthi a) lateral view b) dorsal view Fig. 12. Scanning electron micrograph of the aedeagus of male A. howarthi To the left of center is "elbow." Fig. 13. Androconial patterns of A. howarthi and josepha a) howarthi b) josepha 26 Female Forewing. length 27.42 mm, range 24-32 mm, S-2.68, n-12. The ground color is similar to that of the male. On the dorsal forewing there is a large black spot at the cell end. Otherwise, it is rarely immaculate, usually possessing some marginal and apical dark scaling. The spotting in cells M2 and M3 is more extensive than in the male. In addition, there is a suffused spot in cell Cu2, midway to the margin. The dorsal hindwing is similar to that of the male, but occasionally possesses dark scaling at the vein ends. The ventral surfaces are similar to those of the male. Host-Plant The host-plant, Atamisquea emarginata Miers., is a woody, perennial shrub as tall as 3 m (Bowers, 1980). It grows on calcareous, usually littoral soils and saline flats, occasionally on inland sandy washes. The population at OPCNM is the most northerly known, representing the only known stand of the plant within Arizona and, in fact, within the U.S. All known plants at OPCNM are located at three sites: at Quitobaquito pond, along Aguajita wash within 1 km of the Arizona/Mexico border, and along the Puerto Blanco Drive, approximately 24 km. west of Highway 85 (Bowers, 1980). In personal communications, Bowers stated that the last site is represented by a lone plant. A total of fewer than two dozen plants is known from those areas. 27 The plants grow both in the open on sandy benches along the washes and beneath and among the relatively lush growth of these somewhat riparian areas„ The plant association is mainly mesquite woodland, consisting of Prosopis velutina Woot., Olneya tesota Gray, Condalia warnockii M.C. Johnst., Zizyphus obtusifolia (Hook, ex T. & G.), Dalea spinosa Gray, and Lycium parishii Gray (Peter Warren, personal communication). In general aspect, the dark green coloration of the A. emarginata is deeper than that of the various Lyciums, which it otherwise resembles (Fig. 14). The leaves are dark gray-green on the upper surface and pale silvery-green on the underside. The stems and undersurfaces of the leaves are also lepidote and to some degree glandular (Kearney and Peebles, 1960). They appear tough, almost succulent, in accordance with Type IV leaves, as mentioned by Jordan (1981). This toughness probably aids in water retention, an idea supported by the viable condition of cuttings after long periods of time. Fresh sprigs cut and refrigerated kept their leaves intact at least 72 days. A. emarginata is confined to the Sonoran Desert and to an additional population in Argentina (Hastings, Turner, and Warren; 1972). In Baja California, it is widely distributed northward to approximately 29° north latitude (on lsla Angel de la Guarda). On the mainland it occurs from extreme northern Sinaloa (Fig. 15), northward to the 28 OPCNM site, and never more than approximately 130 km from the coast. Systematics Most of the taxa in the A. josephina complex, i.e., A. josephina, A. josepha, A. howarthi, A. kuschei, and A. sevata, were originally described in the genus Pier is Schrank. They were later placed in the genus Ascia Scopoli and still later in either the genus or subgenus Ganyra Billberg, depending on the author. That was done, in part, to separate the josephina complex from the rest of the genus, e.g., A. monuste. josephina Godard described nominate josephina in 1819 from the Antilles, probably Haiti and Cuba (Salvin & Godman, 1868). It is characterized by the large size, a falcated forewing, a produced hindwing, and a large black forewing cell-spot. Two other closely related Antillean subspecies have been described: paramaryllis Comstock from Jamaica, and krugii (Dewitz) from Puerto Rico. Both of these are somewhat smaller, with the characteristic black forewing cell-spot narrow (Comstock, 1943) Josepha This subspecies is the widespread Mexican and Central American form, described by Salvin & Godman (1868). It differs from nominate josephina primarily in the shape of Fig. 14. Atamisquea emarqinata at OPCNM The plant is more or less centered in the photograph, being behind and to the author1 right. 30 Distribution in the Sonoran Desert of Atamisauea emarclnata Miers Fig. 15. Distribution of Ascia howarthi's larval host plant Amended from Hastings, Turner, & Warren; 1972. I 31 the wings. The forewings are not as falcate as those of josephina and the hindwings are more squared and less produced at the anal angle, it occurs from southern Texas (rarely northward) through eastern Mexico into Central America and some distance north along the west coast of Mexico. Howar thi Dixey (1915 described subspecies howarthi from Tembabichi Bay, Baja California Sur, Mexico. He described it as having more noticeable marginal spotting on the forewing and more pronounced reticulations on the ventral hindwing than typical josephina, as well as being comparatively small (males 58 mm wingspread, females 52-54 mm). Kuschei Schaus (1920) described this subspecies from Mazatlan, Sinaloa, Mexico. It is characterized by a greater extent of the submarginal maculation of the forewings in both males and females. He cited both sexes as having wingspreads of 56 mm, also comparatively small. Sevata C. & R. Felder (1861) described the taxon sevata from Venezuela. Rindge (1948) cited the Baja California population of Ganyra as A. sevata kuschei, a new combination 32 at that time. The original description of sevata mentioned a solid white dorsal surface except for the apical marginal border, wider in females than in males. Fruhstorfer (1908) subsequently named a more northerly subspecies, A. sevata tiburtia, from Guatemala. It differs from true A. sevata in the narrowing or absence of A. sevata's 5 mm forewing border (Fruhstorfer, 1908). The ventral hindwings of A. sevata and A. tiburtia are suffused with shades of violet and pink respectively. No mention of forewing cell-spots is made for either taxon. DISCUSSION There are obvious close superficial relationships among the various members included in the subgenus Ganyra. A closer look will now be given to the exact relationships between those taxa (north of Guatemala) and where the OPCNM population lies in reference to them. Several key points suggest that A. sevata sevata and A. sevata tiburtia are more distantly related to the immediate complex. The lack of the black forewing cell-spot does not in itself suggest removal from the group since Cuban, Jamaican, and Puerto Rican populations of josephina also have this spotting weak to non-existent (Comstock, 1943). However, the presence of a narrow to wide continuous forewing border and pink to purple ventral hindwing coloration do set these taxa apart. More importantly, there are significant structural differences in the form of the male androconia. These average only 0.24 mm in length in sevata, approximately half that of the josephina complex members (Dixey, 1915). Hoffman (1976) sited tiburtia from southern Mexico, a range overlapping that of josepha. This sympatry suggests distinction at the specific level. 33 34 Sevata is here considered a member of the subgenus Ganyra, but outside the josephina complex. A second relationship is that between A. howarthi and A. kuschei. Although kuschei was described from Mazatlan, it was undoubtedly collected far north of there. There are many recent records of a Ganyra from the Mazatlan area, all of which represent josepha not kuschei. The maculation, androconia, and size all suggest this. The location "Mazatla'n" was probably used in a broad sense by Kusche. Phenotypes matching the kuschei description occur from extreme northern Sinaloa northward, where he easily may have collected. Many specimens from Sonora and Arizona, and a few from Baja California closely match the kuschei description, especially for broods during the summer rains. The type locality for kuschei should be amended to San Miguel, near Los Mochis, Sinaloa, Mexico, the southern-most locality for which that taxon is know. The howarthi phenotype, at least ventrally, is restricted to Baja California. But other phenotypes with unmarked ventral hindwings also occur there. These unmarked phenotypes are also widespread in central and northern Sonora and represent the majority of specimens taken at OPCNM. The three phenotypes - heavily marked kuschei, ventrally marked howarthi, and the unmarked population - all blend clinically with each other. When mapped together, 35 their composite distribution duplicates that of the food-plant, A. emarginata (Fig. 16). Kuschei and howarthi, plus the unmarked phenotype are therefore considered synonymous. Similarities in size, androconial pattern, and the distribution of the larval food-plant, all suggest this. Since howarthi has priority, the name kuschei is suppressed as a junior synonym, perhaps to be used as a name for the wet season form. The third most important clarification necessary then is the relationship between josepha and howarthi. This relationship will be examined with regard to phenotype, androconial pattern, genitalia, and food-plant distribution. Josepha and Howarthi Phenotypes On the basis of size, howarthi and josepha represent two very different populations (Table 2). Both sexes of howarthi average in excess of 10 mm smaller in forewing length than those of josepha, There is no size overlap in the males, and only a single, especially small female josepha overlaps the size range of female howarthi. The size mentioned for howarthi by Dixey (1915) seems overinflated. He cited the male wingspread as 58 mm, but the male figured is only partially spread. His figure has a 55 mm wingspread with a 26 mm forewing. In contrast, a well-spread specimen with a 26 mm forewing length has only a 45 mm spread. In the literature, josepha is given the common name of Giant White (Klois, 1951; Pyle, 1981; etc.), 36 ® @© Fig. 16. Distribution of Ascia howarthi in the Sonoran Desert • breeding colonies © influx Table 2. size comparison of Ascia howarthi and Ascia josepha KoT.rarthi Josephina male femal9 male female Forevang length 28.Ul mm 27.U2 ram 38.73 ram 33.82 ram 1J-27 N-12 N.18 Moll Hani^e 23-33 mm 2ij-32 mm 35>-U3 mm 31-Uii mm Total 1 28.10 ram 38.79 mm 11=39 K=29 38 a name hardly fitting the much smaller Sonoran Desert populations. The wing shape is also moderately different in the two taxa. In howarthi the forewings are somewhat shortened and the hindwings are very rounded, in josepha, and apparently even more so in nominate josephina, the forewings are produced or even falcated while the hindwings have the anal angle likewise extended (Fig. 17). In their maculation, the males of howarthi are generally more heavily patterned than those of josepha, especially with forewing marginal and submarginal spotting. Females are very similar in the two taxa, but in- josepha dark morphs are often produced where the ground color is heavily overlain with cinnamon brown. All specimens of this morph seen (Fig. 18) were mid-summer captures and probably parallel the long-day form of Ascia monuste (Pense, 1962). The absence of dark morphs in howarthi is interesting in light of the paucity of records away from breeding colonies. (Appendix A and Fig. 19). Both sexes of howarthi have, on the ventral hindwing, an additional diagnostic mark. Aside from the dark scaling distal to the cross-veins at the base of cells M3 and Cul, there is a darker spot on and around the cross-vein at the base of cell M2. While rarely males of josepha will have that cross-vein darkened, even dorsally, it is not scaled away from the vein as in howarthi. Fig. 17. Size and phenotypic comparison between A. howarthi and josepha The top figures are howarthi, male on the left, female on the right. The bottom figures are josepha, male on the left, female on the right. X r-\ £ " \ t* * i- *• <• -w • fl ( ' Jr _ f t ./i'w/' <*>' : Fig. 18. Phenotypic variation within A. howarthi and josepha Distribution of ,Iosepha - • howarthi - O Fig. 19. Distribution of A. howarthi and josepha in Mexico and southern U.S. 42 Josepha and Howarthi Androconial Patterns While the size and configuration of howarthi and josepha androconia scales are similar (Dixey, 1915), their placement on the wings is vastly different. All androcania of howarthi are confined to forewing cells M3, Cul, Cu2, and 2A (see earlier section and Fig. 13). On josepha, the androconia in these cells are far more extensive. In addition, scent scales are present in cells M2, Ml, and the discal cell. Furthermore, josepha invariably has androconia on the hindwing in cells RS, M2, M3, the discal cell, and occasionally in Cul. Josepha and Howarthi Genitalia The general configurations of the male genitalia of both taxa are similar. Due to the larger size of josepha, the entire genital capsule is proportionately larger in that taxon than in howarthi. However, the length of the saccus is proportionately greater in howarthi after correction for size difference (Table 3). Likewise, the aedeagus is proportional in the two taxa, but the aedeagal elbow length of josepha is significantly greater after correction for size differences. While these slight genitalia differences are not spectacular, they are present and, at least in the case of the saccus length, usable diagnostically in certain circumstances. 43 Josepha and Howarthi Distributipn Because phytophagous insects are closely tied to their larval food-plants, discussion of host-plant and insect distributions will be somewhat intertwined. For the genus Ascia, members of the families Cruciferae, Capparidaceae, and Batidaceae have been reported as host-plants (Howe, 1975). More specifically, Capparis frondosa Jacq. was cited as a host-plant for Ascia josephina josepha (Jordan, 1981). Although this caper is confined to the eastern slope of Mexico, mostly Tamaulipas and Veracruz (Standley, 1961), other members of the genus have wider distributions in Mexico and probably serve as food-plants in other parts of the range of josepha. Members of the genus Capparis, whose distributions include Sinaloa, are C. flexuosa L., C. verrucosa Jacq., and C. indica (L.) (Stanley, 1961). Any of these might serve as larval hosts for josepha since the plant and insect northern distributional limits appear to match one another. None of the members of the genus Capparis has been reported as far north as Sonora. And, records for josepha extend north to Mazatlan, Elota, and Guamuchil, only approximately 100 km southeast of Los Mochis (Appendix B). For howarthi, records for Atamisguea emarginata extend south into northern Sinaloa, near Los Mochis (Fig. 19). This brings the two taxa within 100 km of each other. In fact, a somewhat dubious record of a dark morph female I 44 i'l SI I't'l Table 3. Genitalic comparison between A. howarthi ; ' and A. josepha X Jos "X HOT; Ran-e Jos Han?e How ? value n-10 n-19 Ae.L (a) 3.009 2.258 2.73-3.28 1.98-2.53 .0000 El.I (b) .1390 .1153 .111-.2k .06-.17 ,0000 1C.L (c) 5.k35 L..023 5.06-5.73 3.68-L.50 •0000 3a.L (d) .6710 .5779 .55-.71 •U7-.77 J .0022 sac (s) 12.22 lk. 37 10.S7-13.0k 12.2k-l3.2n j .""00 nr. 1 T-.l (f) 38.10 28.63 } 35—ill 23-33 J .0000 100Ae (?) 7.826 7.932 7.15-8.63 6.39-9.65 .6552 F'.L 100E1 (h) • li930 • ii037 •37-.61 .21-.63 .0279 rr.i 10C-C (i) Ik.29 lk.12 13.33-15.61 12.59-16.61 .6b61 rv.L lOOS a (.1) 1.763 2.03k l.k9-2.00 1.56-2.70 .0167 ?7;.L :i li Table 3. Genitalic comparison between A. howarthi and A. josepha (a) Length of the aedeagus (b) Length of the elbow (c) Length of the genital capsule (d) Length of the saccus (e) Ratio of saccus length to genital capsule length (f) Forewing length (g) Ratio of the aedeagal length to the forewing length (h) Ratio of the elbow length to the forewing length, times a constant (i) Ratio of the genital capsule to the forewing length, times a constant (j) Ratio of the saccus length to the forewing length, times a constant 45 josepha taken near Alamos, Sonora would bring the two populations into overlap. The lack of an apparent cline or of anything resembling intergrades suggests a high integrity to the two populations. While specific records of howarthi from Baja California are too numerous to mention, there being 150 specimens in the SDNHN alone, specific records from the mainland population (which interfaces with A. josepha are given in Appendix A. Taxonomy: A Change in Status There are significant differences between the Sonoran Desert's howarthi and josepha of the remainder of Mexico. These same differences persist when howarthi is compared to josephina as a whole. It therefore appears that the differences are at the species level, not at the subspecies level. There are phenotypic differences including wingshape, maculation, and androconial pattern. There are some slight to moderate genitalia differences. There are significant habitat differences. And there is a difference in host-plant preference. Therefore, I propose to elevate the taxon howarthi to species status. It is most closely related to A. josephina, less so to A. sevata. The genus north of Guatemala and exclusive of the Antilles would be composed of two subgenera and four species as follows: 46 Genus: Ascia Scopoli Subgenus Ascia Scopoli 1. monuste (Linnaeus) a) monuste (Linnaeus) b) phileta (Fabricius) c) cleomes (Boisduval & Leconte) Subgenus Ganyra Billberg 2. sevata (C. & A. Felder) a) tiburtia (Fruhstorfer) 3. josephina (Godart) a) josepha (Salvin & Godman) 4. howarthi (Dixey APPENDIX A COMPLETE DATA FOR MAINLAND ASCIA HOWARTHI POPULATIONS MEXICO Sinaloa - San Miguel (A. Allen) 17 June 1956, 1 9- Sonora - 54 mi. S. Hermosillo (G. Forbes) 12 Mar. 1974 1 cf Peon (R. Bailowitz) 26 Dec. 1984, 1 9 San Jose de Guaymas (R. Bailowitz) 2 Jan. 1985, 1 o* 15 mi. S. Navajoa (D. Mullins) 6 Oct. 1975, 1 o" 2 mi. NE, Atil (R. Bailowitz) 14 Apr. 1985, 1 o» UNITED STATES Arizona - OPCNM, Pima County (R. Bailowitz & K. Kingsley) 22 Oct. 1983, 4 cf»s, 1 2 OPCNM, Pima County (R. Bailowitz) 23 Oct. 1983, 2 o*s Same data, 7 Jan. 1984, 1 cP, 1 9 Same data, 5 Feb. 1984, 1 o" Same data, 22 Apr. 1984, 1 o* Same data, 22 Sept. 1984, 1 o" Same data, 18 Nov. 1984, 1 $ Same data, 2 Mar. 1984, 2 o*s, 1 9 OPCNM, Pima County (K. Roever) 11 Oct. 1970, 3 cfi s * Tucson, Pima County (Fenn) 23 Apr. 1962, 3 cf» s * San Pedro River nr. Benson, Cochise County (K. Roever) 30 Apr. 1965, 1 * Kansas Settlement, Cochise County (K. Roever) 12 Aug. 1963, 1 ? * 18 mi. E. Douglas, Cochise County (K. Roever) 5 May 1962, 1 o", 1 9 * 2 mi. SW Parker, La Paz County (K. Roever) 6 Sept. 1973 1 9 •Probably influx, non-breeding specimens 47 APPENDIX B COMPLETE DATA FOR WEST MEXICAN ASCIA JOSEPHINA JOSEPHA POPULATIONS Colima - Colima (P. Spade) 24 Sept. 1980, 1 d* same data, 23 May 1980, 1 9 same data, 1 Oct. 1979, 1 9 same data, 8 Sept. 1980, 1 o* Madrid (P. Spade) 25 Apr. 1981, 2 2 9 same data, 24 Sept. 1980, 1 Jalisco - Est. Biol., Chamela (Doyen & Chemsak) 20-27 July, 1 9 Sinaloa - Hwy. 40 nr. Concordia (Breedlove) 14 Sept. 1977, 3 d* btw. Culiacan & Mazatla'n (B. & S. Shor) 23 Aug. 1963, 1 9 25 mi. S. Elota (J. Powell) 7 Aug. 1964, 1 o» Mazatl^n (J. Brack) 10 Jan. 1979, 1 cf same data, 1 Jan. 1975, 1 cJ* Hwy 40 nr. Concordia (D. Mullins) 14 Sept. 1977, 1 cf* Hwy. 40, 20 mi. E. Hwy. 15 (D. Mullins) 10 Oct. 1975, 1 cf Sonora - *Rio Cuchajaqui nr. Alamos (S. Prchal) 13 Sept. 1979, 1 9 •locality label somewhat suspect. 48 REFERENCES Bowers, Janice E. 1980. Flora of Organ Pipe Cactus National Monument. J. Ariz.-Nev. Acad. Sci. 15: Issue 1, 11. p. 1-47. Comstock, William P. 1943. The genus Ascia in the Antilles. Amer. Nus. Novit. #1229: 1-7. Dixey, F. A. 1915. New Species and Subspecies of Pierinae. Trans. Entomol. Soc. Land. Part 1: 9-17. Ehrlich, Paul A., & Anne H. Ehrlich. 1961. How to know the butterflies. Dubuque, la, N. C. Brown Co. 162 pp. Felder, C, & R. 1861. Lepidoptera nova Columbiae. Wien. 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