PROCEEDINGS OF THE Denver Museum of Natural History

SERIES 3, NUMBER 3, OCTOBER 15, 1993

A REVIEW OF THE AGAPEMA (: )

RICHARD S. PEIGLER Department of Zoology, Denver Museum of Natural History 2001 Colorado Boulevard, Denver, Colorado 80205-5798

and

ROY O. KENDALL 5598 Mt. McKinley Drive N.E. San Antonio, Texas 78251-3626

ABSTRACT — Agapema is a genus of saturniid ranging in Mexico and the southwestern United States. Agapema galbina, the type-species, was found to be misidenti- fied by authors during the last 21 years. John Pope collected the original type specimens in the lower Rio Grande Valley of Texas, not in western Texas. Agapema platensis, spec. nov., is described from the Edwards Plateau of Texas. Four taxa in far western Texas (dyari, nom. rev.), Arizona and western Mexico (anona, stat. rev.), Baja California (pelora, stat. nov.) and northeastern Mexico (dentifasciata, stat. nov.), previously considered to he subspecies of galbina, are elevated to full species rank based on larval and genitalic dif- ferences. The female and mature larva of dentifasciata and mature larvae of dyari and platensis are described for the first time. Figures and a key to the adult moths are pro- vided. The distribution of each species is plotted on a map. All known records for host- plants and parasitoids are tabulated, and phylogeny, habitats, and other field observa- tions are discussed.

KEYWORDS: Agapema, Arizona, Colorado, Condalia, galbina, Mexico, moths, par- asitoids, Rhamnaceae, , Saturniidae, , Texas

PEIGLER AND KENDALL

The genus Agapema is a holophyletic group of TAMU— Texas A&M University, College small to medium-sized nocturnal saturniid moths that Station range in southwestern North America south to the region SDNHM— San Diego Natural History Museum, of Mexico City. The genus was established by Neumoegen San Diego and Dyar (1894); the type-species is Saturnia galbina by KU — Snow Entomological Museum, Uni- monotypy. The most recent revisionary treatments of the versity of Kansas, Lawrence genus were by Ferguson (1972) and Lemaire (1978). The USNM— United States National Museum of currently accepted classification is: A. galbina galbina Natural History, Smithsonian Insti- (Clemens) from western Texas and southern New tution, Washington, D.C. Mexico, galbina anona (Ottolengui) from southern Arizona and northwestern Mexico, galbina dentifasciata Lemaire from eastern Mexico, galbina pelora Rindge from TAXONOMY Baja California, solita Ferguson from the lower Rio Earlier authors, up to and including Collins and Grande Valley of Texas, and homogena Dyar from north- Weast (1961), were correct in considering that the type- ern Colorado to Mexico City. species of Agapema was the one that formerly occurred The taxonomy has been confused, and the imma- commonly in the lower Rio Grande Valley of Texas ture stages of several of the taxa have not been reported. (Cameron and Hidalgo Counties). Ferguson (1972: 191) This paper provides new information on hostplants, local- wrote: "It has long been accepted that the name galbina ity records, habitats, parasitoids, immature stages, phy- applied to the form found in the lower Rio Grande Valley, logeny, and taxonomy. In contrast to a generic revision, but when I searched for the evidence upon which this which should provide complete synonymies, redescrip- conclusion was based I could find none." He continued tions of all taxa, and a comprehensive bibliography, this with a discussion of an 1854 survey expedition in western paper is a review mainly intended to fill in some gaps Texas by John Pope, who collected the type material in where information has been lacking in earlier publica- Texas, concluding that Pope was never in the lower Rio tions, and to correct errors of earlier authors. Recently, Grande Valley (see Goetzmann 1959). Ferguson then much new material has become available to us on the considered the type-species to be in western Texas, where taxon that inhabits a region of southwestern Texas, partic- he knew Pope had travelled, and redescribed the lower ularly the western portion of the Edwards Plateau. This Rio Grande species as new, i.e..solita. In a subsequent revi- species is described below as new. We have concluded sion of the American representatives of the subfamily that the type-species of the genus has been misidentified , Lemaire (1978) concurred with the taxo- by authors for the past 21 years, and nomenclatural nomic treatment of Agapema by Ferguson. With additional changes are now necessary. We have also concluded that literature we verified that Pope spent time in the south- the available name dyari Cockerell nom. rev. is applicable ern tip of Texas. On a visit to the Field Museum of to the population of far western Texas and southern Natural History in 1990 Peigler found a pair of syntype New Mexico. specimens (Figs. 1–2) in the collection of Ferdinand The following abbreviations are used in the text Heinrich Herman Strecker (1836-1901), revealing that for the collections from which material is cited in this the Agapema found in that region was what Pope had col- study, notably under the subsections for each species enti- lected. Labels in Strecker's handwriting (Figs. 3–4) read: tled "Material Examined. " Label #1: "Capt. Pope, Coll. J. G. Morris" Label #2: "Saturnia Galbina Clem. These two examples AMNH— American Museum of Natural History, are from old coll. of Dr. J. G. Morris, who got them from a New York City Capt. Pope. They are the originals of figs. 4, 5 pt. 12. Lep. CMNH— Carnegie Museum of Natural History, Rho. Het. [continued on reverse side:] they are doubtless Pittsburgh the examples from which Clem. drew his description. " CNC— Canadian National Collection, Bio- We believe these to be true syntypes. Hardwick systematics Research Centre, Ottawa (1978) discussed in detail that Strecker did not have a CSU— Colorado State University, Fort Collins clear type concept and some of the specimens he labelled DMNII— Denver Museum of Natural History, as types are not true type specimens. However, that prob- Denver lem apparently pertains more to names that Strecker pro- FMNH— Field Museum of Natural History, posed himself than to names proposed by other authors Chicago or to old historical specimens that he acquired and proba- LACM— Los Angeles County Museum of bly valued. Gall and Hawks (1990) also discussed type Natural History, Los Angeles material of Strecker, accepting most specimens as valid ROK— Roy O. and Conway A. Kendall and pointing to the accuracy of the illustrations in his Lepidoptera Collection serial publication (Lepidoptera, Rhopaloceres, and

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A REVIEW OF THE GENUS AGAPEMA (LEPID OPTERA: SATURNIIDAE)

Heteroceres), part 12 of which we cite, as it contains the One additional point that convinces us that the text and figures of galbina (Strecker 1875). John Rawlins lower Rio Grande Valley is the true galbina is (pers. comm.), who has intensively studied the Strecker Clemens' (1860) original description. Although most of collection, points out that many more type specimens that 149-word description could refer to most of the exist in that collection that are validly labelled than are species in the genus, the following characters cited by him invalidly labelled. pertain only to the lower Rio Grande species, if, as we To establish if Capt. John Pope visited the believe, the description refers to a male. "Antennae Brownsville area of Texas, it was necessary to consult vari- luteous. Forewings yellowish brown. The marginal por- ous historical documents. For military personnel today, it tion of the wing is whitish, and is tinged on the terminal would only be necessary to consult the military service edge with pale yellowish brown. " All of these characters record for the individual, but service records were not are darker in the other species. kept for Regular Army officers who served before 1863. We therefore cite Agapema solita Ferguson as a new John Pope served in the War with Mexico in 1846-1848 synonym of Agapema galbina (Clemens). We designate the (Cullum 1891: 126). male in the Strecker collection as lectotype of Saturnia gal- Pope was assigned to General Zachary Taylor ' s bina Clemens, and the associated female as paralectotype command. In March 1846 construction of Fort Brown (Figs. 1-2). We agree with Ferguson (1972) and Lemaire began. Pope participated in engagements at Buena Vista (1978) that the true galbina (which they called solita) is a dis- and Monterrey in Mexico (Cullum 1891). After the peace tinct species from the others that occur to the west and treaty with Mexico was signed 2 February 1848, Pope south. This leaves the species in the region of far western probably returned to Fort Brown to work on the Mexican Texas and southern New Mexico in need of a name, for Boundary Survey. This is evidenced by Scudder (1872: which the name dyari is available. Recent authors (Ferguson 68), who described the butterflies Phocides texana and P. 1972, Lemaire 1978) have agreed that all of the taxa that fly sanguinea (Hesperiidae), stating that they had been taken in Texas, Arizona, New Mexico, and Mexico are conspecific, by "Capt. Pope Mexican Boundary Survey from Texas." except for the much larger homogena Dyar. We believe that Although now considered synonyms of P. urania these so-called subspecies are best considered full species (Westwood) and P. palemon lilea (Cramer), respectively, based on the genitalic and particularly larval differences neither has ever been taken in Texas north of Cameron described in this paper. Taxa such as anona and dyari have and Hidalgo Counties. Pope's next assignment, surveying distributions extending hundreds of kilometers, but we a railroad route across the 32 nd Parallel in far western found no geographical variation among populations. Texas, began on 12 February 1854 (Goetzmann 1959), as The nomenclature and distributions of the noted by Ferguson (1972). It therefore appears that Pope species in this genus may thus be listed as follows, includ- was probably in the Brownsville area from late March to ing synonyms: early September 1846, and again late March 1848 to late A. galbina (Clemens), 1860—Cameron and Hidalgo December 1853 or even January 1854. Pope undoubtedly Counties, Texas; undoubtedly Tamaulipas collected the type specimens of galbina during these years A. solita Ferguson, 1972, new synonymy in the lower Rio Grande Valley. A. anona (Ottolengui), 1903, status revised- Strecker (1875) stated: "I received six examples, southern Arizona; south into Mexico (Sierra five males, one female, from south-western Texas, on the Madre Occidental) at least to Zacatecas border of the Rio Grande, but most of them before A. galbina ab. interrupta Draudt, 1929 coming into my possession had suffered to such an A. dyari Cockerell, 1914, new status—Chihuahua; extent, from ravages of mites or other insect depreda- Trans-Pecos region of western Texas and south- tors, as to be utterly worthless." The sex ratio of the type ern New Mexico series suggests that the specimens were collected at A. pelora (Rindge), 1961, new status—Baja light instead of reared from cocoons. The only pair that California Sur, Baja California Norte Strecker retained was also damaged, but his illustrations A. dentifasciata Lemaire, 1973, new status—Sierra do not reflect it. The salvaged pair that he kept and Madre Oriental: San Luis Potosi, Nuevo Leon, used as models for his illustrations were labelled as to Hidalgo their source. We figure this pair of specimens and the A. platensis new species—Edwards Plateau and label in Strecker's handwriting (Figs. 1-4). Strecker ' s South Texas Plains: Kinney, Val Verde, La omission of the words "From the Smithsonian Salle, Maverick, Zavala, Zapata, Dimmit, and Institution" when he cited Clemens' text, suggests that Mason Counties, Texas; undoubtedly Coahuila he had decided to retain the specimens himself. A. homogena Dyar, 1908—Rocky Mountains Ferguson (1972) noted that the types could not be locat- (northern Colorado to western Texas) along ed in the USNM and speculated that they may not have Sierra Madre Occidental (to Mexico City) been returned.

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PEIGLER AND KENDALL

SYNOPSES OF THE SPECIES 8. Ground color solidly dark blackish brown 9 Below are synopses for each of the seven species, - Ground color pale or smoky gray or brown presenting new information on distribution of most of 11 the species, and the first description of the female of denti- fasciata and larvae of three species. New data on host- 9. Eyespots of forewing touching postmedian plants and parasitoids are also given in subsequent sec- line dentifasciata tions. We do not provide detailed redescriptions of larvae - Eyespots of forewing not touching post- or adults for those taxa that were well-described and illus- median line 10 trated by Ferguson (1972) and Lemaire (1978). It should be noted that various shades of gray on the wings of all 10. Eyespots oval, apical red dash straight ..galbina species fade to brown or beige, and white fades to yellow- - Eyespots circular, apical red dash curved ish. These changes of color are obvious when comparing anona fresh material to older specimens, and should not be interpreted as taxonomic or even phenotypic differences. 11. Eyespots touching postmedian line, post- The moths may be separated by the following key. median area not dentate dyari - Eyespots not touching postmedian line; post- median area dentate platensis Key to the adults of Agapema (The female of pelora is unknown.) 1. Postmedian lines on forewing and hindwing distinct, light, narrow bands, sharply contrast- Agapema galbina (Clemens) ing with median and postmedian areas; anal Ferguson (1972) gave detailed descriptions and margin of hindwings tinted pink homogena excellent color figures of both sexes, including under- Postmedian lines scalloped, white, broad sides (see our Figs. 1-2, 13-14). The type-locality of solita bands, fading into median or postmedian is Esperanza Ranch. Packard (1914, pl. 59) figured a areas; no pink tint on hindwings 2 pair from Esperanza Ranch collected by Jacob Doll. The type-locality for galbina is Brownsville, Cameron 2. Antennal rami long, yellow or dark brown; County, Texas. forewings in form of isoceles triangle, falcate Both of us knew and visited the late Perry A. Glick males, 3 on several occasions. He was a long-term resident and — Antennal rami short, yellow; forewings usually professional entomologist in Brownsville who collected more rounded females, 8 Lepidoptera. He told us that this was formerly com- mon at lights, but he had not seen it since the late 1960s. 3. Antennae yellow galbina Glick and others have speculated (see Ferguson and — Antennae dark brown 4 Knudson 1986) that widespread insecticide applications were the primary cause of many populations of 4. Wing coloration blackish brown, with white Lepidoptera disappearing in the lower Rio Grande Valley. markings; outer edge of postmedian area very We favor alternative hypotheses. Peigler believes that gal- dentate 5 bina is one of those species limited by climate, so that Wing coloration gray brown or washed out; northernmost populations are exterminated by periodic outer edge of postmedian area not dentate or freezes and then must recolonize from the south. While a weakly so 6 resident entomologist in Texas who collected yearly in the Brownsville area for several years, Peigler observed 5. Median area of hindwing whitish anona the saturniid Rothschildia lebean (Guerin-Meneville) — Median area of hindwing darkly suffused (=forbesi Benjamin) to disappear in the mid 1980s after dentifasciata being common in the late 1970s and early 1980s. This hypothesis offers hope that galbina may return if there has 6. Eyespot in forewing centered in median area not been too much habitat destruction in Tamaulipas platensis south of the Rio Grande River (=Rio Bravo). However, - Eyespot in forewing touching postmedian line sanctuaries such as Bentsen-Rio Grande State Park and 7 Santa Ana National Wildlife Refuge may be among the few remaining areas in that region of Texas that have not 7. Apices of forewing and hindwing pointed or been altered by agriculture. Peigler has looked for abrubtly curved pelora Agapema in these preserved areas on several trips without — Apices of forewing and hindwing very success. Collins and Weast (1961) reported that in some rounded dyari years hundreds of galbina cocoons could be found,

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A REVIEW OF THE GENUSAGAPEMA (LEPIDOPTERA: SATURNIIDAE)

whereas none were found in other years. Obviously, agri- (California Insect Survey; M. J. Smith, pers. comm.). The culture and urbanization have been more significant than most southern and eastern distributional record for insecticides in reducing populations of this moth. anona is Zacatecas. In the field, Kendall observed masses of cocoons The eggs are probably the overwintering stage of galbina on ebony trees in a suburb of Brownsville (Tuskes and Collins 1981), although larvae can feed on being attacked and destroyed by large, mound-building the evergreen hostplants (see section on hostplants ants (not fire ants, Solenopsis) 19–22 March 1961. below) in early spring, which is January through March in Unfortunately, no ants were collected for future identifi- these areas. Females are obviously very adept at locating cation. The impact of ants on populations of this moth hostplants at very low density; R. D. Weast (pers. comm.) remains undetermined. located many cocoons on Condalia near Arivaca, Arizona, The moths fly in October and November. Eggs which were on a few widely scattered shrubs over a large hatch in December and January. Larvae are winter feed- area, growing among many non-hosts. M. J. Smith (pers. ers (Collins and Weast 1961), maturing just before comm.) reports that the moths emerge in July in Condalia sheds leaves and refoliates. The mature larva was Guadalupe Canyon, Arizona, whereas in nearby areas described by Ferguson (1972). with higher elevations they fly in October and November. Material examined — Larvae in alcohol, Peigler observed empty cocoons and half mature larvae in Brownsville, Texas, 16 November 1922, F. H. Benjamin the field in northern Durango on 6 March 1992. A male, (USNM); 7 December 1939 (USNM). One male, one the only resulting adult, emerged 15 July 1992. female, no locality, emerged October 1967 (DMNH). Edwards (1888) described eggs, newly hatched Two males, Brownsville, Texas, ex-pupa 26 October 1966 larvae, and cocoons of what was possibly this species, and "ca. 1967", P. A. Glick (DMNH). Two females, although he did not indicate the source locality of the paratypes of solita, San Benito, Texas, 5 and 7 October material. The mature larva was described by Ferguson 1955, Otto Buchholz (AMNH). Two males, one female, (1972). His description was based on material preserved Esperanza Ranch, Brownsville, Texas, Jacob Doll, Exch. in alcohol that we amend as follows based on a color slide A.N.S.P., C.M. Acc. 20359 (CMNH). Two males, two of a living larva sent to us by M. J. Smith, phis material females, "S. West U.S. " or "Texas", Marloff collection observed in the field. The subspiracular lines and subdor- (CMNH). One male, one female, Brownsville, Texas, sal stripes are snow white, in contrast to the yellow mid- Jacob Doll (CMNH). One male, two females, Brownsville, dorsal stripe that is formed by clusters of pinacula. These Texas (one labelled "August") (CMNH). One male, yellow markings are not arrowhead-shaped as they are in Brownsville, Texas, Frank Johnson collection (CMNH). dyari and platensis. The scoli are reduced compared to One male, Texas (CMNH). One male, no data (CMNH). many other Saturniinae, but are the largest within the One female, no field data, H. Edwards collection 1882 > J. genus Agapema. Sewer collection > Ehrman collection (CMNH). One Field observations made by Peigler in Durango female, Weslaco, Hidalgo County, Texas, 1939, P. T. were as follows. Of 30 cocoons observed on Condalia eri- Riherd (TAMU). One female, Cameron County, Texas coides, these were never massed, and in fact, no two were (CSU). One male, one female, Augusta, [erro- seen to be in contact. They were not nearer than within 8 neous, probably where transported cocoons emerged], 20 cm of each other on the same branch, and were always and 14 August, ex-larvae, Engel collection (CMNH). One positioned with the exit valve upward, from 45° to 900 male, San Benito, Texas, 1955, P. A. Glick (ROK). (vertical). Seven were seen on a single plant; most plants had 3-4 cocoons. Cocoons were attached 10-60 cm above Agapema anona (Ottolengui) ground level. One mature and several half-mature larvae This species (Figs. 19–20) is well-known and were seen on three shrubs. Their yellow markings pro- remains common, even in certain suburbs of Tucson, vided camouflage among the yellow flowers of the host- Arizona (M. J. Smith, pers. comm.). It was described and plant. These larvae ate flowers in preference to leaves. figured by Packard (1914, pl. 63), Ferguson (1972), Material examined — One female, San Bernardino Lemaire (1978), and other authors. The male figured by Ranch, 1144m, Cochise County, Arizona, August, F. H. Holland (1903: pl. 9, fig. 3) under the name A. galbina is Snow (KU). One male, Rancho Bonito, La Abra Valley, this species. Peigler searched for it in areas south of Organ Pipe Cactus National Monument, Arizona, C. F. Tucson in July 1986, but found only one empty cocoon. Harbison (FMNH). One male, Zacatecas [the state or Resident lepidopterists are able to collect it routinely. town?], Mexico, 8 September 1979, at light, Terry W. This insect also occurs in southwestern Arizona near No, Taylor (S. E. Stone collection). One male, Sierra Vista, where M. M. Collins (pers. comm.) observed males at Arizona, emerged 1992, cocoon sent to Peigler from R. D. light in late December at a ranger station in Organ Pipe Weast (U. and L. Paukstadt collection). One female, Cactus National Monument. It is recorded from the Bates Well, Pima County, Arizona, 10 November 1939, Mexican state of Durango, 34 km north of Durango City Engel collection (CMNH). Three males, four females,

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PEIGLER AND KENDALL

Redington, Arizona, Chrissman collector [probably "Mr. this taxon, but his drawing (Fig. 24a, b) of the genitalia of Morris Chrisman" cited in Packard 1914], W. J. Holland a male specimen from Del Rio should be assigned to collection (CMNH). Two males, Arizona, 1904, J. Doll platensis (see below). Ferguson (1972) did not have larval (KU). Larvae, empty cocoons, ca. 10 km SE of La material to describe; we provide a description as follows. Resolana, Highway 45, Meseta de la Zarca, 1900 m, Mature larva — I lead and integument black; tho- Durango, 6 March 1992, on Condalia ericoides, one male racic legs brownish black; prolegs brownish black with emerged 15 July 1992 (pupated 14 March), Ward S. Sear, orange lateral patch; dorsal scoli brownish orange, with John D. Boone, and R. S. Peigler (DMNH). Larva, six black projections, each with a short black spine; sub- Tucson, Arizona, July 1988 (DMNH). One male, base of dorsal scoli smaller and flatter than dorsal scoli, lighter Madera Canyon, Pima County, Arizona, ex-cocoon 29 scarlet, each with five black spines arising from a dark October 1975 (TAMU). One female, Tucson, Arizona, ex- spot; subspiracular scoli flattened with seven black projec- larva July 1988, reared 11 October 1990 from cocoon by tions, each with a long white seta; spiracles black; dorsum R. S. Peigler (DMNH). One male, one female, Highway black, covered by many small faint pinacula, some of 82 between mile markers 46 and 48, Cochise County, which fuse into patches or slender transverse lines and Arizona, emerged 21 and 15 November 1983 from short longitudinal dashes (i.e. subspiracular stripes); mid- cocoons on Condalia spathulata, Stephen E. Stone dorsal stripe is pinacular patches in a series of white (DMNH). One male, one female, Madera Canyon, Santa arrowhead-shaped markings, pointing posteriorly; sub- Rita Mountains, Santa Cruz County, Arizona, emerged 26 dorsal stripe broken, subspiracular stripes running con- December and 7 November 1972 from wild cocoons, tinuously and unbroken on each side; white setae origi- Kenneth C. Hansen (DMNI I). nating from pinacula; ventrum brown, covered with pinacula, each with a white seta, some setae with black Agapema dyari Cockerell median sections. This species (Figs. 5, 16–17) has been described Material examined — One female, roadside park and illustrated by several authors, including Ferguson 16 km south of Marathon, Brewster County, Texas, 16 (1972) and Lemaire (1978). For the past 21 years it has October 1983, caught in spider web on Condalia ericoides, been erroneously considered to be the true galbina, usu- R. O. and C. A. Kendall (ROK). Eight males, nine ally treated as the nominotypical subspecies. A pair from females, roadside park, 3 km southeast of Orla, Reeves Chihuahua was figured by Cockerell (in Packard 1914) County, Texas, 21 April 1981, reared from larvae and under the name anona on plate 59, but he proposed the cocoons on Condalia ericoides, adults emerged 16 and 30 name dyari for this pair in the text of the same work. August, 9 and 22 September, 4 October 1981, 13 August, Ferguson (1972) designated the female as lectotype (Fig. 25 October 1982, 9 and 11 October, 10 November 1983, 5) and C. Lemaire sent us a color slide of this specimen. R. O. and C. A. Kendall (ROK, DMNH). Adults, Highway Forewing length of wild collected males (at light or ex- 652, 3 km west of and 8 km east of Delaware River, wild cocoons) ranges from 25 mm to 32 mm. Culberson County, Texas, 21 April 1981, 52 larvae mainly Jimmie C. Coleman (pers. comm.) has seen and on C. ericoides, a few feeding on nearby Rhus microphylla, collected this species on several occasions, and offered plus one viable cocoon, adults emerged 3, 6, 7, 8, 9, 14, the following data: south of Guadalupe Mountains 25, 27, and 30 September 1981, 20 and 25 October 1982, National Park, Culberson County, Texas, intersection of 19 and 23 October 1983, 30 August 1985, R. O. and C. A. roads 62 and 54, 18–19 October 1982, five specimens Kendall (ROK). One male, Post, Garza County, Texas, 17 attracted to UV light; rest stop north of McKittrick November 1985, L. Simpson (ROK ex San Angelo State Canyon Road, Highway 62, Culberson County, 19 University). One male, Forsan, Howard County, Texas, 13 October 1982, saw 16 specimens; ca 1.5 km south of November 1981, D. Norton (ROK ex San Angelo State Sierra Blanca, Hudspeth County, Texas, 18 October 1982, University). One male, one female, San Angelo, Tom collected female at light, and several egg rings in field. Green County, Texas, 7 November 1987, P. Gordy (CSU). Localities and dates for collection by the late Five males, Antelope Lodge, Alpine, Brewster County, Andre Blanchard are: localities of Alpine, September Texas, 5 October 1981 at UV light, J. F. Doyle III (Doyle 1963, Government Springs at 1190 m, Dugout Wells, collection). Two males, no data, Sweadner collection Chisos Basin, and Grapevine Spring all in Big Bend (CMNH). One female, 5 km east of Hueco Inn, 1586 m, National Park, Brewster County, Texas, September 1963, Hueco Mountains, Hudspeth County, Texas, ex-pupa on September 1964, September 1965, October 1966; Shafter, catclaw 12 October 1960, Lee D. Miller (CMNH). One Presidio County, Texas, 16 and 19 October 1973; male, Old Fort Davis National Historic Site, Jeff Davis Carlsbad Caverns National Park, Eddy County, New County, Texas, 6 October 1969 at UV light, A. and M. E. Mexico, 19 September 1963, A. and May Elise Blanchard. Blanchard (ROK). One male, "West Texas", 13 October All of the specimens figured by Ferguson (1972, 1962, D. G. Ford, collector; Perry A. Glick collection col. pl . 13) under the name A. galbina galbina belong to (DMNH).

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A REVIEW OF THE GENUSAGAPEMA (LEPIDOPTERA: SATURNIIDAE)

Agapema platensis Peigler and Kendall, sp. n. Female — Differing from male as follows. This species (Figs. 7-11) is an endemic of the Forewing: length 33-34 mm; wingshape more rounded; Edwards Plateau and the northern portion of the South eyespots larger; median area of hindwing dusted with Texas Plains (Correll and Johnston 1979, Map 1). light gray-brown; stronger notching on outer edge of Through the efforts of state park superintendent Kelly B. postmedian area of hindwing. Bryan, we received much new material of adults, larvae, Diagnosis — Comparisons are made here to the and field observations. The only other name that has other five species in the galbina group. The large size and been proposed in the genus as currently defined is A. gal- tendency to have a lot of white areas, particularly on spec- bina ab. interrupta Draudt (1929); names proposed as imens that have flown, are characteristics shared particu- aberrations are not available according to the larly by platensis and pelora, and to a lesser extent by males International Code of Zoological Nomenclature (1985, of galbina. The female of platensis is the lightest in overall Art. 1(b) (5)). It is preferable to use a reared instead of a coloration known in the genus. The eyespots are centered wild-collected specimen to designate as the holotype more in the median area for platensis than in other because much of the distinctive maculation characteriz- species in which the eyespots usually contact the post- ing this species is lost after the moths have flown. median line. They also tend to be larger and more Interestingly, the same phenomenon of shedding wing rounded than in the other species. The darker antennae scales occurs in peigleri Lemaire, also an of the males of platensis separate them from the males endemic of the Edwards Plateau, a trait not shared by of galbina. most other members of its group (Peigler and Stone Mature larva — Head and integument black (dark 1989). Tuskes and Smith (1984) noted change in appear- brown in material in alcohol); thoracic legs black; prolegs ance from loss of wing scales between specimens of light brown with dark brown lateral patch; dorsal scoli homogena that had and had not flown. The pair of platensis carmine, with six black projections, each with a tiny black (from Lake Walk, near Del Rio, and Eagle Pass) reported spine; subdorsal scoli smaller and flatter than dorsal scoli, by Ferguson (1972: 190) were tentatively assigned by him lighter carmine, each with five black projections having a to the far western Texas taxon because he had an inade- black or white spine; subspiracular scoli flattened, not quate sample available for study. clearly delineated from surrounding area, pinkish, with Male — Head: frons with dense, long tuft of gray- seven to eight black projections, each with a long white ish black scales; antennae stramineous, with blackish sen- seta; spiracles black; dorsum blackish brown, covered by sillae on rami, giving antenna overall color of brown. numerous white pinacula, forming a middorsal series of Thorax and abdomen: covered by long, thin, grayish arrowhead-shaped patches, pointing posteriorly; pinacula black scales, interspersed with longer, thicker brown so numerous between subdorsal (broken) and subspirac- scales. Legs dark brown, tufted with long scales, the ular (unbroken) white lines that the area appears to be a longest ones lighter brown. Forewing: length 27-31 mm broad light band when viewed without magnification; (average 29 mm); antemedian area white, usually with a transverse white lines (two per segment) on dorsum gray-brown triangle or patch; antemedian disjunct, ca. prominent; white setae covering entire body, arising from 900; median area white, with diagnostic dusting of gray- pinacula; ventrum uniformly beige with faint pinacula brown throughout (lost when flown), space between ante- throughout. median and postmedian lines on anal margin variable; Material examined — HOLOTYPE: male, eyespot large, composed of black, blue, yellow rings, not Kickapoo Cavern State Natural Area, ca. 41 km north of touching postmedian line; postmedian line thin, broken, Brackettville, Kinney County, Texas, reared in San gray-brown; postmedian area dark gray-brown, weakly Antonio ex-ovo on Condalia hookeri by R. O. Kendall, notched on outer edge; marginal area white, tan, dark emerged 6 October 1989. ALLOTYPE: female, same data gray brown, with crimson and black marking at tip of as holotype, emerged 1 November 1989. The holotype forewing. Hindwing: Antemedian line obsolete; median and allotype are deposited in the Denver Museum of area large, solidly white; eyespot smaller than that of Natural History. PARATYPES: Kickapoo Cavern State forewing, of same color and pattern; postmedian line Natural Area, at light: Kelly B. Bryan, collector: one male, undulate and very faint; postmedian area dark gray- 3 October 1988; one male, 29 October 1988; one male, 10 brown, inner and outer edges notched, marginal area as October 1989; one male, 21 October 1989; one male, one in forewing; a few crimson scales sometimes visible, corre- female, 29 October 1989; three males, 1 November 1989; sponding to forewing marking. Genitalia: Saccus squared one male, 2 November 1989; one male, 5 November 1989; or rounded, never pointed or elongated. Uncus larger seven males, 6 November 1989; 14 males, one female, 7 than in dyari. Sharper, stouter, longer median tooth on November 1989; four males, 8 November 1989; one male, valve than in all other species except anona. Valve in pro- 11 November 1989; one male, 14 November 1989; one file very slender proximally. In general appearance, most male, 15 November 1989; four males, one female, 21 closely resembling pelora. November 1989; two males, 26 November 1989 (ROK).

Denver Museum of Natural History Series 3, No. 3, p. 7, October 15, 1993

PEIGLER AND KENDALL

Kickapoo Cavern State Natural Area, reared ex-ovo in San egg mass contained 102 ova. Extremely cold weather Antonio by R. O. Kendall on Condalia hookeri: one male, "burned" the leaves of the Condalia viridis, and the larvae emerged 1 October 1989; one male, 2 October 1989; one had to wait until new leaves budded out to continue feed- male, 3 October 1989; one male, 7 October 1989; one ing, and thus pupated in early April, about a month later male, 18 October 1989; one male, 22 October 1989; two than normal. The pupal stage of 20 reared individuals males, 24 October 1989; one male, one female, 25 ranged from 154 to 1,325 days. Flight in nature ranges October 1989; two males, 26 October 1989; one female, from 3 October to 3 December, peaking late October into 27 October 1989; two males, one female, 28 October mid-November (a bimodal pattern observed in 1990; see 1989; one female, 29 October 1989; one female, 1 below), depending on locality and climatic conditions. November 1989; one female, 7 November 1992; one Twenty specimens emerged 19 August to 1 November. female, 17 November 1992 (ROK). Kickapoo Cavern Between 12 October and 3 December 1990, D. Stuart and State Natural Area, reared ex-larvae on Condalia viridis by K Bryan counted 110 males and eight females attracted to K. B. Bryan: one female, emerged 19 August 1989; one a security light at Kickapoo Cavern State Natural Area. male, 24 October 1989; one male 26 October 1989 Males (numbers in parentheses) were observed on the fol- (ROK). 31 males, Devils River State Natural Area (=Dolan lowing dates in October: 12 (2), 16 (1), 17 (3), 18 (4), 20 Creek Ranch), Val Verde County, Texas, 27 October 1990 (3), 22 (2), 23 (2), 25 (14), 26 (1), 27 (7), 28 (5), 29 (3), at UV light, J. F. Doyle III (Doyle collection). One male, 31 (3); November: 12 (3), 13 (5), 14 (10), 15 (21), 16 one female, Kickapoo Cavern State Natural Area, 25 (11), 17 (1), 22 (7), 23 (1); December: 3 (1). Females October 1990, K. B. Bryan (Doyle collection). One male, were observed as follows: one each on 18 and 20 October, Dolan Creek Ranch, Highway 277/377 ca. 29 km north of and 17 and 22 November; two each on 25 October and Del Rio, Val Verde County, Texas, 29 October 1979, Dale 16 November. Benham (ROK). One male, near Crystal City, Zavala County, Texas, UV light, 1964, J. F. Raney (ROK). One Agape= pelora (Rindge) female, Seminole Canyon State Park, near Langtry, Val Rindge (1966) described this moth (Fig. 15) from Verde County, Texas, 27 September 1981 at light, Edward three males from three localities in Baja California Sur. C. Knudson (Knudson collection). Two males, Seminole The biomes for the three different localities where the Canyon State Park, Val Verde County, Texas, 20 October type series was taken were described in detail by Fox 1985, J. Rawlins and R. Davidson (CMNH). Two males, (1963). The female and immature stages remain Seminole Canyon State Park, Texas, 8 October 1989, G. unknown. Apparently, larvae were collected in the 1890s M. Chamberlain (TAMU). One male, Mason, Mason by the Frenchman L. Diguet in Baja California, but no County, Texas, 9 November 1969, B. L. Hofmann descriptions were published. Bouvier (1936) considered (TAMU). One male, Dimmit County, Texas, 23 October that material to be anona. Seven specimens loaned to us 1933 in light trap, S. E. Jones (TAMU). One male, Artesia from the SDNHM extend the known range much farther Wells, La Salle County, Texas, 11 November 1971, at UV north, and enable us to expand on the variation as follows. light, A. and M. E. Blanchard (USNM). One female, Forming length 30–35 mm (average 32.3 mm; n=10). The Eagle Pass, Maverick County, Texas, 5 October 1963, at largely (lark forewing more sharply contrasts with the UV light, A. and M. E. Blanchard (USNM). One male, white hindwing in the fresher material. The postmedian Del Rio, Val Verde County, Texas, 4 October 1963, at UV band of the hindwing tends to be narrower in the ones light, A. and M. E. Blanchard (USNM). from Baja California Norte than those from farther south. Etymology — This species is endemic to the It is interesting to note that the 1941 material was appar- Edwards Plateau and the adjoining section of the Southern ently collected by American armed forces personnel. Texas Plains. The name platensis means "of the plateau." Material examined — One male, Puerto Chileno, Field and lab observations — Information on host- Baja California Sur, 25 November 1961 (holotype) plants, structure and location of cocoons, and parasitoids (CMNH). One male, La Paz, Guaycura Hotel grounds, of this new species is given below under the respective sub- 4 December 1961 (paratype) (CMNH). One male, sections. K. B. Bryan made many observations in the field, Rancho Palmarito, 3 December 1961, at UV light, geni- and Kendall recorded data from rearings in San Antonio. talia slide no. 1455 D. C. Ferguson (paratype) (AMNII). Eggs deposited and kept outdoors in captivity on 20 One male, La Paz, 24 10 ' N, Baja California Sur, 22 October 1989 hatched on 24 November (incubation 32 October 1941, F. F. Gander (SDMNH). Three males, La days). David Stuart and K. Bryan located 20 egg rings dur- Paz, 6 and 8 November 1941, C. Harbison (SDNHM). ing mid-December 1989 and marked the location with Two males, 41 km northeast of El Arco, 700 m, Baja flagging tape. In mid-January 1990 these were checked California Norte, 11–15 December 1987, Norris with the following results: 16 had hatched; two were Bloomfield (SDNHM). One male, 16.4 km west of unhatched and discolored (possibly parasitized); two were Rancho Progresso (S. Francisquito), Baja California missing (possibly from predation). A large wild-collected Norte, 6–7 November 1987, N. Bloomfield (SDNHM).

Denver Museum of Natural History Series 3, No. 3, p. 8, October 15, 1993

A REVIEW OF THE GENUS AGAPEMA (LEPIDOPTERA: SATURNIIDAE)

Agapema dentifasciata Lemaire emerged 1 July, 7, 8, 22 August, and 2 September 1980, R. This species (Figs. 6, 12) is apparently endemic to O. and C. A. Kendall (ROK). Five adults, larvae, near the western slopes of the Sierra Madre Oriental in Mexico. Estancia Roberto, Highway 61, ca. 11, 26, and 39 km east- Lemaire (1978) recorded this species from the state of San northeast of San Roberto junction of Highway 57, Nuevo Luis Potosi, September 1966 (holotype), and 31 km south- Leon, 7 November 1978 ex-larvae and ex-cocoons, R. O. east of the city of San Luis Potosi, 1950 m, August 1974 and C. A. Kendall (ROK). One female, roadside park on (LACM). At the Matehuala locale cited below, ants identi- Highway 57, ca. 21 km east-southeast of Saltillo, Coahuila, fied as Liometopum apiculatum Mayr (Formicidae: 15 September 1977, R. O. and C. A. Kendall (ROK). Dolichoderinae) (det. D. R. Smith) were observed killing a pupa within a cocoon. A male of dentifasciata was illus- Agapema homogena Dyar trated by Sonthonnax (1904: pl. 12, Fig. 1) under the This species (Fig. 18) flies in the southern Rocky name Saturnia Galbina; Sonthonnax cited Mexico as part Mountains from Routt and Larimer counties, Colorado of the range of the species. Hoffmann's (1942) report of down the mountains through New Mexico, Arizona, west- galbina in the state of Ilidalgo should belong under this ern Texas, continuing along the Sierra Madre Occidental species, but his citation of the state of Durango must be to the vicinity of Mexico City (Hoffmann 1942, Ferguson referred to anona. Based on locality, we refer a badly dam- 1972, Lemaire 1978, Tuskes and Smith 1984). The species aged female cited below from Coahuila to this taxon. We was described from a pair of specimens, both figured by give descriptions below of the previously unknown female Packard (1914: pl. 59). The lectotype female, designated and larva. We also figure a male in fresh condition, better by Ferguson (1972), was collected 9 June 1894 in Fly Park, than the holotype figured by Lemaire (1978). 3050 m, Chiricahua Mountains, Arizona; the paralectotype Female — Head, legs, thorax, and abdomen cov- male was collected in March 1908 in the vicinity of Mexico ered with chocolate brown scales, with longer, strami- City by Roberto Miller. Resident lepidopterist Donald neous scales on thorax. Antennae: yellow. Forewing: Bowman has collected adults at light in Colorado as fol- Antemedian and median areas solidly gray brown, except lows: Golden, 1830 m, Jefferson County, 29 June; Lookout white on veins in median area; antemedian line broad, Mountain, 2135 m, Jefferson County, 2-30 June; Lawson, whitish, blurred, and angled at 90°; postmedian line Clear Creek County, 2440 m, 17-27 June; and larvae near whitish, straight, broad; eyespot large, oval, yellow, blue, Steamboat Lake, Routt County, in early July 1977. The black, contacting postmedian line; postmedian area moth was collected by David L. Eiler at light at an eleva- solidly gray brown, strongly and evenly notched on outer tion of ca. 2745 m in Cimarron Canyon State Park, Colfax edge; marginal area sharply contrasting white, brown, County, New Mexico, 11 June 1990 (News Lepid. Soc., gray brown; apex with crimson and black markings. Mar/Apr 1992, p. 13). All other known localities for this Hindwing: Antemedian area fading into median area, species were cited by Ferguson (1972) and Tuskes and light gray brown; eyespot interrupting postmedian line Smith (1984). The latter authors described and figured which is broad, whitish, angled, and less sharp than in the larval stages, of which there are only four instars. forewing; postmedian area dark gray brown, with faint Material examined — Two males, Silver Saddle whitish band, outer edge strongly and evenly notched; Motel, Boulder County, Colorado, 1678 in, 17 and 18 marginal area as in forewing. June 1961, M. R. MacKay (CNC). One male, 16 km west Mature larva — Ilead and integument black; tho- of El Salto, 2745 m, Durango, 11 April 1964, W. C. racic legs brownish black; prolegs brown, lighter distally; McGuffin (CNC). One male, Alto, near reservoir, Lincoln dorsal scoli scarlet, with six black projections, each with a County, New Mexico, 16 June 1965 at light, Willis and tiny white or black spine; subdorsal and subspiracular Atchley, collectors (KU). One female, 24 km west of scoli smaller and flatter than dorsal scoli, lighter scarlet, Aguilar, Las Animas County, Colorado, 2288 in, 25 June each with six long white setae, each arising from a dark 1982, at light, W. Huffman (DMNH). One male, Lawson, spot; spiracles black; dorsum black, covered by numerous Clear Creek County, 2440 in, Colorado, 18 June 1984, D. white pinacula, many of which fuse into patches or trans- Bowman (DMNII). One male, one female, one cocoon, verse and longitudinal bands; patches of pinacula indi- Bear Canyon Picnic Area, Santa Catalina Mountains, 1830 cate trace of middorsal stripe when viewed without magni- m, Pima County, Arizona, emerged 16 and 19 May 1984, fication; white subdorsal and subspiracular stripes, latter M. D. Van Buskirk (DMNH). Larva, General Hitchcock more prominent and unbroken; white setae covering Campground, 1708 m, Santa Catalina Mountains, Pima entire body, generally arising from scoli or pinacula; ven- County, Arizona, 17 August 1982 (preserved 28 trum very light brown, covered with pinacula, each with a September), on Rhamnus californica, M. J. Smith white seta, some setae with black median sections. (DMNH). One female, #556, Fort Collins, Colorado, 20 Material examined — Two males, two females, lar- June 1892, at light, C.F.B. (CSU). One male, Colorado, vae, Highway 60, 58 km east-northeast of Matehuala, David Bruce collection (CSU). One male, one female, Nuevo Leon, 7 March 1980 ex-larvae on Condalia ericoides, three cocoons, Glacier View Meadow, 2300 m, Larimer

Denver Museum of Natural History Series 3, No. 3, p. 9, October 15, 1993

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County, Colorado, 1 June 1989, ex-larvae on Ribes cereum, Although the genus Agapema ranges far down into H. E. and M. A. Evans (CSU). One male, one female, Mexico (Fig. 21), the map by Beutelspacher (1978) Glacier View Meadow, Colorado, 20 June 1991, at light, demonstrates that all the species technically belong to the H. E. and M. A. Evans (CSU). Nearctic faunal region, including the areas of Mexico City and the southern portion of Baja California. On the map PHYLOGENY by Brown and Lowe (1983), the following biotic provinces Previous authors have considered the genus are inhabited by the various Agapema. Agapema anona Agapema to be closely allied to Saturnia, a Holarctic group occurs in the Arizona Upland Subdivision of the Sonoran in the tribe Saturniini ranging across Eurasia and in west- Desertscrub, as well as Semidesert Grassland. On the Baja ern North America. Michener (1952) classified Agapema California Peninsula, pelora is an inhabitant of the as a subgenus of Saturnia, a plan rejected by all subse- Vizcaino Subdivision of the Sonoran Desertscrub. quent authors despite their agreement that the two Biogeography of that peninsula was discussed in detail by groups are closely related. The very hirsute larva of Fox (1963) and Brown et al. (1992). In western Texas and Agapema cannot be used as a character to support generic- southern New Mexico, localities where dyari flies occur level separation from other Saturnia because several among Semidesert Grassland and Chihuahuan Asiatic Saturnia also have very hirsute larvae (Bouvier Desertscrub. Thus, anona and pelora are members of the 1936, Nardelli 1986). The scoli on larvae of Agapema fall fauna of the Sonoran Desert; the area of Ajo and Organ into Type 2 of Nassig (1989), having short, stout spines, in Pipe Cactus National Monument has a biogeographical agreement with most other species in the Saturnia com- link with Baja California (Halffter 1976), yet the Agapema plex. Ferguson (1972) cited several differences in adult there is anona, not pelora. Agapema dyari and platensis morphology to justify maintaining Agapema as a genus belong to the Chihuahuan Desert (see maps in Helms separate from Saturnia. Despite the fact that two species 1983: 6-7). The area in northern Durango State where of Copaxa (copaxoides (Dyar) and muellerana (Dyar)) were Peigler collected anona was a level, grassland plateau with originally described as belonging in Agapema (see Condalia ericoides as a dominant shrub. Lemaire 1978), we consider Agapema to be more closely On similar maps delineating ecoregions by allied to the Saturnia-complex of genera. Omernik and Gallant (1987), the range of galbina falls Hogue et al. (1965) considered that Agapema within the Southern Texas Plains. Correll and Johnston and the Nearctic Saturnia were all derived from a com- (1979) cited South Texas Plains, Rio Grande Plains, and mon ancestor that dispersed across the Bering land mass, Tamaulipan Brushlands as synonyms. Common plants of with subsequent divergence into two genera (or subgen- this biome include mesquite (Prosopis spp.), several era) and several species. Tuskes and Collins (1981) species of Acacia, and cenizo (Leueophyllum frutescens accepted this hypothesis but Ferguson (1972: 187) ques- (Berl.) I. M. Johnst.) (Vora 1990). Agapema dyari occurs in tioned it. The genus Agapema itself is undoubtedly a holo- the Southern Deserts and the Western High Plains phyletic group. We propose that the most divergent (Omernik and Gallant) or the Trans-Pecos Mountains species, i.e. homogena, represents the sister-group to the and Basins (Correll and Johnston). Agapema platensis is remaining species, i.e. the galbina complex. This hypothe- endemic to the Edwards Plateau and the Southern Texas sis is supported by the obvious differences in phenology, Plains (Correll and Johnston) or the Central Texas wing pattern and size, suite of hostplants, larval structure, Plateau and northwestern part of the Southern Texas broader geographical distribution, and montane habitat Plains (Omernik and Gallant). Plants that are common in of homogena. these regions of central and western Texas include and Condalia. The latter authors put localities for anona ECOLOGICAL OBSERVATIONS among Southern Deserts and Southern Basin and Range Habitat and biogeography—The montane harnogena ecoregions. Creosote bush (Larrea tridentate (DC.) Gov.) occurs at higher altitudes than do populations of other is a dominant plant in these areas. Cacti of the genus species in the genus. In southern Arizona, the altitudinal Opuntia are also common throughout all of these habi- ranges of homogena and anona are separated by an entire tats. The distributions of Agapema in Texas parallel those life zone ( woodland zone) where neither species of species of Henileuca: peigleri on the Edwards Plateau, occurs (M. J. Smith, pers. comm.). The result is that all slosseri Peigler and Stone on the Western High Plains, taxa in the genus are allopatric, either geographically (the grotei Grote and Robinson in both regions, and conwayae various representatives of the galbina complex) or by altitu- Peigler in the Trans-Pecos Mountains and Basins dinal habitat preference (homogena compared to all the (Kendall and Peigler 1981, Peigler and Stone 1989). others). The ecosystem for homogena is subalpine forest; The montane biomes for homogena differ consid- pines (Pinus spp.) are dominant in the areas. For all other erably from those of the species in the galbina complex. species, the ecosystems are grassland plateaus, chaparral, These areas are generally higher in altitude and are char- or deserts. acterized by oak and pines, from Colorado to Mexico

Denver Museum of Natural History Series 3, No. 3, p. 10, October 15, 1993

A REVIEW OF THE GENUSAGAPEMA (LEPIDOPTERA: SATURNIIDAE)

City. Tuskes and Smith (1984) listed associated plants ing. The color of cocoons of all species in the genus is in the Arizona localities for this moth; in Colorado the golden brown. Exposed cocoons are often bleached to dominant plants, in addition to the oaks and pines, near white, and we agree with Tuskes and Smith (1984) include Rhus aromaticaAit., Cercocarpus rnontanus Raf., and that moisture darkens the cocoons. The cocoons of all two species of Ribes. (Salix) and cottonwood species in the galbina complex are probably indistinguish- () also grow commonly in all of these areas if in able, but those of homogena tend to be browner and more riparian zones. slender and compact, with less open netting. A pre- Speciation in the galbina complex appears to have formed exit is at the anterior end, as in all Saturniini. resulted from isolation. The Sierra Madre Oriental sepa- The location of the cocoons in the habitat varies rates the ranges of galbina and dentifasciata. Likewise, the with the species. Tuskes and Smith (1984) wrote that the central plateaus in north-central Mexico may form a bar- cocoons of homogena are formed "in cracks or crevices rier between the range of the latter species and that of among rocks, tree trunks, or man-made structures." In anona which ranges down the eastern slopes of the Sierra galbina, the cocoons are massed on the bark of the ebony Madre Occidental. The fact that anona has not been trees, usually on the main trunk up near the first large found on the western slope of the range, including areas branches (see subsection below on hostplants). The that are well-collected, supports the hypothesis that cocoons of anona and dyari may be clustered on the main mountains form effective barriers to dispersal. Mountains trunks deep within the thorny hostplant, or scattered over in southern New Mexico down into Chihuahua undoubt- the entire hostplant singly or in groups of two to four, but edly separate the ranges of dyari and anona. always on main stems surrounded by smaller thorny Eggs — The ova of Agapema are laid in clusters stems. K. B. Bryan (pers. comm.) has collected hundreds around twigs of the hostplant. This unusual pattern of of cocoons of platensis in the field, and reports that they oviposition is shared by other saturniids that are ecologi- are more often at ground level than those of dyari. The cally (i.e. Hemileuca) or phylogenetically (i.e. Saturnia cocoons of platensis are usually at the base of the trunk of pavonia (L.)) allied to Agapema. the hostplant, sometimes massed at the first crotch on the Larvae — The caterpillars of Agapema closely main stem, often under leaf litter on the ground below resemble those of certain Lasiocampidae, as noted by the hostplant, or even on fenceposts and between lime- Ferguson (1972: 189), having black integuments with stone rocks near the hostplants. The Kendalls found white and red markings, gracile form, and long and dense cocoons of dentifasciata on Condalia in clumps, with a few hair; this resemblance is certainly convergent but not positioned singly up to 25 cm from the main clumps. mimetic. As in many saturniids, larvae of Agapema are well As far as is known, all cocoons of Copaxa are retic- camouflaged on their hostplants, yet appear colorful and ulate (Wolfe 1993), either single or double. In the structurally ornate when not on their hostplants. M. M. Saturnia complex three main types of cocoons exist per- Collins (pers. comm.) reports that the larvae of homogena taining to gross structure: single, reticulate (Saturnia are conspicuous, and suggests that this plus their gregari- jonasi (Butler), S. japonica (Moore), ous behavior may indicate they are distasteful. (Kupido)); double, reticulate (Saturnia mendocino Since the larvae of Agapema are winter and early Behrens, S. walterorum Hogue and Johnson); or of a solid spring feeders (galbina complex) or live at high altitudes non-reticulate structure (Saturnia naessigi (de Freina) stat. (homogena), the black integument serves to absorb heat et comb. nov., S. pyri, S. albofasciata (Johnson), S. pavonia, from the sun augmenting the ability of the larvae to be S. lindia Moore) (see Nassig 1983). Cocoons of species in active at lower temperatures. The young larvae of the the S. pyretorum Westwood species-group may be either many species of Hemileuca are also black, facilitating ther- solid (Taiwan) or reticulate (Assam). Cocoons of the dis- moregulation in the early spring. Like larvae in that tantly related Ceranchia (Saturniinae: Saturniini) from genus, the younger larvae of Agapema are gregarious, but Madagascar are gold colored, double, and reticulate, thus when in the last instar scatter over the entire hostplant or greatly resembling those of Agapema except that cocoons across the ground to other hostplants. The hostplants of Ceranchia are much larger (ca. 10 cm long). (Condalia) of Agapema are in effect evergreen, since new Seitz (1927: 315), in mentioning the perforated leaves emerge before all of the old ones have fallen. cocoon of a rainforest saturniid, postulated that the func- Cocoons — The cocoons of Agapema are double, tion of reticulation is drainage. Some of the Asiatic both the inner and outer cocoon being reticulate. The Saturnia and Neotropical Copaxa occur in biomes with pupa can be seen within the cocoon. The weave of the heavy rainfall, yet others having reticulate cocoons are outer cocoon is much coarser and more irregular than found in drier temperate climates and arid regions, that of the inner one. The amount and quality of the host- Agapema being good examples. Alternatively, it has been plant available in the last larval instar greatly influences postulated that perforations in cocoons may provide ven- the size and presence of the outer cocoon, because in tilation (Tuskes and Collins 1981), but the fact that retic- captivity the outer cocoon is often poorly formed or lack- ulated cocoons occur among species in very cold climates

Denver Museum of Natural History Series 3, No. 3, p. 11, October 15, 1993

PEIGLER AND KENDALL

(e.g., Siberia; mountains of Colorado) discredits this idea. Larvae of galbina feed on Condalia hookeri, proba- Although no satisfactory ecological explanation for retic- bly the only species in this genus of plants that is available ulation in cocoons is available, this type of cocoon appar- in the lower Rio Grande Valley. Larvae and cocoons of ently confers survival advantages in view of the fact that so platensis have been found in nature only on Condalia many species have this feature in many biomes on several viridis (K. Bryan, pers. comm.), but will feed on C. hookeri continents. M. M. Collins (pers. comm.) postulates that in captivity and that is the only species of Condalia that the reticulation could be a preadaptation to invade grows in Mason County, where platensis is recorded to deserts, and serve to both drain and ventilate. occur. The Kendalls collected larvae and cocoons of denti- The tendency to remain in pupal diapause two or fasciata on C. ericoides in Nuevo Leon and completed the more years is not common among Saturniidae. It occurs rearing in captivity on C. hookeri. In Arizona, larvae of in Agapema, plus groups that are phylogenetically related anona have been collected in nature on Condalia lycioides, ( (Denis and SchiffermCdler) and S. cephalar- C. spathulata, and C. ericoides. Peigler observed this species iaeRomanoff, W. A. Nassig, pers. comm.) and the ecologi- on C. ericoides in northern Durango. No hostplant records cally related Hemileuca (Hemileucinae). Diapause of two have been reported for pelora. or more years also sometimes occurs among Attacini A few reports of hostplants for Agapema in the lit- (Saturniinae). Unfortunately, virtually all data on this erature are erroneous. Weast (1962) reported Lycium, phenomenon are derived from material in captivity Solanaceae, as a host for anona in southern Arizona, and under artificial conditions. this has been perpetuated by several subsequent authors. A remarkable case of convergence is also seen in Throughout the world, Saturniidae do not feed on Olceclostera seraphim (Dyar) (Apatelodidae) in western Solanaceae, even polyphagous species, as plants in this Texas (Culberson County), the golden, reticulated family are well-known to contain alkaloids and other cocoons of which could easily be mistaken for Agapema chemical defenses. Godfrey (1984), in correcting the except that they occur on desertwillow (Chilopsis linearis same hostplant record for a notodontid moth in Arizona, glutinosa (Engelm.) Fosberg, Bignoniaceae), which is not pointed out that Condalia lycioides and Lycium look almost a hostplant for Agapema. The cocoons of Olceclostera angel- identical. The name lycioides means "looks like Lycium. " ica (Grote), which are found in western Texas (Val Verde "Greasewood" has been cited as a hostplant for County) on their hostplant Gregg ash ( greggii anona in Arizona in much of the earlier literature Gray, Oleaceae), also closely resemble those of Agapema. (Holland 1903, Packard 1914, Draudt 1929-1930). Later Hostplants — All valid records of hostplants for authors have assumed this to be either Sarcobatus vermicu- Agapema are summarized in Table 1. Agapema homogena latus (Hook.) Torr., Chenopodiaceae (Ferguson 1972, feeds in nature on Rhamnus californicus var. ursina in Stone 1991, as maximiliani Nees) or Larrea, Zygo- Arizona (Tuskes and Smith 1984), and Salix exigua (D. E. phyllaceae (Arnett and Jacques 1981, as Laria). The latter Bowman, pers. comm.) and Ribes cereum in Colorado plant is more correctly called "creosote bush" (Correll (Stone 1991). It has been reared successfully in captivity and Johnston 1979). We doubt these records and do not on Rhamnus frangula and Ribes alpinum (Stone 1991), include them in Table 1. All are likely based on a single both Eurasian plants. Collins and Weast (1961) first cited misidentification of Condalia at the turn of the century Ribes as a food for Agapema but did not specify which moth (see Packard 1914: 159). or plant species. They also cited wild cherry (, Some authors have cited "ebony" or "ebonywood" Rosaceae) and privet (Ligustrum, Oleaceae) as hostplants (i.e. Collins and Weast 1961) as a hostplant for galbina in for rearing in captivity for the genus Agapema. With the the lower Rio Grande Valley. This confusion results from specialization on Rhamnaceae by the genus, we might the fact that the larvae move from Condalia hookeri, the expect to find larvae of homogena on Ceanothus fendleri A. true hostplant, to ebony trees to make their cocoons in Gray, a common rhamnaceous plant in the foothills and masses on the trunks, as pointed out by Ferguson (1972) mountains of Colorado and Arizona. and observed by the Kendalls. Ebony is Pithecellobium ebano All other species of Agapema feed almost exclu- (Berlandier) C. H. Muller, Leguminosae, of which P. flexi- sively on Condalia (=Microrhamnus) as far as is known, caule (Benth.) Coult. is a junior synonym (Everitt and with a few exceptions. Larvae of dyari were found in Drawe 1993). There is a report of larvae of anona being Culberson County, Texas, by R. O. and C. A. Kendall to reared from dwarf hackberry (Ferguson 1972). Based on be feeding on Rhus microphylla but a larger number of lar- the description of Celtis pallida Torr., Ulmaceae, in vae were eating Condalia ericoides within a few meters. Kearney and Peebles (1942: 228) and the figure in Everitt Kendall offered larvae (eggs from a female taken at and Drawe (1993: 176), we believe that this record proba- light) of dyari from Jeff Davis County the following four bly also represents a misidentification of Condalia. Rhamnaceae, but only the last two were accepted: Kendall offered larvae of dyari leaves of Celtis laevigata Karwinskia humboldtiana, Colubrina texensis, Condalia eri- Willd. in 1966 which were refused. Many desert shrubs coides, and Condalia hookeri. have tiny rounded leaves and spiny stems, so some

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A REVIEW OF THE GENUS AGAPEMA (LEPIDOPTERA: SATURNIIDAE)

misidentifications by entomologists are understandable. (4) January, and 1 (4), 2 (1), 11 (3), 14 (1), and 19 (1) As discussed by Tuskes and Collins (1981) February. Both sexes were obtained (in some species of Saturnia (California and Eurasia) and Agapema differ in Anastatus only females are known) . their suite of hostplant preferences. This is basically valid, Peigler obtained or reared the following para- but both groups occasionally use Anacardiaceae, as seen sitoids, records of which are previously unpublished. in the following examples. Saturnia walterorum feeds on Voucher specimens for all of these below are in the Rhus in nature, and in captivity on Schinus (Stone 1991) Denver Museum of Natural History: and Pistacia chinense Bunge (Peigler, unpubl.). Claude Cocoons of anona collected in March 1992 by Lemaire (pers. comm.) collected larvae of S. pavonia in Peigler in northern Durango contained reddish puparia the wild in southern France on Pistacia terebinthus L. in (already emerged) of an unidentified tachinid. The struc- 1982. The Kendalls found larvae of A. dyari feeding on ture and pattern of the spiracular plates indicate this is Rhus microphylla in the field. There is also overlap with likely the same species cited above as "Lespesia sp. near tex- Rhamnaceae, as S. atlantica Lucas and S. albofasciata ana. " There are also some of these tachinids plus their (Johnson) are recorded to feed on species in this plant puparia in the Denver Museum of Natural History, associ- family (Stone 1991), the main group used by Agapema. ated with 1967 moths of galbina. Lespesia samiaewas reared Parasitoids — The following parasitoids have from anona from the vicinity of Tucson, based on the key been recorded to attack Agapema: Euphorocera sp. near to adults by Sabrosky (1980) and the deep pits and rugose floridensis (Buck and Keister 1955, 1956; Arnaud 1978), rims of the spiracular plates of the puparia. These were presumably from galbina since Buck and Keister cited reared from host cocoons in 1984 collected on Condalia "southern Texas" as the source of their stock; Lespesia sp. spathulata by S. E. Stone. Twelve cocoons of anona from near texana, from homogena in Arizona (Peigler 1985, Sierra Vista, Arizona, sent to Peigler by R. D. Weast see also Sabrosky 1980, Tuskes and Smith 1984); Lespesia yielded five adults of Lespesia samiae in early December sp. (archippivora group) from anona in Arizona (Peigler 1992. Of the 12 cocoons, eight were parasitized, most con- 1985); Cotesia electrae, from galbina or anona (Collins and taining a single parasitoid; some flies emerged before Weast 1961, as Apanteles electrae); Enicospilus texanus receipt of the lot. In Lespesia, the maggots sometimes in dyari from Reeves County, Texas, collected by Kendall form puparia between the inner and outer cocoons of the (Gauld 1988); and an unidentified ophionine ichneu- host, but usually within the inner cocoon. monid from anona in Arizona, almost certainly E. A male of Enicospilus texanus was reared from a texanus, hyperparasitized by Microdontomerus fumipennis cocoon of anona from Tucson, Arizona, in summer 1989 (Peigler 1985). by Peigler. The Arizona specimens of texanus reared from Kendall reared the following parasitoids, records Agapema and Hemileuca are darker than the ones from of which are previously unpublished: Texas, as noted by Gauld (1988). Specimens of Cotesia elec- Cotesia electrae reared from larvae of dyari from trae were reared from larvae of homogena in Tucson, Reeves County, Texas. The 92 parasitoids (numbers in Arizona, 2-5 April 1988. parentheses) emerged on the following dates in April All of the above records for parasitoids attacking 1981: 21 (3), 22 (17), 23 (8), 24 (3), and 25 (61). These Agapema are listed in Table 2. When records of saturniid tiny black wasps emerge from white cocoons that are hosts and parasitoids are viewed collectively, it is notewor- attached to the back of the host larvae. There are usually thy that some parasitoids that attack species in the sat- several individuals per host. Nine wasps were reared from urniid subfamily Hemileucinae (such as , three larvae of platensis from Kinney County, Texas, on Hemileuca, ), also attack Agapema. Those that 20-22 February 1989, three per host. attack species in the saturniid subfamily Saturniinae (such Parasitoid cocoons of Enicospilus sp., probably tex- as Hyalophora, Antheraea, and Actias) generally do not anus were found in dentifasciata 58 km east-northeast of attack Agapema. Since Agapema belongs to the subfamily Matehuala, Nuevo Leon by R. O. and C. A. Kendall. No Saturniinae, host selection by the parasitoids thus appears adult parasitoids were obtained. The female of E. texanus to be determined more by the ecology than the phy- that Gauld (1988) reported from dyari was collected in logeny of the host. This phenomenon pertains to certain April 1981 and emerged 25 January 1983. parasitoids in the Ichneumonidae, Braconidae, and possi- Lycogaster pullata nevadensis (det. A. S. Menke), bly others, but not Tachinidae. Moreover, Enicospilus tex- one wasp emerged 9 April 1980 from cocoon of dentifasci- anus has also been reared from the cocoons of Olceclostera ata collected 58 km east-northeast of Matehuala, Nuevo seraphica by the Kendalls, which, as mentioned above, are Leon. Specimen in Kendall collection. Wasps in this small structurally and therefore ecologically like those of family are generally hyperparasites of tachinids. Agapema. This is the only known host for texanus that is Anastatus reduvii were reared from eggs of platensis not a saturniid (see Gauld 1988), but seraphica belongs to from 41 km north of Brackettville, Texas, and eclosed as the superfamily Bombycoidea, as does Saturniidae. It also follows in 1989 (numbers in parentheses): 26 (6) and 27 has a hirsute larva like those of Agapema. Some species of

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PEIGLER AND KENDALL

the hemileucine genus Automeris are ecologically more history, conservation biology. Lepidoptera Research like Saturniinae, and are thus attacked by the parasitoids Foundation, Beverly Hills, Calif. v + 129pp., 8col. pls. associated with Saturniinae (Gauld 1988). BUCK, J. and M. KEISTER. 1955. Cyclic CO2 release in diapausing Agapema pupae. Biological Bulletin ACKNOWLEDGMENTS 109:144-163. We were loaned valuable material by David K. 1956. Host-parasite relations in Agapema pupae Faulkner (San Diego Natural History Museum), Douglas (Lepidoptera, Saturniidae). Annals of the C. Ferguson (United States National Museum of Natural Entomological Society of America 49:94-97. History), John E. Rawlins (Carnegie Museum of Natural CLEMENS, B. 1860. Contributions to American lepi- History), Frederick H. Rindge (American Museum of dopterology — No. 4. Proceedings of the Academy Natural History), and Boris C. Kondratieff (Colorado of Natural Sciences of Philadelphia 12:156-174. State University). Peigler was allowed access to study col- COLLINS, M. M. and R. D. WEAST. 1961. Wild silk moths lections by Charles D. Michener (University of Kansas), J. of the United States, Saturniinae. Collins Radio Co., Donald Lafontaine (Canadian National Collection), and Cedar Rapids, la. iii + 138pp. entomology curators at the Field Museum of Natural CORRELL, D. S. and M. C. JOHNSTON. 1979. Manual of History. Michael M. Collins, J. Rawlins, and F. Rindge pro- the vascular plants of Texas. Second ed. Texas vided constructive reviews of the manuscript. Additional Research Foundation, Renner. xv + 1881pp., 1 color data and specimens were kindly provided by the late map. Andre Blanchard, Donald E. Bowman, Jimmie C. CULLUM, G. W. 1891. Biographical register of the offi- Coleman, the late Perry A. Glick, Michael J. Smith, and cers and graduates of the U. S. Military Academy at Stephen E. Stone. Particular thanks are extended to Kelly West Point, N. Y., from its establishment, in 1802, to B. Bryan for providing abundant material and field obser- 1890 with the early history of the United States vations, enabling us to verify the correct identities of all Military Academy. Vol. 2: Nos. 1001-2000. Third ed. the species in Texas. Material collected by the junior Iloughton, Mifflin and Co., Boston. author was under Permit No. 26-89 with approval of David DRAUDT, M. 1929-1930. Saturnidae [sic], Pages 713-827, H. Riskind, Director of Natural Resources Program, col. pls. 101--142, in A. Seitz, ed., Die Gross- Texas Parks and Wildlife Department; that of J. F. Doyle Schmetterlinge der Erde, vol. 6: Die amerikanischen was collected under Permit No. 28-90. J. E. Rawlins sup- Spinner mid Schwarmer. A. Kernen, Stuttgart. plied the photographs of the syntypes of Saturnia galbina. EDWARDS, H. 1888. Early stages of some North Nancy Jenkins and Rick Wicker (Denver Museum of American moths. Entomologica Americana 4:61-62. Natural History) produced other photographs in this EVERITT, J. H. and D. L. DRAWE. 1993. Trees, shrubs, paper. Some of the parasitoids were identified by tax- and cacti of south Texas. Texas Tech University Press, onomists at the Systematic Entomology Laboratory, U. S. Lubbock. x + 213pp. Department of Agriculture (Beltsville, Maryland). FERGUSON, D. C. 1972. Bombycoidea, Saturniidae, fasc. 20.2B, in R. B. Dominick et al., eds., The moths of LITERATURE CITED America north of Mexico, pp.153-269, pls.12-22. E. W. ARNETT, R. H., Jr. and R. L. JACQUES, Jr. 1981. Simon Classey, London. and Schuster ' s guide to . Simon and Schuster, . and E. C. KNUDSON. 1986. Four new United New York. 511 pp. States records of moths from Texas. Journal of the BEUTELSPACHER, C. R. 1978. Familias Sphingidae y Lepidopterists' Society 40: 353-354. Saturniidae (Lepidoptera) de Las Minas, Veracruz, FOX, R. M. 1963. Reports on the Margaret M. Cary and Mexico. Anales del Instituto de Biologfa, Universidad Carnegie Museum expedition to Baja California, Nacional Autonoma de Mexico 49 (Ser. Mexico, 1961. Annals of Carnegie Museum Zool.) :219-229. 36:181-192. BOUVIER, E.-L. 1936. Etude des Saturnioides normaux, GALL, L. F. and D. C. HAWKS. 1990. Systematics of famille des Saturniides. Memoires du Museum moths in the genus Catocala (Lepidoptera: National d'Histoire Naturelle, Paris (n. ser.) vol. 3, Noctuidae). I. Type material in the Strecker collec- fasc. 1:1-354, 12pls. tion, with lectotype designations. Fieldiana, Zoology, BROWN, D. E. and C. H. LOWE. 1983. Biotic communi- N. Ser., No. 59: iii + 16pp. ties of the Southwest (large color map). General GAULD, I. D. 1988. The species of the Enicospilus ameri- Technical Report RM-78. Rocky Mountain Forest canus complex (Hymenoptera: Ichneumonidae) in and Range Experiment Station, Forest Service, U. S. eastern North America. Systematic Entomology Dept. Agric. 13:31-53. BROWN, J. W., II. G. REAL and D. K. FAULKNER. 1992. Butterflies of Baja California: faunal survey, natural

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A REVIEW OF THE GENUSAGAPEMA (LEPIDOPTERA: SATURNIIDAE)

GODFREY, G. L. 1984. Notes on the larva of Cargida NEUMOEGEN, B. and H. G. DYAR. 1894. Preliminary pyrrha (Notodontidae). Journal of the Lepidopterists ' revision of the Bombyces of America north of Society 38: 88-91. Mexico. Journal of the New York Entomological GOETZMANN, W. H. 1959. Army exploration in the Society 2:109-132, 147-174. American West 1803—1863. Yale Univ. Press, New OMERNIK, J. M. and A. L. GALLANT. 1987. Ecoregions Haven, Conn. 509pp. of the southwest states; Ecoregions of the south cen- HALFFTER, G. 1976. Distribution de los insectos en la tral states. (color maps, 1987-795-479). United States Zona de Transition mexicana; relaciones con la ento- Environmental Protection Agency. mofauna de Norteamerica. Folia Entomologica PACKARD, A. S. 1914. Monograph of the bombycine Mexicana 35:1—64. moths of North America, part 3 (edited by T. D. A. HARDWICK, D. F. 1978. An analysis of the heliothidine Cockerell). Memoirs of the National Academy of types (Noctuidae) of Herman Strecker with lectotype Sciences 12:ix + 1—516. designations. Journal of the Lepidopterists ' Society PEIGLER, R. S. 1985. Recent records for parasitism in 32:49-54. Saturniidae (Lepidoptera). Nachrichten des HELMS, C. L. 1983. The Sonoran Desert. KC Entomologischen Vereins Apollo, Frankfurt (N. F.) Publications, Las Vegas, Nev. 48pp. 5:95-105. HOGUE, C. L., F. P. SALA, N. McFARLAND and C. . and S. E. STONE. 1989. Taxonomic and biologi- HENNE. 1965. Systematics and life history of Saturnia cal notes on the Hemileuca rnaia (Saturniidae) com- (Calosaturnia) albofasciata in California (Saturniidae). plex with description of a new species from Texas and Journal of Research on the Lepidoptera 4:173—184. New Mexico. TyO to Ga 40:149-166. HOFFMANN, C. C. 1942. Catalogo sistematico y zoo- RINDGE, F. H. 1966. Reports on the Margaret M. Cary- geografico de los Lepidopteros mexicanos, tercera Carnegie Museum expedition to Baja California, parte: Sphingoidea y Saturnioidea. Anales del Mexico, 1961, 4. The family Saturniidae Instituto de Biologia, Universidad Nacional (Lepidoptera). Annals of Carnegie Museum Autonoma de Mexico 13:213—256. 38:137-142. HOLLAND, W. J. 1903. The moth book. Doubleday, SABROSKY, C. W. 1980. A revised key to the Nearctic Page, and Co., New York. xxiv + 479pp., 48co1. pls. species of Lespesia (Diptera: Tachinidae). Annals of KEARNEY, T. H. and R. H. PEEBLES. 1942. Flowering the Entomological Society of America 73:63-73. plants and ferns of Arizona. United States SCUDDER, S. H. 1872 [1871]. A systematic revision of Department of Agriculture, Miscellaneous some of the American butterflies: with brief notes on Publication 423:1069pp., 29phot. pls. those known to occur in Essex County, Massachusetts. KENDALL, R. O. and R. S. PEIGLER. 1981. Hemileuca Peabody Academy of Science, Fourth Annual Report grotei (Saturniidae): its morphology, natural history, for the year 1871:24-84. spacial and temporal distribution. Journal of the SEITZ, A. 1927. Saturnidae [sic], Pages 313-316, in A. Lepidopterists ' Society 35:41—50. Seitz, ed., Die Gross-Schmetterlinge der Erde, vol. 14: LEMAIRE, C. 1978. Les Attacidae americains...The Die afrikanischen Spinner and Schwarmer. A. Attacidae of America (=Saturniidae) Attacinae. C. Kernen, Stuttgart. Lemaire, Neuilly-sur- Seine, France. 238pp., 49p1s. SONTHONNAX, L. 1904. Essai de Classification des MICHENER, C. D. 1952. The Saturniidae (Lepidoptera) Lepidopteres Producteurs de Soie, fast. 4. A. Rey and of the Western Hemisphere, morphology, phylogeny, Cie, Lyon. 86pp. and classification. Bulletin of the American Museum STONE, S. E. 1991. Foodplants of world Saturniidae. of Natural History 98:335— 502, p1.5. Lepidopterists ' Society Memoirs, Number 4: xv + NARDELLI, U. 1986. Zur Kenntnis der Praimaginalstadien 186pp., 1 col. pl. and zur Zucht von Caligula thibeta Westwood, 1853 STRECKER, H. 1875. Lepidoptera, Rhopaloceres and (Lepidoptera, Saturniidae). Nachrichten des Heteroceres, indigenous and exotic; with descrip- Entomologischen Vereins Apollo, Frankfurt (N.F.) tions and colored illustrations, part 12:101—108, col. 7:11-23. pl.12. Owen's Steam Book and Job Printing Office, NASSIG, W. A. 1983. Zur Kenntnis von Caligula lindia Reading, Penn. (Moore) (Lep.: Saturniidae). Nachrichten des TUSKES, P. M. and M. M. COLLINS. 1981. Hybrid- Entomologischen Vereins Apollo, Frankfurt (N.F.) ization of Saturnia mendocino and S. walterorum, and 3:89-109. phylogenetic notes on Saturnia and Agapema . 1989. Wehrorgane and Wehrmechanismen bei (Saturniidae). Journal of the Lepidopterists' Society Saturniidenraupen (Lepidoptera, Saturniidae). 35:1—21. Verhandlungen der Westdeutscher Entomologentag, Dnsseldorf 1988, pp.253- 264.

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TUSKES, P. M. and M. J. SMITH. 1984. The life history WEAST, R. D. 1962. Two new foodplants of southwestern and immature stages of Agapema homogena Saturniidae. Journal of the Lepidopterists' Society (Saturniidae) . Journal of the Lepidopterists' Society 16:61— 62. 38:134—137. WOLFE, K. L. 1993. The Copaxa of Mexico and their VORA, R. S. 1990. Plant phenology in the Lower Rio immature stages (Lepidoptera: Saturniidae). Grande Valley, Texas. Texas Journal of Science 42: Tropical Lepidoptera 4: Supplement 1, 26 pp. 137—142.

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PIMIMPPL"F- A REVIEW OF THE GENUSAGAPEMA (LEPIDOPTERA: SATURNIIDAE) TABLE 1. RECORDED IIOSTPI.AN'1'S FOR AC:APEAfA.

Botanical Name and Plant Family Common Name A gaperna: Region References

IRhaminis californirus var. tlrsina California buckthorn, honrogena Arizona Tuskes and Smith 1984 (Greene) Wolfe, Rhatnnaceac coffeeber7

RhaInnas fraiig-ula L. buckthorn horrtogena captivity Stone 1991 (=Frangula alnus Mill.)

Rilres cereum I)ougl., squaw currant homogena Colorado Stone 1991 Saxifragaceae

Ribes al/nrtum L. irioutttain currant hnrrtogena captivity Stone 1991

Salix exigu a Null., gray sandbar willow hnmogerui Colorado D. Bowman pers. comm. Salicaccac

Cnndrrlia spalhulata A. Gray, squaw hush, anan.a. Arizona trerguson 1972 Rhainnaceac enstilla

Condalia lyvioider (A. Gray) Weberh. anona Arizona Ferguson 1972

Condalia viridis I. SI. Johnst. pla term is southwestern Texas K. B. Bryan pers. comm.

Condalia elcuides (Gray) javelina hush, anonla Arizona Ferguson 1972 M. C. John.st. tecomblate anon(1 Durango, Mexico l'eigler unpubl. dyrlri western Texas Kendall unpubl. dealifasCiala Nuevo Leon, Mexico Kendall unpubl.

C;ondalirr honlreri M. C. Johnst• bra.sil. gaMMina southern 'I exas Collins and Weast 1961 (= C. ohm ala Hooker) capnl negro dvari captivity Kendall unpubl. denli fascia1a captivity Kendall unpubl. (tlalensis captivity Kendall unpubl.

Rh us rnieroplrylla Engeltn., desert sumac, dyari Western Texas Kendall unpubl. ;lnacardiaccae scrub siunac, correosa

Fr' Denver Museum of Natural History Series 3, No. 3, p. 17, October 15, 1993 PEIGLBR AND IENDAU

TABLE 2. PARASITOIDS REARED FROM AGAPEMA.

Parasitoid Classification Hosts Locality References

DIPTERA

Euplw/'Ocera sp. near Tachinidae: gall/ina southern Texas Buck and Keister 1955, 1956 jlOlidensisTownsend Exoristinae

LesjJesia sp. near Tachinidae Iwmogena Arizona Peigler 1985 ll'x(,ma (Webber) Exoristinae galbina southern Texas Peigler unpub!. anona Arizona Peigler unpub!. anona Durango, Mexico Peigler unpub!.

LesjJfsia smniae (Webber) anona Arizona Peigler unpub!.

Lespesia sp. anona Arizona Peigler 1985 (arehippivora group)

HYMENOPTERA

Golesia eketme (Viereck) Braconidae: Agapemasp. ? Collins and Weast 1961 Microgastrinae dyari Reeves County, Texas Kendall unpub!. platensis Kinney County, Texas Kendall unpub!. homogena Arizona Peigler un pub!,

Enicospilus texanus Ichneumonidae: anona Arizona Peigler 1985 (Ashmead) Ophioninae dentifasciata uevo Leon, Mexico Kendall unpubJ. d)Ia1'i Reeves County, Texas Gauld 1988

L)'eogasler pullata Trigonalidae tachinid in uevo Leon, lexico Kendall unpub!. nevadensis Cresson dentifasciata

Microdontomerus Chalcidoidea: Enicost)ilus in Arizona Peigler 1985 fumipennis Crawford Torymidae anona

Anastatus "eduvii (Howard) Chalcidoidea: Eupelmidae platensis Kinney County, Texas Kendall unpubJ.

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A REVIEW OF THE GENUSAGAPEMA (LEPIDOPTERA: SATURNIIDAE)

^ftc^ '-e tr)n•,(!!f`Ie.s f.."' ^'`.j<. ftg Z^, r • orb ofPr..Z 2rt-erl-11,1-4.he ftot. trce^* /r'rw^ v C,...-y<' ,ern , .4,1' atl.t, Pa^ e, 4trryt cC7 s :,o t t., ^Gtsc,-icl.'{< Asefr .,f 14 y ►y, s .t-/^c. c-,o Rho i.0'

Leptdoptcra Type Lepidoptera Type Photograph No. 395 J'^/Lf AA//e Photograph No. 395 1'0,_4 Field Museum Field Museum ( 'old ' T Q 3 1r /,r 4

5.

Figures 1–5. 1. Agapema galbina, lectotype, male, 5. A. dyari, lectotype, female, Chihuahua, Mexico, W. Brownsville, Texas (FMNH). 2. A. galbina, paralectotype, Schaus collection (USNM). Photos 1–4 by J. E. Rawlins, 5 female (FMNH). 3–4. Labels for pair in Figs. 1–2 above. by C. Lemaire.

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iinitllllliiilII I 6 7 '8 9 1 0 111 1 —J

Figures 6-11. 6. Agapema dentifasciata, female, near light 29 October 1989 (ROK). 9-11. Reared ex-ovo on Matehuala, Nuevo Leon, Mexico, ex-cocoon on Condalia Condalia hookeri by R. O. Kendall. 9. Holotype, male, ericoides, emerged 22 August 1980, R. O. and C. A. Kendall emerged 6 October 1989 (DMNH). 10. Paratype, male, (ROK). 7-11. Agapema platensis, Kickapoo Cavern State emerged 2 October 1989 (ROK). 11. Allotype, female, Natural Area, Kinney County, Texas. 7. Paratype, male, at emerged 1 November 1989 (DMNH). Photo by Nancy light 7 November 1989 (ROK). 8. Paratype, female at Jenkins (Denver Museum of Natural History).

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Figures 12—20. 12. Agapema dentifasciata, male, near on Condalia ericoides in April 1981 by R. O. and C. A. Matehuala, Nuevo Leon, Mexico, ex-cocoon on Condalia Kendall. 16. Male, emerged 9 September 1981 (DMNH). ericoides, emerged 1 June 1980, R. O. and C. A. Kendall 17. Female, emerged 9 October 1983 (DMNH). 18. A. (ROK). 13. A. galbina, male, no locality data, emerged 17 homogena, male, Santa Catalina Mountains, Pima County, October 1967 (DMNH). 14. A. galbina, female, paratype Arizona, 1830 m, emerged 16 May 1984 (DMNH). 19—20. of A. solita, 7 October, Otto Buchholz collection A. anona, Cochise County, Arizona, cocoons collected on (AMNH). 15. A. pelora, male, ca. 16 km W Rancho Condalia spathulata by S. E. Stone. 19. Male, emerged 21 Progresso, Baja California Norte, Mexico, at light 6—7 November 1983 (DMNH). 20. Female, emerged 15 November 1987, Norris Bloomfield (SDNHM). 16—17. A. November 1983 (DMNII). Photo by Rick Wicker (Denver dtiari, near Orla, Reeves County, Texas, larvae collected Museum of Natural History).

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A A. galbina • A. dyari n A. anona ♦ A. dentifasciata o A. pelora A. platensis *A. homogena

Figure 21. Map of Mexico and the southwestern United Teller, Boulder, Clear Creek, States showing known localities for the species of Jefferson, and El Paso Counties Agapenaa. Records for A. homogena in Larimer, Routt, of Colorado are not shown.

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