J. Burns.
The Polytypic Genus Celotes (Lepidoptera: Hesperiidae: Pyrginae) from the Southwestern United States and Northern Mexico.
Psyche 81:51-69, 1974.

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THE POLYTYPIC GENUS CELOTES
(LEPIDOPTERA: HESPERIIDAE: PYRGINAE)
FROM THE SOUTHWESTERN UNITED STATES
AND NORTHERN MEXICO*
BY JOHN M. BURNS
Museum of Comparative Zoology, Harvard University Although most species of American hesperiids are hard to deter- mine, some are not; and none is more immediately distinct than the "streaky skipper," Celotes nessus (Edwards). Ever since it was described (in genus Pholisora) in 1877, nothing else like it has been known. Indeed, since the turn of the century -despite abortive attempts of various workers to jam it into polytypic genera (such as Py rgus [then commonly called Hesperia] , Systasea, and Antigonus) -it has properly stood alone in the monotypic genus Celotes (God- man and Salvin 1899). I feel, therefore, a measure of remorse in now describing a second species of Celotes that closely resembles the first.
Although I have accumulated data on Celotes since New Year's Eve of 1961, when I first recognized it as polytypic, I still see no clear clues to the evolutionary differentiation of C. nessus and the species to be described: they overlap fully in space and in time, and at least partly in choice of larval foodplants. The new species is a multivoltine mallow-eater occurring in a montane strip that runs northwest-southeast through the middle of the range of C. nessus. Because C. nessus is both peculiar and familiar, the following description is comparative. And because original descriptions can be too tedious for words, this one is largely visual. In preparing it, I have examined the genitalia df all specimens of Celotes available to me and possessed of an abdomen - a total of 529 individuals, of which 97 represent the new species. Wherever possible, I have also examined the metathoracic pouch of males. Each pinned specimen studied has received a sex-and-determination label.
The new specific name is a noun in apposition. Celotes limpia new species
Holotype. - c3 , Limpia Canyon, 5000 feet, Davis Mountains, + miles WNW of Fort Davis, Jeff Davis County, Texas, May 2, *Manuscript received by the editor March 1, 1974. Pwht U-:S 1-69 (1974). hup //psyche rinclub org/Si/81-05i.html



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52 Psyche [March
1959 (J. M. and S. N. Burns) [Museum of Comparative Zoology no. 318881.
Size. -On average, limpia is larger than nessus; and, in both species, females are larger than males, although size varies consid- erably (table I ). In nessus, the wingspread of Texas specimens exceeds that of Arizona specimens by at least I mm. Table 1. Maximum wingspread in well-mounted specimens of Celotes from Texas.
Species Sex Number Wingspread (mm)
measured Mean Range
nessus
Ì
Fades. - Fig. IE-H. Very like nessus (fig. IA-D), but with slightly larger and more conspicuous hyaline spots and a generally paler aspect.
Noteworthy in the latter connection is a set of pale areas between dark marks on the proximal 60% of the ventral secondary (i.e. from the three hyaline spots of the secondary inward) : these pale areas are enlarged and whitened in limpia and hence usually more pro- nounced than in nessus. In dorsal view, unworn (particularly reared) specimens of limpia may show considerable grayish overscaling on the body and adjacent wing bases. The usually paler shades of brown and tan in limpia tend to make it reflect more and appear colder and harder and sometimes more contrasty. All of these average differences involving color are perceptible chiefly when limpia is compared with nessus from Texas; nessus from Arizona (where limpia does not occur) often essentially duplicates the facies of limpia - except insofar as the remarkably small average size of Arizona nessus gives its pattern a special sharpness. Wear and fading Data for specimens in Fig. 1 :
A, B - Sitting Bull Falls, 4650 feet, Guadalupe Mountains, Eddy County, New Mexico, V-26-1959 (J. M. and S. N. Burns). C, D -Topotype. San Antonio, Bexar County, Texas, VI-30-1963 (J. M. Burns).
E,
F - Holotype. Limpia Canyon, 5000 feet, Davis Mountains, 4 mi. WNW of Fort Davis, Jeff Davis County, Texas, V-2-1959 (J. M. and S. N. Burns).
G, H - Allotype. As for holotype, except V-1-1959.



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Burns - Polytypic Genus Celotes
Fig, 1. Facies of Cehtes.
c. nessus
B ÌÔ 8
[ * D
-+ $
C. limpta
F -+ 8
G H ÌÔ Q
4 J,
dorsal ventral




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raise additional hell with these subtle color characters, but no matter : the genitalia {q.v.) last and last.
Male scent-spreading sex characters. - Males of limpia, like those of nessus, have a narrow costal fold on the primary and a metathoracic system of ventral pouch and tibia1 tufts. The meta- thoracic pouch of hpia (fig. 2B) bears short linear scales over most or all of its dorsal surface (except at the lateral margin). In nessus (fig. aA) , linear scales are altogether absent (particularly in Arizona males) or, if present (as in Texas males), are much less dense and/or proximal in distribution; even in extreme cases, they never attain the distal apex of the pouch, and the pouch is therefore clad-en- tirely or for its greater part - with much broader, flat-lying shingle- like scales (cf. figs. 2A and 2B). Because the linear scales of Simpiu are creamy, whereas some to all of the broad shinglelike scales of nessus are gray to brown, the general color of the dorsal surface of the metathoracic pouch differs in the two species. (Interior morphol- ogy of costal folds has not been studied.) Fig. 2. Metathoracic pouch of the male; abdomen removed, dorsopos- teriar view. A. Crhfes MSSUS from Austin, Trails County, Texas, V-26- 1966 (J. M. Barns). B. Cdotes Wia, a paratype reared out EX-10-1966 from a larva collected on fiutilon malacam, 15 mi. SE Redford, 2500 ft., Presidio County, Texas, VIII-18-1966 (K. 0. Kendall). Male genitalia. - Fig. 4. Although basically similar to nessus (fig. 3)) there are abundant differences, of which the most salient include the following. (I) In limpia, a long thin flattened projection, like a curved spatula, arises from the dorsoposterior rim of the body of the valva and starts dorsad but at once curves caudad, extending far back to become the caudalmost valval element and to end in a



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19741 Burns - Polytypic Genus Celotes 55 slightly flared tip with a peripheral array of fine sharp teeth (fig. 4A-C). Its homolog in nessus is a robust spike arising in a similar position, except that the distinctive high dorsal curvature of the body of the valva in nessus places the origin of the spike well down on the medial surface of the valva (fig. 3A-C) ; this tapering spike extends chiefly dorsad, but usually bends slightly caudad at its apex, which is pointed and often entire or bifurcate (fig. 3A,B), but sometimes trifurcate, or very rarely quadrifurcate; although occasionally broad- ened or lengthened and bent strongly caudad apically, the spike never approaches the form of the spatula in limpia. (2) Anterior end of tegumen, in limpia, large and well-developed, projecting far cephalad (fig. 4E,F) ; but in nessus, extraordinarily reduced (fig. 3E.F). (3) In limpiu, uncus broad, and the paired terminal prongs of the uncus heavy and stubby (fig. 4F) ; but in nessus, uncus relatively narrow, and its terminal prongs more delicate (fig. 3F). Female genitalia. - Fig. 6. Sterigma of limpid altogether more massive (fig. 6A,B) and, viewed ventrally, more nearly square in outline (fig. 6A) ; heavy central sclerotization in lamella postvagi- nalis like a wide triangle tapering quickly toward ostium bursae (fig. 6A). Sterigma of nessus (fig. 5) less massive (fig. 5A,B) and, in ventral view, narrower posteriorly, suggesting in outline a cau- dally truncated triangle (fig. 5A) more than a square; heavy central sclerotization in lamella postvaginalis limited to a comparatively narrow midventral strip that tapers but little toward ostium bursae (fig. 5A). Ostium bursae of limpia a relatively narrow curved slit, like a crescent bowed dorsad (fig. 6C), but that of nessus large and more or less round, like a manhole (fig. 5C). Spatial distribution. -A widespread species, nessus ranges from about the 97th meridian in southcentral Oklahoma and central Texas west to northwestern and southeastern Arizona and south in Mexico to at least southern Sonora, southern Chihuahua and Coahuila (fig. 7). By contrast, so far as known, limpia occurs only along the southern Rocky Mountain axis in Trans-Pecos Texas and along the Mexican counterpart of this axis, the northern segment of the Sierra Madre Oriental, as far south at least as southern Coahuila (fig. 8). Having been taken in numbers in the Guadalupe, Davis, and Chisos mountains of Trans-Pecos Texas, limpia may confidently be expected (I) in many other west Texan ranges-such as the Delawares, Chinatis, and SantiagosÌ that are more or less associated with the scattered southern Rocky Mountain system, (2) northward in at least the New Mexican extension of the Guadalupes, if not in the Sacramentos and more northern chains, and (3) southward in Chi-



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Fig. 3.
Male genitalia of Celotes nessus. A. Lateral view of right valva. B. Medial view of right valva.
C. Dorsal view of both valvae. D. Dorsal view of aedeagus. E. Left lateral view of uncus, tegumen, gnathos, vin- culum, and saccus. F. Dorsal view of uncus, tegumen, vinculum, and saccus. [A and B, drawn from male from Palo Duro Canyon, 2800 feet, Randall County, Texas, V-10-1959 (J. M. an~d S. N. Burns) ; C to F, from male from Austin, Travis County, Texas, V-26-1966 (J. M. Burns).]



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19741
Burns - Polytypic Genus Celotes
57
Fig. 4.
Male genitalia of Celotes limpia. A to F as in fig. 3. [A and B, drawn from holotype; C to F, from paratype from type locality, V-4-1959 (J. M. and S. N. Burns).]




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Fig. 5.
Female genitalia of Celotes nessus. A. Ventral view of bursa copulatrix, sterigma, and ovipositor lobes. B. Right lateral view. C. Ven-
troposterior view. [Drawn from female from Austin, Travis County, Texas, IV-18-1967 (J. M. Burns).]




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19741
Burns - Polytypic Genus Celotes
\
(J
Fig. 6. Female genitalia of Celotes limpia. A. Ventral view of bursa copulatrix, sterigma, and ovipositor lobes. B. Left lateral view. C. Ventro- posterior view. [Drawn from paratype from Limpia Canyon, 4700 feet, Davis Mountains, 5 mi. NE of Fort Davis, Jeff Davis County, Texas, V-4- 1959 (J. M. and S. N. Burns).]




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Fig. 7. Spatial distribution (based on material examined) of Celotes nessus - southwestern United States and northern Mexico. ~ikers and the borders of states are shown. Type locality indicated by arrow. huahua, as well as Coahuila, and, in all likelihood, beyond these states. Indeed, either limpia or nessus (see Nomencla/uraZ epilogue below) was long ago reported from Durango city (Godman and Salvin 1899), and both species probably range well down the Central Plateau of Mexico. The distribution of limpia appears to lie within that of nessus.
Over its much broader distribution, nessus occurs from sea level to somewhere between 5000 and 6000 feet; limpia, with its strictly interior distribution, is now known from an elevational range of 1900-5700 feet. In montane regions of the southwestern United States, both skippers are species of alluvial fans, foothills, lower canyon reaches, and lower elevations generally; they avoid higher elevations.
The distribution of nessus in the United States is not perfectly known. The skipper may conceivably occur in the New York and Providence mountain region of southeastern California, in southern Nevada, in southern Utah, possibly in extreme southwestern or south- eastern Colorado, and certainly in considerably more of New Mexico



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19741
Burns - Poiytypic Genus Celotes
Fig. 8. Spatial distribution (based on material examined) of Celote~ /im$ia - Trans-Pecos Texas. Mountains and the borders of counties are shown.
Type locality indicated by arrow, The range as plottad spans a north-south distance of about 220 miles; but, since preparing this map, I have examined one male from southern Coahuila, Mexico, about 250 miles southeast of the Big Bend.
than fig. 7 indicates. On the other hand, this map probably gives a close approximation of the eastern distributional limit of nessus- This meridional eastern limit coincides neatly with that of many other organisms - including other pyrgine skippers of the southwestern United States and northern Mexico such as Pyrgux philetas Edwards (Burns and Kendall 1969) and Erynnis meridianus Bell (Burns 1964, and unpublished) -and reflects an important biogeographic barrier.
There are two freaky records of streaky nessus east of the 97th meridian. The first [Burton, Georgia, V-2 I -I 91 I (J. C. Bradley) (Cornell University collection) ] was published by Harris ( I 950) and repeated [as "Georgia (once!)"] by Klots ( 195 I ), but was not even mentioned later by Harris (1972) and should be discounted. The second [Tallulah, Louisiana, between VIIX- I 926 and X- I 93 I (collected in the day at an altitude of 20 feet by an airplane)] was published by Glick (1939) ; but though it has been repeated (Lam- brernont 1954; Mather and Mather 19581, the feat itself has not.



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Temporal distribution. - Both Zimpia and nessus are multivoltine, and both fly at the same times. Nearly all of the dated wild-caught adults of CeZotes that I examined were collected from March to September: in Texas, every half-month interval from the beginning of March to mid-September includes records of both Zimpia (N = 68) and nessus (N = I 58) ; however) for Arizona nessus (N = 183)~ the months of May and June together yield but 3 records, which leaves a major gap between one large cluster of records in March and April and another in July) August, and the first half of Septem- ber. The only other dated wild-caught specimens examined are 5 nessus from far southern Arizona collected in mid-January and 5 nessus from the last half of September and the first half of October from central and) as might be expected) extreme southern Texas (the lower Rio Grande Valley). In the experience of Kendall ( I 965) ) nessus flies from early March to mid-November in Texas. The fact that temporal distribution is essentially continuous through the warm season for both Zimpia and nessus in Texas but distinctly bimodal for nessus in Arizona probably relates to the dif- ferent patterns of rainfall in the two states. For the most part, Texas has much more evenly distributed precipitation than does Arizona, where it occurs in two widely disjunct winter and summer periods. The second flight period of nessus in Arizona coincides with the summer rainy season, to which it is presumably geared. In the course of his field studies and laboratory rearings, Kendall (1965) and unpublished) has observed that full-grown larvae of CeZotes are able to enter diapause (facultatively) to beat heat and drought) as well as to get through the cold of ,a winter season. Kendall has shown in the laboratory that such diapause can be broken by pro- viding moisture, whereupon larvae pupate and produce adults promptly. Given this capacity to be flexible, variable larva1 growth rates, and the usual vicissitudes of weather, individuals of a single generation must often get out of synchrony with the result that gen- erations overlap broadly and irregularly, forming no definite number per year in either species.
There are no indications that Zimpia and nessus are temporally displaced with respect to each other where they spatially coexist in Trans-Pecos Texas. Rather, my records show that adults of both species have repeatedly been collected in association - unwittingly, of course, since the collectors were unaware of the existence of more than one species of CeZotes. On nine occasions altogether) five dif- ferent collectors (or pairs of collectors) have caught a total of 37



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19741 Burns - Polytypic Genus Celotes 63 limpia and nessus at the same place on exactly the same date- the places occurring in Jeff Davis and Brewster counties and the dates in March, May, June, July) and August. Many other records of both species at a single locality differ by only one to a few days. Larval foodplants. - Numerous studies in Texas by R. 0. and C. A. Kendall show that both limpia and ne.ssus feed on various species of Malvaceae. For both skippers, the Kendalls have witnessed oviposition - directly on larvd foodplants - in the wild, and have collected numerous eggs and larvae from which they have reared adults.
In the field, Zimpia eats Abutilon malacum Wats., A. incanum (Link) Sweet, Sphaeralcea angustifolia var. lobata (Wmt.) Kern.) and Wissadula holosericea (Scheele) Gke., and, in the laboratory, also Malvastrum american,um ( L. ) Torr. and Althaea rosea Cav. In the field, nessus eats A butilon incanumJ Sphaeralcea angustif olia var. lobataJ Wissadula hofiosericea, W. amplissima (L.) R. E. Fries, and Sida+J&.es Gray, and, in the laboratory, also Althaea rosea. Both skippers very JikqJpeat other mallows, as. well. Documentation: Eggs and larvae of Celotes were found on Sphaeralcea angustifolia var. Zobata at Davis Mountains State Park, Jeff Davis County, V-1-61, by R. 0. and C. A. Kendall. At the laboratory in San Antonio, those larvae that were not preserved were reared perforce on Althaea rosea. One adult was obtained from a larva on V-28-61 and two from eggs on VI-6 and 9-61. Although all were reported as nessus (Kendall 1965)~ I find that only the adult that emerged VI-6 is nessus whereas those that emerged V-28 and VI-g are limpia (paratypes).
From 3 larvae found by the Kendalls on WissaduZa holosericea in Musquiz 'Canyon on state highway I 18, Jeff Davis County) VIII- 14-66, I adult (a paratype of limpia) was obtained on IV-14-67. From 2 larvae found by the Kendalls on Abutilon incanum near Rio Grande Village in Big Bend National Park, Brewster County, X-12-67, 2 adults of limpia (paratypes) were obtained (after a lengthy diapause) on VI-19 and 22-68. At the same locality, 111- 27-68, the Kendalls saw a female of Celotes oviposit on A. incanum but got only the egg and not the female; from this egg, an adult of limpia ( paratype) was obtained on V-29-68. The Kendalls collected 23 larvae on Abutilon maZacumJ VIII- 18-66, at a point on ranch road 170 overlooking the Rio Grande, about 15 miles southeast of Redford, Presidio C,ounty; and, using A. incanumJ Malvastrum americanumJ and especially Wissadula



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holosericea as larval food in the laboratory) they reared 22 adults of limpia (all paratypes) (one seemingly parasitized laxva was pre- served). At exactly the same spot, X-I 1-67, the Kendalls collected about 40 more Celotes larvae on A. malacum and reared 23 adults (apparently Zimpia, but none of them examined by me). Foodplant records for nessus are legion (Kendall 1959, 1965 and unpublished). Most of them are of Abutilon incanum; and these come from many counties in central Texas: Bexar (innumerable records), Blanco) Bosque) Comal, Crockett, Kerr, Kimble) Llano, Maverick, McCulloch, Medina, Nueces, San Patricio, Travis) Uvalde, and Val Verde, I have examined much of the nessus ma- terial reared on A. incanum.
The one valid record of nessus found as an egg on Sphaeralce~ angustifolia var. lobata in Jeff Davis County) Trans-Pecos Texas, and reared to an adult in the laboratory on Althaea rosea was dis- cussed above; the other two adults reared from this lot of eggs and larvae on Sphaeralcea are actually limpia, though they, too, were reported ( Kendall I 965 ) as ness,us.
Three larvae found on Sida filipes in San Antonio, Bexar County) V-9-63) and kept on this plant, produced adults of nessus (not seen by me) on VI-8) 14) and VII-I 1-63 (Kendall 1965). A larva found by the Kendalls on WissaduZa holosericea in San Antonio, Bexar County, V-24-65) yielded an adult of nessus (seen by me) on VI-I 1-65.
Three larvae found on W. amplissima in extreme southern Texas at the Laguna Atascosa National Wildlife Refuge, Cameron County, IV-21-62) produced adults of nessus (all seen by me) on V-I 5, VI-4, and VIII-20-62 (Kendall 1965 ) .
Hypodigm, - Of limpia: the holotype c? specified earlier) an allo- type 9 with identical data (except V-1-59) y and paratypes comprising 64 8 309 from Trans-Pecos Texas and I 8 from Mexico. Paratypic data arranged in the system described by Burns ( 1964 : 19-20), with names of recurring collectors abbreviated: J. M. and S. N. Burns to B, H. A. Freeman to F, and R. 0. and C. A. Kendall to K. MEXICO.
COAHUILA. 25 mi. N Saltillo, IX-19-69> I 8 (J. A. Scott).
UNITED STATES. TEXAS. Brewster County : Alpine, V-20- 26, I 8 (0. C. Poling) (MCZ) ; V-21-26, I (0. C. Poling) (MCZ) ; VI-4-42, I ~7 (I?) ; VI-5-42) I 8 (I?) ; VI-6-42? 3 d (I?) ; VI-7-49, 18 (F); VII-10-49, 19 (F); VIII-9-52? 2d (F); VII-28-53, I d (F) ; VI-16-60, I? (F) ; 111-14-61, I 8 (W. S.



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19741 Bums - PoZyty$ic Genus CeZotes 6 5 McAlpine) ; 111-27-61) 7 d (W. S. McAlpine) (2 in PMY) ;
VI-5-61) 1d
(F). 8 mi. N Study Butte) 3500 ft.) VII-15-63) 1d' (H. Clench) (CM), Big Bend National Park: The Basin) 5500 ft.) IV-30-65) I? (K) ; IX-8-65, I 3 (K). Oak Spring, 5700 ft.) IX-9-65, 19 (K). Rio Grande Village) 1900 ft.) 111-27-68) 3 d (K) ; V-29-68> I d reared from egg (K) ; VI-19, 22-68) 2d reared from larvae (K). Window) 5500 ft.) VIII-16-66) 3 d (K). Culberson County: 2 mi. N Pine Springs) 5700 ft.) Guadalupe Mts.) VII-19-63) 6d (H. Clench) (CM). Jeff Davis County: I I mi. N Alpine) VI-20-68) 5 d
(J. A. Scott). 13 mi. NW Alpine on state highway 118) IV-17-60, I cJ' I? (K. Roever). Big Aguja Canyon) IV-2-68) 5~3
(K). Davis Mts. State Park) V-28-61) I d reared from larva (K) ; VI-9-61? 19 reared from egg (K). Ft. Davis, VIII-27-54, I d (R. M. Bohart). Limpia 'Canyon) 4800 ft.) Davis Mts., I mi. N Ft. Davis, IV-28-59) I d (B) (MCZ). Limpia Canyon, 5000 ft.) Davis Mts.) 4 mi. WNW Ft. Davis) IV-28-59> 29 (B) (Mcz) ; v-1-59) 2d 19 (B) (Mcz); v-2-59) 29 (B) (MCZ) ; V-4-59) I (B) (MCZ), Limpia Canyon) 4700 ft.) Davis Mts., 5 mi. NE Ft. Davis) V-4-59> I 6 19 (B) (MCZ).
Musquiz Canyon on state highway 118) VIII-14-66) 2d (K) ; IV-14-67, 19 reared from larva (K). Presidio County: 30 mi. SE Presidio along Rio Grande) 111-26-66) 29 ( D. J. Lennox) . I 5 mi.
SE Redford on ranch road 170) 2500 it.) VIII-29) 30) 31-66 and