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Volume 47 1993 Number 1
ISSN 0024-0966
JOURNAL
of the
LEPIDOPTERISTS’ SOCIETY
Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS
24 February 1993
THE LEPIDOPTERISTS’ SOCIETY
EXECUTIVE COUNCIL
Ray E. STANFORD, President HIROSHI INOUE, Vice President FLoypD W. PRESTON, Immediate Past President IAN KITCHING, Vice President M. DEANE BOWERS, Vice President ROBERT J. BORTH, Treasurer
WILLIAM D. WINTER, Secretary
Members at large:
Karolis Bagdonas Charles V. Covell, Jr. Eric H. Metzler Steven J. Cary Linda S. Fink Robert K. Robbins Stephanie S. McKown Scott E. Miller J. Benjamin Ziegler
EDITORIAL BOARD
PAUL A. OPLER (Chairman), FREDERICK W. STEHR (Member at large) JOHN W. BROWN (Journal), WILLIAM E. MILLER (Memoirs) STEPHANIE S. MCKOWN (News)
HONORARY LIFE MEMBERS OF THE SOCIETY
CHARLES L. REMINGTON (1966), F. MARTIN BROWN (1973), E. G. MUNROE (1978), ZDRAVKO LORKOVIC (1980), IAN F. B. COMMON (1987), JOHN G. FRANCLEMONT (1988), - LINCOLN P. BROWER (1990), DoUGLAsS C. FERGUSON (1990),
HON. MIRIAM ROTHSCHILD (1991), CLAUDE LEMAIRE (1992)
The object of the Lepidopterists’ Society, which was formed in May 1947 and for- mally constituted in December 1950, is “‘to promote the science of lepidopterology in all its branches, .... to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures’ directed towards these aims.
Membership in the Society is open to all persons interested in the study of Lepi- doptera. All members receive the Journal and the News of the Lepidopterists Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. There are four numbers in each volume of the Journal, scheduled for February, May, August and November, and six numbers of the News each year.
Active members—annual dues $25.00 Student members—annual dues $15.00 Sustaining members—annual dues $35.00 Life members—single sum $500.00 Institutional subscriptions—annual $40.00
Send remittances, payable to The Lepidopterists’ Society, to: Robert J. Borth, Treasurer, 6926 North Belmont Lane, Fox Point, Wisconsin 53217, U.S.A.; and address changes to: Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, California 90007-4057 U.S.A. For information about the Society, contact: William D. Winter, Sec- retary, 257 Common St., Dedham, Massachusetts 02026-4020, U.S.A. (617-326-2634). To. order back issues of the Journal, News, and Memoirs, write for availability and prices to the Publications Manager: Ronald Leuschner, 1900 John St., Manhattan Beach, Cali- fornia 90266-2608, U.S.A.
Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for $40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, California 90007-4057. Second-class postage paid at Los Angeles, California and additional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, California 90007- 4057.
Cover illustration: The head of a small tineid moth (Tineidae) from Arizona, with all scales removed, illustrating the primitive mouthparts. Original pen and ink drawing by Elaine R. S. Hodges, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. :
JouRNAL OF Tue LeEPIDOPTERISTS’ SOCIETY
Volume 47 1993 Number 1
Journal of the Lepidopterists’ Society 47(1), 1993, 1-7
PRESIDENTIAL ADDRESS, 1992: MEGATRENDS AND THE LEPIDOPTERISTS’ SOCIETY’
FLOYD W. PRESTON 832 Sunset Drive, Lawrence, Kansas 66044-23873
With the approach of the 50-year anniversary of the Lepidopterists’ Society in 1997, it is appropriate that we begin to think about what type of Society we want for the next half century. I hope that this address will help to identify a few major goals that I believe the Society should pursue.
I joined the Society in 1948, missing charter membership by one year. As a member through its first 45 years, I have a vantage point from which to look back upon our Society and, perhaps, from which to offer some thoughts about its future. It is reassuring to read the premises upon which our Society was founded as stated by Charles L. Remington and Harry K. Clench in the May 1947 issue of the Lepi- dopterists’ News. Three points stand out above the rest: 1) the Society would be devoted to the scientific study of Lepidoptera; 2) its mem- bership would be international; and 3) the Society would include both amateurs and professionals. Harry Clench was able to reaffirm the primacy of these premises twenty-five years later when he wrote the history of the Society in our Commemorative Volume (1945-78). These same principals guide us today, and I hope that they will continue to be central to the Society in the future.
Technological, economic, and political changes are occurring on a vast scale and at a rapid pace. It is never easy to establish policy to accommodate future changes; however, I would like to raise a few
' Presidential address presented at the 43rd Annual Meeting of the Lepidopterists’ Society, East Lansing, Michigan, 27 June 1992.
2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TABLE 1. Influence of major trends on the Lepidopterists’ Society.
Megatrend Impact on Lepidopterists’ Society 1. Industrial society to informational society Publication methods 2. High tech/high touch Butterfly houses Butterfly counts 3. Centralization to de-centralization Regional lepidopterist societies
questions to our membership, realizing that it probably will require the next five years to evaluate and answer them. I am ever mindful of Mark Twain’s famous quip that “predictions are very difficult, especially about the future.’ My rationale for anticipating the future comes from Megatrends, a book written ten years ago by John Naisbitt. Many, but not all, of his predictions have come true. More importantly, Naisbitt provided social and psychological unifying themes to explain major or ‘““megatrends” in the world.
I have chosen three of these themes that have influenced and will continue to influence our Society and the study of Lepidoptera. These are listed in Table 1 with an indication of the type of impact they will have or have had upon our Society.
Industrial Society to Informational Society
Whether the Lepidopterists’ Society has made the transition from an industrial to an informational Society may be debated, but new tech- nology undoubtedly has permitted us to transmit and process infor- mation more quickly and efficiently than ever before. There are few members left in our Society who received the first issue of the News in May 1947. It was mimeographed using a typing stencil. Such stencils now are obsolete and almost unavailable. Everyone who remembers typing on stencils, with their messy correction fluid and their high premium on perfect typing, probably is glad that they have disappeared. Our editor of the News now produces camera-ready copy with a com- puter by a process that wasn’t even dreamed of in 1947.
Looking into the future, we should realize that what we see today is but a point on a continuum of change in information processing tech- nology—new approaches are being developed continually. The Journal of Research on the Lepidoptera (JRL) now is offering the JRL cumu- lative index on computer disk. Should we consider something similar? Should we consider the use of compact disks (CDs) not only for indices but also for volumes of the Journal? Is our material copyrighted so that the Society could benefit from such a medium? A modest way to begin
VOLUME 47, NUMBER 1 3
computerization may be to advertize in the News for volunteers to assist Richard Arnold with the compilation and data entry for a cumulative “Season Summary.” Should we begin to develop regional data bases on Lepidoptera or at least coordinate the standardization of such data bases? Scott Miller strongly suggested this possibility last year in his talk at Tucson. Should we develop a standardized computer format for articles for the News and/or Journal? If the Society plans to make educational materials available to schools, should the material be sup- plied in machine readable format? Many secondary school libraries now have computer data bases search capability.
High Tech/High Touch
The second theme, high tech/high touch, involves the connection between growth of high technology industries, with their mechanization and impersonalization, and growth of industries that emphasize personal awareness, person-to-person contact, and personal involvement. I be- lieve that this theme is manifested in the growing popularity of butterfly houses. People hear much about the cause and effect behind environ- mental issues such as ozone depletion, global warming, and pollution. They can achieve some temporary relief from the chaos and experience a little bit of the “Garden of Eden” when they walk through a butterfly house. This same high tech/high touch action-reaction also may explain the increasing popularity of butterfly counts—people want to become involved directly.
How does all this relate to the Lepidopterists’ Society? I believe we should take advantage of the growing interest in butterfly houses. We should ask ourselves if we are doing all we can to encourage contri- butions to our Journal and News for providing the basic background science for mass rearing of Lepidoptera, whether for butterfly houses or potential environmental restoration projects. Terry Domico, a free- lance wildlife biologist, recently asked me “for how many of our but- terflies and moths do we know enough of the biology that we could undertake a mass-rearing and re-population effort?’’ He related an interesting story to me in regard to this question. In preparation of a book on the insects of Borneo, he visited Lepidopterists’ Society member David Goh at his butterfly farm in Malaysia, where Goh is mass rearing species for butterfly houses. In a few weeks, Domico recorded the life histories of four species of Troides that occur in Borneo for which there were no published life history data. This indicates to me that much of the little life history information we have may actually reside, unpub- lished, in the minds of amateurs. There is a need to get such information into the public domain.
4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Centralization to De-centralization
The third theme that I have selected from Naisbitt’s book is the trend from centralization to de-centralization. Naisbitt relates this trend to changes in government and business practices, and explains it in terms of the growing sophistication and diversity of the populace. For the Lepidopterists’ Society, I see this theme exhibited in the proliferation of regional lepidopterist organizations. Currently there are nine of these in the U.S. alone, some more formally structured than others. The growth of these organizations has occurred during a period when mem- bership in our own Lepidopterists’ Society has remained more or less constant. A significant portion, but by no means all, of the members of these regional groups are also members of the Lepidopterists’ Society. We should not view this movement with alarm nor should we attempt to “coordinate” or control it; we should encourage such groups to form. Our Society can only benefit from such a stance. My wife and I have visited five of the nine organizations, several repeatedly. There is great vitality and enthusiasm there, especially evidenced in local meetings and field trips. This is truly the grassroots of our organization.
What I see in these local groups is a greatly under-utilized resource for making significant contributions to the study of Lepidoptera. There are people willing and able to take part in short and long term survey projects. There are local floral and faunal experts. What is needed is a recognition of this resource and creative leadership to channel its energy toward prioritized local projects. Such efforts will be of extreme value if and when conservation projects are initiated by local and other agen- cies. This need for professional leadership also was expressed last year by Scott Miller.
These regional lepidopterist organizations represent one of the best means to provide the personalized one-on-one encouragement and ed- ucation for young people aspiring to learn more about the science of lepidopterology. They are an excellent recruiting mechanism for our Society and the profession.
It is not clear how the Society can further the aims of local lepidop- terist organizations without encroaching upon their autonomy. For- malizing and expanding our publication of their activities may help. A forum for local group representatives could be provided at the Annual Meeting. Articles in the News could highlight model activities such as local studies or re-population efforts.
In keeping with the international scope of our Society, we should cover activities of organizations similar to ours in other part of the world. Our “News From Europe” editor, Willy De Prins, is exploring
VOLUME 47, NUMBER 1 5
Fic. 1. Pyrrhopyge hygieia rufipectus (Godman and Salvin) (Hesperiidae). Upper photo from 35 mm color slide taken in the Rio Napo area of eastern Ecuador by Joseph T. and Suzanne L. Collins ca. 1970. Lower photo from 35 mm slide taken of color monitor display of the digitized version of the butterfly. Digitization was done at 900 dots/inch using 24 bit color.
6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
increased communication links with SEL (Societas Europaea Lepidop- terologica).
High Technology
As an example of how technology might impact the study of Lepi- doptera in the future, I would like to describe briefly the exciting and rapidly advancing field of high resolution digital imaging.
Biology is a visual response discipline, whether one thinks of field observation, type specimens, or microscopic studies. High resolution color photography is the current medium of choice to record visual data. Sometime within the next five to fifteen years, we will have high resolution digital television in our homes. This will mean easy access to a display device, and because the signals are digital, microcomputers will be capable of analyzing them. Images may be transmitted with high fidelity for publication or analysis where phenotypic information is sufficient. More importantly, new types of quantitative phenotype analysis would be possible.
As an introductory example, I would like to present an image created at the National Center for Supercomputing Applications (NCSA) at Champaign-Urbana. Figure | is in two parts. The upper part was prepared from a conventional 35 mm color slide taken in 1970 in the Rio Napo area of eastern Ecuador. The lower portion was prepared in two steps. The original slide was digitized at 900 dots/inch (dpi) in 24 bit color (16.8 miilion different colors). The digital record of the image was then displayed on a special, very flat, high resolution color monitor and photographed with a 85 mm camera. The resulting 35 mm slide and the original slide were then composited by the printer to produce Fig. 1. The high fidelity of the lower image demonstrates the great promise of digital storage and retrieval techniques for archiving color pictorial information. The price one pays for this fidelity is the need for very large memory storage capability (approximately 70 million bytes per image). Reasonable fidelity currently is widely available in 8 bit color (256 colors). When the high resolution or high fidelity color equipment becomes readily available, images can be placed on compact disks (CDs) and distributed like books, i.e., loaned or sold. Even more important is the possibility of using the images for studies never before possible, such as viewing an image as a predator might view it if the spectral response of the predator’s eye (color palette) is known.
In conclusion, a wonderful symbiotic relationship exists within our Society between amateurs and professionals. We now see a similar relationship evolving between our Society and regional lepidopterist groups that share our goals. As a Society, we have benefited directly
VOLUME 47, NUMBER 1 7
from technology in our increased publication capabilities and indirectly from technology’s tendency to foster antithetical activities such as but- terfly houses, butterfly counts, and participation in local survey and conservation projects. We can look forward to our 50th anniversary with great confidence and high expectations.
ACKNOWLEDGMENTS
I thank Joseph T. Collins and his wife Suzanne L. Collins for loaning me the color slide of Pyrrhopyge hygieia and Stephen R. Steinhauser for identifying the specimen. Special thanks go to Kenneth A. Bishop of the University of Kansas Department of Chemical and Petroleum Engineering, and Jay Alameda, industrial consultant, National Center for Supercomputing Applications (NCSA), University of Illinois, for their pioneering efforts to digitize the original image and prepare the 35 mm slide from the display of the digitized image. Thanks also go to Ken Blair of Allen Press for insuring the faithful renditions of the two slides forming Fig. 1.
Received for publication 8 July 1992; accepted 8 July 1992.
Journal of the Lepidopterists’ Society 47(1), 1998, 8-16
THE NATURE OF ANT ATTENDANCE AND THE SURVIVAL OF LARVAL ICARICIA ACMON (LYCAENIDAE)
MERRILL A. PETERSON’ Department of Zoology, University of Washington, Seattle, Washington 98195
ABSTRACT. I examined ant attendance and its importance to larval survivorship in a facultatively myrmecophilous butterfly, Icaricia acmon (Westwood and Hewitson) (Ly- caenidae), in a population that uses two host plant species, Eriogonum compositum Doug. and E. strictum Benth. (Polygonaceae). Third and fourth instar larvae of I. aemon were tended by three ant species: Tapinoma sessile (Say), Formica neogagates Emery, and an unidentified Formica species. Third instar larvae were tended less frequently than fourth instar larvae on both plant species, and T. sessile was the attendant ant species for a higher proportion of third instar than fourth instar larvae developing on E. com- positum. Over the duration of the study, all switches of attendant ant species on individual plants were from early T. sessile attendance to later F. neogagates attendance. An exclosure experiment revealed that ant attendance had no significant effect on larval mortality.
Additional key words: Tapinoma, Formica, facultative myrmecophily, Washington, Eriogonum.
The association of lycaenid larvae with ants (Formicidae) has pro- vided researchers with model systems for studying the costs and benefits of mutualisms. Recent work has focused on both ecological and evo- lutionary aspects of these mutualisms, including host-parasitoid inter- actions (Pierce & Mead 1981), oviposition behavior (Atsatt 198la, Pierce & Elgar 1985), and the evolution of host choice (Atsatt 1981b, Pierce 1985). To understand the evolution of these mutualisms, it is critical to examine the costs and benefits to both partners. Several field studies have documented the importance of ants to the survival of larval ly- caenids (Pierce & Mead 1981, Pierce & Easteal 1986, Pierce et al. 1987, Fiedler & Maschwitz 1989b), but few have experimentally addressed this issue in facultative ant-lycaenid associations (but see Pierce & Mead 1981, Pierce & Easteal 1986). This is particularly surprising when one considers that most lycaenid-ant associations are facultative (see Fiedler 1989a for a recent review). Although the mutualisms studied so far provide excellent ecological and evolutionary case studies, it is impor- tant to recognize how variability in the intensity of ant-lycaenid asso- ciations might affect generalizations about these systems. To broaden our understanding of facultative ant-lycaenid mutualisms, I conducted an ant exclosure experiment to determine the importance of ant atten- dance and host plant choice to larval demography in a population of Icaricia acmon (Westwood and Hewitson) (Polyommatinae: Polyom- matini) in central Washington.
' Present address: Section of Ecology and Systematics, Cornell University, Ithaca, New York 14853.
VOLUME 47, NUMBER 1 9
STUDY SITE AND ORGANISM
The population of Icaricia acmon lutzi (dos Passos) that I studied occupies a bench of the Yakima River, 11.2 miles north of Yakima, Washington at 46°47'’N, 120°27'W and 375 m elevation. The habitat represents the Artemisia /Festuca zone described by Franklin and Dyr- ness (1978), but has fewer Artemisia and more Eriogonum (Polygonace- ae) species than typical Artemisia /Festuca communities. The four com- monest Eriogonum species were E. compositum Dougl., E. strictum Benth., E. microthecum Nutt., and E. elatum Doug]. The climate in this region is arid to semiarid with relatively warm summers and cold winters. The mean annual precipitation in nearby Yakima is 20 cm with only a small amount (2.9 cm) of this falling during June through August. Average temperatures in the Yakima area in July and January are 21.7°C and —2.5°C, respectively (Franklin & Dyrness 1973).
Although populations of Icaricia acmon in this area of Washington have been called hybrids with I. lupini (Boisduval) (Goodpasture 1973), regional systematists refer to all populations in the state as I. acmon (J. P. Pelham pers. comm.). Icaricia acmon ranges throughout much of western North America, and has been recorded feeding on Polygonum and at least seventeen species of Eriogonum (both Polygonaceae) as well as Lotus, Astragalus, Lupinus, and Melilotus (all Fabaceae) (Scott 1986). Subspecies lutzi ranges from British Columbia south to central Oregon and east to central Colorado (Goodpasture 1973). In Washing- ton, I. acmon lutzi specializes on Eriogonum, having been recorded from E. compositum, E. pyrolifolium Hook., E. sphaerocephalum Dougl., and E. strictum, and has one or two broods, depending on the site (Peterson, unpubl. data). Larvae diapause in the third instar through winter and resume feeding when their host plant leafs out in the spring. Larvae feed on leaves or flowers by chewing holes in the surface of the structures and inserting their heads to mine out the internal tissues. Several ant species tend the third and fourth instar larvae of I. acmon, but earlier instars are not tended (Peterson pers. obs.). California pop- ulations of I. acmon have four instars and larvae have both a honey gland and eversible tentacles, structures associated with myrmecophily (Ballmer & Pratt 1988). Washington populations are similar in these regards (Peterson pers. obs.). It is not known when these structures first appear in development or if pupae of I. acmon are tended by ants.
METHODS
I determined host plant species of the study population of I. aemon by following ovipositing females and searching for eggs and young larvae. I observed 73 alightings on Eriogonum compositum and E.
10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
strictum, two of the three commonest Eriogonum species at the site. These alightings resulted in 10 ovipositions, with all eggs laid singly. I saw no encounters with other Eriogonum species at the site. In addition, I found eggs and larvae only on these two plant species.
To test the effect of ant attendance on larval demography, I selected 24 pairs of E. compositum and E. strictum in an area approximately 70 m X 380 m. Each pair was composed of one plant of each species, with individuals in a pair matched for size and occurring within 0.25 m of, but not in contact with, each other. To exclude ants, I encircled half of the pairs with rings of aluminum flashing which I had painted with Fluon® AD 1 (Northern Products, Inc., Woonsocket, RI), a sub- stance which forms a slippery coating on which arthropods cannot get traction. The base of the flashing was buried 1-2 cm below the soil surface and did not disturb the roots of the plants. In addition, because the soil at the study site had excellent drainage, the flashing could not have retained precipitation. Finally, the flashing was placed far enough from the plants to eliminate shading, and thus should have had no impact on plant quality. I left the remaining pairs without an arthropod exclosure as a control. Onto each of the plants, I placed either ova or early instar (first or early second) larvae of I. acmon. I placed two individuals on each plant, for a total of 96 individuals (18 ova and 83 larvae). It is unlikely that at such low densities, larvae would cannibalize each other, especially in light of the fact that in numerous rearings at much higher densities, I have seen cannibalism rarely in this species.
From 7 June to 18 September 1986, I conducted midday censuses at approximately weekly intervals; I resumed the censuses from 17 March to 26 April 1987 (19 censuses total: 13 in 1986 and 6 in 1987). During censuses, I noted the number of larvae on each plant and the species and number of attendant ants. Following the onset of the experiment, 35 larvae appeared in addition to those I had placed on the plants. Twenty-nine of these appeared between the onset of the experiment and 6 July, and the remaining six appeared by 6 August. Presumably, these larvae were already on the plants when I started the experiment or were from eggs that I did not see. I left these additional larvae on the plants because it was impossible to tell individual larvae apart. The distribution of these larvae was as follows: 9 larvae on E. strictum control plants, 11 on E. strictum exclosure plants, 18 on E. compositum control plants, and 2 on E. compositum exclosure plants. When larvae reached late 4th instar (the final instar), I collected them and raised them in the laboratory to obtain an estimate of the frequency of larval parasitism. On E. compositum, all of the larvae that reached late 4th instar did so from 18-26 April 1987. One larva matured by 6 July 1986 on E. strictum, but the remainder that reached late 4th instar did so
VOLUME 47, NUMBER 1 a!
from 5-26 April 1987. One of the ant-exclosure pairs was destroyed during the experiment, and larvae on these plants were excluded from the analysis of survivorship. To eliminate problems of autocorrelation, I analyzed ant attendance data with t-tests of mean per-plant attendance rates. Overall attendance rates were calculated using data from only the eight census dates on which at least one larva was tended. The reason for this is that on the days when no larvae were tended, it is likely that the lack of attendance had more to do with foraging con- ditions than larval attractiveness. Larvae were tended on 14, 21, 29 June and 6 July 1986 and 5, 12, 18, and 26 April 1987. For analyses of survivorship, I performed G-tests of independence, applying Williams’ 2 x 2 correction (Sokal & Rohlf 1981). Vouchers of I. acmon are deposited in lot #1198 in the Cornell University Entomology Collection.
RESULTS Ant Attendance
Three species of ants tended I. acmon larvae during this experiment: Tapinoma sessile (Say) (Dolichoderinae), Formica neogagates Emery, and an unidentified Formica species (both Formicinae). In all instances, only one ant species tended larvae on a single plant at a given time. Third and fourth instar larvae were tended by these ants, but I saw no first or second instar larvae tended during the censuses.
Tapinoma sessile is a small (2.5-3.5 mm long) dolichoderine ant which tended larvae singly or in groups of up to four ants. When disturbed during censuses, these ants ran to the base of the plant, abandoning the larva they were tending. Formica neogagates is larger (3.5-4.5 mm long) than T. sessile, tended I. acmon singly or in pairs, and was a much more aggressive tender than T. sessile. When disturbed, they assumed an alarm-defense posture (Wilson 1971) and would bite any object placed near them. The unidentified Formica sp. was similar in size and behavior to F. neogagates. J did not collect any specimens of this species. Because I saw only two of these ants and because of their similarity to F. neogagates, I combined these two species in the analysis of the composition of attendant ants by instar.
I saw no ant attendance from 13 July 1986 through 29 March 1987 (recall there were no censuses from 18 September 1986 to 17 March 1987); during this time larvae were quiescent and fed rarely. Midday soil temperatures were well in excess of 50°C throughout much of the summer and this may have restricted ant activity. On days when I saw ant attendance, third instar larvae were tended less than fourth instar larvae on both host species (Table 1). In addition, the species com position of attendant ants varied with instar on E. compositum, with the di-
12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TABLE 1. The incidence of ant attendance! of third and fourth instar Icaricia acmon larvae on Eriogonum compositum and E. strictum on the eight census dates when ants were observed.
Plant species Mean % tended SE N E. compositum? 8rd instar 43.3 7.0 ll Ath instar 88.9 6.5 7 E. strictum> 3rd instar 10.9 9.1 iil Ath instar 53.6 15.8 7
1 To avoid problems with autocorrelation, I determined the percentage of larvae tended on each control plant. Values are the means of these percentages.
at = 2.66, 16 df, P < 0.025.
bt = 2.52, 16 df, P < 0.025.
minutive Tapinoma sessile tending a greater proportion of third instar than fourth instar larvae (Table 2). On E. strictum, the composition of attendant ants varied similarly, but the differences were not statistically significant (Table 2). Most plants of both species had the same attendant species throughout the study. Interestingly, the six plants that had more than one ant species during the experiment all had Tapinoma sessile tending larvae early in the study and Formica neogagates tending larvae later.
Survivorship
Ants had no statistically significant effect on the survivorship of larvae to late fourth instar on either host species (Table 3). These data include those extra larvae that appeared after the onset of the experiment, and percentages are from data combined for all plants with similar treat- ments. All disappearances were interpreted as deaths for these measures of survivorship because it is unlikely that larvae leaving the plants in
TABLE 2. The percentage! of tended third and fourth instar Icaricia acmon larvae on Eriogonum compositum and E. strictum that were tended by Tapinoma sessile. The
remainder of the larvae were tended by Formica.
Mean % tended
Plant species by Tapinoma SE N
E. compositum?
3rd instar 92.9 Coll ill
Ath instar 30.6 16.3 Me E. strictum>
8rd instar 10.0) 25.0 11
Ath instar 40.0 24.5 t
1 The percentage of overall attendance was determined for each control plant and values are the means of these percentages.
at = 8.69, 11 df, P < 0.005. bt = 0.78, 5 df, P > 0.40.
VOLUME 47, NUMBER 1 13 TABLE 8. Survivorship of Icaricia acmon larvae to late 4th instar on ant-excluded and control plants of Eriogonum compositum and E. strictum.
Plant species Ant-excluded Control
E. compositum?
Survived 4 10 Died AM 25 % survival 15.4 28.6 E. strictum>
Survived LS 11 Died 20 20 % survival 45.5 Ope 4 Goorr = 1.46, P > 0.
= 0.2. Eee 06P > 0.5.
this harsh environment would survive. Only one parasitoid individual, an unidentified braconid, emerged from the larvae I raised in the lab.
DISCUSSION
The pronounced variability in this association over the course of larval development is notable; third instar larvae on both plant species ex- perienced lower overall attendance rates than fourth instar larvae, and the species composition of attendant ants changed with larval devel- opment. Although these results may reflect seasonal changes in the absolute and relative abundances of ants, it seems more likely that larger larvae are more attractive to ants. Fiedler (1989b) showed in a lab study that fourth instar larvae of Lycaena tityrus (Lycaenidae) are tended more frequently and with greater vigor than third instar larvae and suggested that an increase in the number and size of pore cupola organs was the cause for this increase. In addition to the pore cupola organs, the honey gland has been clearly demonstrated to play an important role in recruiting ants to lycaenid larvae (Fiedler & Masch- witz 1989a), and it is likely that large larvae produce more honeydew than small larvae. Finally, lycaenid larvae produce calls that serve to recruit ants (DeVries 1991a), and it is again likely that larger larvae could produce louder calls. Any or all of these factors could be important in determining the attractiveness of larval Icaricia acmon: pore cupola organs are widespread in the Lycaenidae (Henning 1983, Fiedler 1988); I. acmon is known to possess a honey gland (Ballmer & Pratt 1988); and in his survey of lycaenids, DeVries (1991a) found larval calling in several members of the Polyommatinae. If large larvae of Icaricia acmon are indeed more attractive to ants, the large, aggressive Formica neogagates may simply usurp these larvae from the small, docile Tap- inoma sessile.
14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
In examining the survivorship data, it is important to recognize that in this study and all other similar studies, the effect of ants on larval survivorship may be somewhat obscured because terrestrial predators were excluded along with ants. The only way to remove ants without removing these predators is to locate all ant nests and selectively isolate or remove them from half of the study area. It was impossible to perform this experiment because the nests of both F. neogagates and T. sessile are difficult to find. Nonetheless, this is the first experiment showing that lycaenid mortality may be the same in the presence and absence of ants and terrestrial predators. Because of the small sample sizes in this experiment, analyses lacked the power to detect small differences in survivorship between the treatment and control. Had the difference in survivorship between the treatment and control been greater than 25%, the analysis would have detected it. This magnitude of difference is comparable to that found by Pierce and Easteal (1986) for tended and untended larvae of Glaucopsyche lygdamus (Lycaenidae). Because survivorship of I. aemon larvae on E. strictum tended to be higher in the absence of ants, I feel that insufficient statistical power cannot explain entirely the apparent absence of a beneficial effect of ant at- tendance.
It is quite possible that I would have seen an effect of ant attendance on larval survival had I performed this study in a different place or time. DeVries (1991b) pointed out that variation in the abundances of natural enemies could influence whether riodinid and lycaenid larvae benefit from their associations with ants. In addition, host plant quality, ant abundance, and lycaenid abundance could be important in deter- mining overall benefits. Although temporal and spatial variation in benefits has not been examined in lycaenid-ant associations, Cushman and Whitham (1989) found that the benefits enjoyed by membracids from ant attendance varied markedly over a three year study period. It is clear from the high survivorship of larvae in this experiment that predation and parasitism pressures were low throughout the study pe- riod. Had I performed the experiment in a year or region of high larval mortality, I may have seen a difference between the treatments.
Finally, the differences in behavior of the two ant species suggest that they may differ in their effectiveness as defenders of larvae. Al- though the data here cannot address this issue, Bristow (1984) found that the ant species offering the best protection differed between an aphid and a membracid on New York ironweed. It would be interesting to perform a selective ant removal experiment to test whether the large, aggressive Formica species are more effective defenders of I. aemon larvae than are the smaller T. sessile.
VOLUME 47, NUMBER 1 15
ACKNOWLEDGMENTS
I thank P. Kareiva for much thoughtful advice on all aspects of this project. Jon Pelham kindly provided information on Washington populations of I. aemon. W. Carson, C. B. Cottrell, A. Herzig, G. Meyer, D. Peck, N. Pierce, R. Root, F. Sperling, and four anonymous reviewers contributed valuable comments on earlier drafts of this paper. C. McCullough offered useful advice on the statistical analysis of attendance data. I also thank L. Goncharoff, W. Morris, and J. Pearson for providing help and companionship in the field. R. Sugg identified the ants, for which I am grateful. This work was partially supported by an NSF REU grant to P. Kareiva.
LITERATURE CITED
ATSATT, P. R. 198la. Ant-dependent food plant selection by the mistletoe butterfly Ogyris amaryllis (Lycaenidae). Oecologia 48:60-63.
1981b. Lycaenid butterflies and ants: Selection for enemy-free space. Am. Nat. 118:638-654.
BALLMER, G. R. & G. F. PRATT. 1988. A survey of the last instar larvae of the Lycaenidae (Lepidoptera) of California. J. Res. Lepid. 27:1-81.
Bristow, C. M. 1984. Differential benefits from ant attendance to two species of Homoptera on New York ironweed. J. Anim. Ecol. 53:715-726.
CUSHMAN, J. H. & T. G. WHITHAM. 1989. Conditional mutualism in a membracid-ant association: Temporal, age-specific, and density-dependent effects. Ecology 70:1040- 1047.
DEVRIES, P. J. 1991a. Call production by myrmecophilous riodinid and lycaenid but- terfly caterpillars (Lepidoptera): Morphological, acoustical, functional, and evolu- tionary patterns. Am. Mus. Novitates 3025:1-23.
1991b. Mutualism between Thisbe irenea larvae and ants, and the role of ant ecology in the evolution of myrmecophilous butterflies. Biol. J. Linn. Soc. 43:179- 195.
FIEDLER, K. 1988. The preimaginal epidermal organs of Lycaena tityrus (PODA, 1761) and Polyommatus coridon (PODA, 1761) (Lepidoptera: Lycaenidae)—A comparison. Nota Lepid. 11:100-116.
1989a. European and North West African Lycaenidae (Lepidoptera) and their
associations with ants. J. Res. Lepid. 28:239-257.
1989b. Differences in the behaviour of ants towards two larval instars of Lycaena tityrus (Lepidoptera: Lycaenidae). Deut. Entomol. Z. 36:267-271.
FIEDLER, K. & U. MascHwitz. 1989a. Functional analysis of the myrmecophilous relationships between ants (Hymenoptera: Formicidae) and lycaenids (Lepidoptera: Lycaenidae) I. Release of food recruitment in ants by lycaenid larvae and pupae. Ethology 80:71-80.
1989b. The symbiosis between the weaver ant, Oecophylla smaragdina, and Anthene emolus, an obligate myrmecophilous lycaenid butterfly. J. Nat. Hist. 23: 833-846.
FRANKLIN, J. F. & C. T. DyRNEsS. 1973. Natural vegetation of Oregon and Washington. Forest Service, USDA, Portland, Oregon. 417 pp.
GOODPASTURE, C. 1973. Biology and systematics of the Plebejus (Icaricia) acmon group (Lepidoptera: Lycaenidae) I. Review of the group. J. Kansas Entomol. Soc. 46:468- 485.
HENNING, S. F. 1983. Chemical communication between lycaenid larvae (Lepidoptera: Lycaenidae) and ants (Hymenoptera: Formicidae). J. Entomol. Soc. South. Afr. 46: 341-366.
PiERCE, N. E. 1985. Lycaenid butterflies and ants: Selection for nitrogen-fixing and other protein-rich food plants. Am. Nat. 125:888-895.
PiERCE, N. E. & S. EASTEAL. 1986. The selective advantage of attendant ants for the larvae of a lycaenid butterfly, Glaucopsyche lygdamus. J. Anim. Ecol. 55:451-462.
16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
PIERCE, N. E. & M. A. ELGAR. 1985. The influence of ants on host plant selection by Jalmenus evagoras, a myrmecophilous lycaenid butterfly. Behav. Ecol. Sociobiol. 16: 209-222.
PIERCE, N. E. & P. S. MEAD. 1981. Parasitoids as selective agents in the symbiosis between lycaenid butterfly larvae and ants. Science 211:1185-1187.
PIERCE, N. E., R. L. KITCHING, R. C. BUCKLEY, M. F. J. TAYLOR & K. F. BENBOW. 1987. The costs and benefits of cooperation between the Australian lycaenid butterfly, Jalmenus evagoras, and its attendant ants. Behav. Ecol. Sociobiol. 21:237—248.
ScoTT, J. A. 1986. The butterflies of North America. A natural history and field guide. Stanford Univ. Press, Stanford, California. 583 pp.
SOKAL, R. R. & F. J. ROHLF. 1981. Biometry. W. H. Freeman and Co., New York. 859 pp.
WILSON, E. O. 1971. The insect societies. Belknap Press, Cambridge, Massachusetts. 548 pp.
Received for publication 23 September 1991; revised and accepted 12 September 1992.
Journal of the Lepidopterists’ Society 47(1), 1993, 17-21
BISTON BETULARIA (GEOMETRIDAE), THE PEPPERED MOTH, IN WIRRAL, ENGLAND: AN EXPERIMENT IN ASSEMBLING
Cyrit A. CLARKE
Department of Genetics and Microbiology, University of Liverpool, P.O. Box 147, Liverpool L69 3BX, England
FRIEDA M. M. CLARKE 43 Caldy Road, West Kirby, Wirral L48 2HF, England
AND
BRUCE GRANT
Department of Biology, The College of William and Mary, Williamsburg, Virginia 23185
ABSTRACT. The melanic form of the peppered moth, Biston betularia form “‘car- bonaria,” has continued to decline in frequency, comprising only 25.8% of a sample of 933 moths trapped on the Wirral peninsula near Liverpool, England, in 1991. The large sample was made possible, in part, by the use of two assembling traps and a mercury vapor trap. The assembling traps held either females of the North American subspecies, Biston betularia cognataria, or native British B. betularia females, thus allowing a com- parison of the relative effectiveness of the two forms in attracting local males. Our results indicate that British B. betularia males do not discriminate between the mating phero- mones released by the females of the two races.
Additional key words: “carbonaria,’ “cognataria,’ pheromone, industrial melanism.
Previous papers by Clarke et al. (1985, 1990) document the fall in frequency of the melanic form of the peppered moth, Biston betularia f. “carbonaria,’ at Caldy Common, West Kirby, on the Wirral peninsula near Liverpool, England, during the years 1959 through 1989. Figure 1 extends the census showing a slight “hiccup” in 1990 with “carbo- naria’ up from the previous year from 29.5% to 33.1%. The 1990 sample, however, was limited to only 154 moths, including 51 “car- bonaria,’ 6 intermediates (=f. “insularia’’), and 97 pale typicals. In 1991 “carbonaria”’ dropped precipitously to 25.8%, the lowest figure so far recorded.
The continued decline evidently reflects major habitat modification resulting from reduced industrial pollution accompanying the Clean Air Acts begun in the 1960’s although it remains unclear what ecological factors are involved. The typical form of the moth, once widely thought to gain protection from predators by its resemblance to gray foliose lichens, is rapidly replacing ‘“‘carbonaria’”’ as the common form in the virtual absence of such lichens (Clarke et al. 1985, Grant & Howlett 1988). Clarke et al. (1985) also noted a gradual lightening of the trees in the absence of industrial soot, and Grant and Howlett (1988) have
18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
total catches 283 431 994 478 394 610 207 318 269 480 631 407 689 860 444 219 933
cara bead oon al a eae al el anal Sr oral a Sie | males oe %of 95
carbonarla 90
85 80 75
70
Oe) CO “OS GS "BV We) 7 77 ‘75 ‘77 ‘79 ‘81 '83 ‘85 ‘87 ‘89 ‘91 year
Fic. 1. The decline in the proportion of the melanic form of the peppered moth, Biston betularia f. “carbonaria,” in West Kirby, Wirral, England. “Carbonaria” is ex- pressed as a percentage of a total in which typicals and “insularia’”’ are combined. The total number of B. betularia trapped in the 33 year period was 15,969 of which only 86 were female (0.54%).
further suggested that the comeback of birch trees (Betula pendula) following the Clean Air Acts may afford suitable hiding places for both the typicals and the melanics. Atmospheric sulfur dioxide concentrations at West Kirby have dropped markedly during this same period, and with minor fluctuations have remained low in recent years (see fig. 2 in Clarke et al. 1990). The mean winter SO, levels (ug per cubic meter) were 20.17 in 1988, 25.25 in 1989, 31.89 in 1990 and the mean for January and February 1991 was 44.39. Whatever the cause, the decline in the frequency of the melanic variant of this species at this location continues.
A second component of our study involved the method of trapping. All 1990 moths were trapped by MV light (mercury vapor lamp) because we had no female Biston betularia available that year for use in an assembling trap (AT). The AT is a cage containing virgin females to which local males are attracted by airborne pheromones released by the captive females, i.e., the female odor serves to lure males to the trap. In 1991 we used both MV and AT methods, and we also had the
VOLUME 47, NUMBER 1 19
opportunity of altering the assembling techniques by using virgin fe- male Biston betularia cognataria (Guenée) deriving from a large num- ber of pupae brought by one of us (BG) from Virginia, USA.
Biston betularia cognataria is the North American equivalent of Biston betularia (L.). It has typical and melanic (f. “swettaria’’) forms, as well as intermediates grouped together as “‘insularia’’ (see West 1977). However, typical cognataria generally are darker in appearance than British typicals, and, in fact, have been described as resembling inter- mediate-grade British “‘insularia’” (see Kettlewell 1973:plate 9.1). Ket- tlewell (1973) regarded cognataria and betularia as distinct species because of differences in color and in behavior in the early stages, there often being bivoltinism in the former but never in the latter. Rindge (1975), on the other hand, concluded that the evidence indicates that cognataria and betularia are members of the same biological species. They can be fully interbred and there is no disturbance of the sex ratio in the hybrids, and both male and female genitalia are identical or nearly so, as is the structure of the male antennae.
We used cognataria typicals from Virginia (about 20 at a time in the core of the trap and frequently replenished with fresh females as they emerged) to attract British B. betularia males throughout June 1991. Towards the end of this month the cognataria emergences were tailing off and the females old. Then our British B. betularia virgins (deriving from 20 pupae kindly supplied by Tony Liebert) began to eclose, and we used them in a different trap during July. Our original intention was to produce a quantitative bio-assay of relative pheromone effectiveness, but as we were unsuccessful in coordinating eclosions of the two kinds of females, we were unable to run both assembling traps simultaneously throughout both months. While the period of overlap when both cognataria and betularia females were available as lures in separate traps extended into July, we must emphasize that the number of individuals present in the cores, and their ages, unfortunately, were not controlled.
Nevertheless, the qualitative evidence that the cognataria pheromone is a powerful attractant to betularia males is compelling (Table 1). In June, when only cognataria females were releasing pheromone from the traps, 329 betularia males were drawn in as compared to only 109 males captured by the MV during that same period. During July, when both kinds of females were “calling” simultaneously from separate assembling traps, the numbers of males lured to the traps did not differ significantly by chi-square (144 to the cognataria-AT versus 171 to the betularia-AT, x? = 2.31, df = 1, 0.10 < P < 0.25). To our knowledge such data on comparative assembling have not been published previ- ously, and, as we have no reason to conclude that the mating pheromones
20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
TABLE 1. Biston betularia catches at West Kirby, Wirral, England, 1990 and 1991. Only the MV trap was used in 1990 as no virgin females were available. The 1991 combined catch figures also are subdivided to show numbers taken by each of the three traps used: a) MV trap; b) assembling trap using betularia females; c) assembling trap using cognataria females.
Year Trap used Total catch Carbonaria Insularia Typical % carbonaria 1990 MV only 154 ol 6 97 33.1 1991 All traps combined 933 241 37 655 25.8 199la MV trap 289 78 15 196 26.9 1991b betularia-AT 171 4] 4 126 23.9 1991c cognataria-AT 473 W22 18 333 25.8
of cognataria and betularia are different, our findings are consistent with Rindge’s (1975) assessment that cognataria and betularia are con- specific.
It is true that pheromones occasionally are not entirely species spe- cific, because Clarke (1979) showed that Orgyia thyellina Butler (Ly- mantriidae) females from Japan regularly assembled Orgyia antiqua (L.) males in the Wirral, though this is probably an exception to the general rule. The point, however, must be made that Priesner (1975) thought the pheromones of all Orgyia species were similar, but the findings were made by antennogram techniques which are not partic- ularly sensitive. If the pheromones in the genus Orgyia were alike there would be mating chaos where species fly together, and we know from observation that O. recens Hubner and O. antiqua do not attract each other (Greenberg et al. 1982). |
A final point in the 1991 series relates to the placement of the three traps. Jones, Majerus, and Timmins (unpublished) have produced evi- dence that in five polymorphic moth species in which melanism is thought to be of ancient origin, great sampling differences in morph frequencies over very short distances occur depending on the local environments in which traps are placed. They suggest that such habitat selection is not likely to be present where the melanism is of recent origin, as in Biston betularia, and our 1991 data support this view. Specifically, our MV trap was completely in the open, the cognataria- AT under a very thick, old oak tree, and the betularia-AT in a relatively exposed position near a small birch tree. Yet, all three morph frequencies were proportionately represented in all three trapping locations. In fact, the three data sets are remarkably homogeneous by G-test (G = 3.28, dit— 450) 5) <aR = 0275):
In summary, the 1991 sample of 933 Biston betularia at West Kirby shows a return to the lowering of the proportion of f. “carbonaria,” and we report that much of the assembling data resulted from using the American subspecies, B. b. cognataria.
VOLUME 47, NUMBER 1 . I
ACKNOWLEDGMENTS
We are grateful to Tony Liebert for providing B. betularia pupae; Sally Thompson for technical assistance throughout the study; R. J. Agass, Director of Housing and Environ- mental Protection, Metropolitan Borough of Wirral, for the SO, levels; M. E. N. Majerus for sending us a draft of his submitted manuscript; and the Nuffield Foundation for support.
LITERATURE CITED
CLARKE, C. A. 1979. Some observations on Orgyia thyellina Butler and Orgyia antiqua (L.). Entomol. Rec. 91:315-316.
CLARKE, C. A., G. S. MANI & G. WYNNE. 1985. Evolution in reverse: Clean air and the peppered moth. Biol. J. Linn. Soc. 26:189-199.
CLARKE, C. A., F. M. M. CLARKE & H. C. DAWKINS. 1990. Biston betularia (the peppered moth) in West Kirby, Wirral, 1959-1989: Updating the decline in f. carbonaria. Biol. J. Linn. Soc. 39:323-326.
GRANT, B. & R. J. HOWLETT. 1988. Background selection by the peppered moth (Biston betularia Linn.): Individual differences. Biol. J. Linn. Soc. 33:217-232.
GREENBERG, S., A. H. WRIGHT & C. A. CLARKE. 1982. Contrasting results in assembling experiments using Orgyia thyellina Butler, O. recens Hiibner and O. antiqua (L.). Entomol. Rec. 94:25-27.
KETTLEWELL, B. 1973. The evolution of melanism. Clarendon Press, Oxford. 423 pp.
PRIESNER, E. 1975. Electroantennogram responses to female sex pheromones in five genera of Lymantriidae (Lepidoptera). Z. Naturforsch. 30:676-679.
RINDGE, F.H. 1975. A revision of the New World Bistonini (Lepidoptera: Geometridae). Bull. Am. Mus. Nat. Hist. 156:69-155.
WeEsT, D. A. 1977. Melanism in Biston (Lepidoptera: Geometridae) in the rural Central Appalachians. Heredity 39:75-81.
Received for publication 17 March 1992; revised and accepted 31 August 1992.
Journal of the Lepidopterists’ Society 47(1), 1998, 22-31
TERRITORIALITY ALONG FLYWAYS AS MATE-LOCATING BEHAVIOR IN MALE LIMENITIS ARTHEMIS (NYMPHALIDAE)
ROBERT C. LEDERHOUSE
Department of Entomology, Michigan State University, East Lansing, Michigan 48824
ABSTRACT. A central New York population of Limenitis arthemis (Drury) was studied during June and July 1983. Males emerged a week before females with an overall male-biased sex ratio of 1.43:1.00. Of the 30 marked males, 7 (23%) were recaptured within the study area with an average longevity of 10.3 days. No marked female was recaptured. During midday mate-locating behavior, males perched an average of 82.5%, flew for 14.0%, and encountered other individuals for 3.5% of the time. Conspecific males were encountered at a rate of 8.3/h. Conspecific encounters averaged significantly longer than heterospecific encounters (12.8 vs. 5.8 sec). Marked males favored certain areas for perching but changed areas fairly frequently resulting in dynamic territories. Nearly half the perches were on sumac, and 68% were 1-3 m above the ground. Favored territories provided good vantage of female flyways.
Additional key words: activity budget, mark-recapture, protandry.
Territoriality in butterflies is a male tactic to locate receptive females (Powell 1968, Baker 1972, Davies 1978, Lederhouse 1982a, Wickman 1985a,b). Commonly, males defend landscape features such as hilltops and ridges that have high female visitation rates despite their lack of concentrated larval or adult resources (Shields 1968, Lederhouse 1982a, Alcock 1983, 1985, Alcock & O’Neill 1986, Alcock & Gwynne 1988, Rutowski et al. 1989). In addition, areas along butterfly flyways are defended (Baker 1972, Douwes 1975, Bitzer & Shaw 1988). Males of several species defend favorable microhabitats where females may raise their body temperature to facilitate activity (Davies 1978, Knapton 1985). Defense of feeding or oviposition resources appears to be un- common in butterflies (Baker 1972, Rutowski & Gilchrist 1988, Leder- house et al. 1992), except where they overlap with emergence sites (Dennis 1982).
Two subspecies of Limenitis arthemis (Drury) occur in eastern North America. Limenitis arthemis astyanax (Fabr.) is a Batesian mimic of the aposematic, distasteful pipevine swallowtail, Battus philenor (L.) (Papilionidae) (Platt et al. 1971, Codella & Lederhouse 1990). North of the geographic range of B. philenor, Limenitis arthemis arthemis has medial white wing bands. Wing banding is believed to offer pro- tection from predators through disruptive coloration although this has not been demonstrated experimentally (Silberglied et al. 1980). Nearly complete genetic mixing of the two subspecies occurs (Platt & Brower 1968, Platt 1983) except where geographic barriers to gene flow are
VOLUME 47, NUMBER 1 | 23
present (Waldbauer et al. 1988). In a band between 40° and 45°N latitude, females breed at random with males without regard to degree of male wing banding (Platt 1983).
Males of Limenitis species are notoriously aggressive (Pyle 1981, Lederer 1960). Male Limenitis weidemeyerii Edwards perch and en- gage passersby in territorial defense (Rosenberg 1989, Rosenberg & Enquist 1991). Male L. arthemis perch in the sun on trees and tall bushes and periodically patrol (Clark 19382, Ebner 1970). Although fidelity of male L. a. arthemis (Ebner 1970) and male L. a. astyanax (Harris 1972) to particular perches has been noted, Opler and Krizek (1984) stated that male L. a. astyanax do not seem faithful to particular sites. This study looked at male mate-locating behavior and population structure of Limenitis arthemis in a region of subspecies overlap. In particular, I investigated male longevity, location, aggressiveness, and activity patterns in relation to vegetation structure and local topography.
MATERIALS AND METHODS
The mate-locating behavior of Limenitis arthemis was studied near Brooktondale, Tompkins County, New York, during June and July 1983. The study site consisted of a gravel road with hedgerows on each side (Fig. 1). The general area was a mosaic of tilled fields and wooded areas.
Butterflies were captured with a net, marked individually, and re- leased immediately at the site of capture. Redundant marks were placed on both the right and left sides of the dorsal and ventral wing surfaces using red or green felt-tipped pens following a modification of Leder- house (1978). The identity of a butterfly could be determined at a distance of 3 m or less through observation of a perched or feeding individual. The term “‘recapture’”’ is used to denote the identification of a marked butterfly either by capture or observation on any day following the date marked.
The presence of Limenitis arthemis in the study area was monitored for a minimum of an hour on most sunny days of the study from 1030 to 1630 EDT. New individuals were captured and marked, and marked individuals were identified. The behavior of marked focal individuals was monitored continuously for 15 minute periods. The duration of each activity was recorded to the nearest second and the location noted. All Limenitis arthemis activity was recorded for consecutive 15 minute periods in each of three subunits favored by males. The order in which these subunits were observed was determined by random draw. Be- havioral observations were conducted during the periods of greatest Limenitis arthemis activity (1100 to 1500 EDT).
24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fic. 1. Map of the Brooktondale, New York, study area.
RESULTS
During the study, 30 males and 21 females were marked. Males first appeared about a week before females; 47% of the males were captured before the first female (Fig. 2). This protandry was significant (Kol-
VOLUME 47, NUMBER 1 , 25
—-A-— Male --V-- Female
Cumulative frequency
0 6 12 18 24 30 Days after first capture
Fic. 2. Cumulative frequencies of male and female captures at the Brooktondale, New York, study area per six day intervals. Day 1 of the generation was 21 June.
mogorovy-Smirnov two sample test, P < 0.01). Just over 23% of the 30 marked males were recaptured within the study area at least once. Of 7 males recaptured at least once, 71% were recaptured more than once. These 7 males were seen on an average of 4.0 (SE = 0.7) different days. Among multiply recaptured males, the average duration between first and last capture was 10.3 days (SE = 2.0), with three males observed over 17, 16, and 18 day periods. No marked female was recaptured. The overall sex ratio was male-biased (1.43:1.00). Using the method of Manly and Parr (1968) and Manly (1969), the male population was estimated to be 9.3 (SD = 2.0) males for day 7 and 9.0 (SD = 8.0) for day 12 of the study. Only 7% of males and 5% of females had the unbanded astyanax-like wing pattern.
Mate-locating behavior by males occurred primarily between 1100 and 1600 EDT. Activity budgets were calculated for a composite male based on 195 min during a total of 11 observation periods of 6 different marked males. Observations were made on sunny days between 1230 and 1500 EDT. The composite male perched 82.5% (SE = 3.8), flew 14.0% (SE = 8.2), and encountered other individuals 3.5% (SE = 0.9) of the time. Encounters with other species occurred at a rate of 1.9 per h; conspecific males were encountered at a rate of 8.3 per h. Hetero- specific encounters were usually brief chases, averaging 5.8 sec (SE =
26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
0.7, n = 6). Conspecific encounters were either longer chases or spiral flights and averaged 12.8 sec (SE = 1.0, n = 29). The difference in duration of conspecific and heterospecific encounters was significant (t-test, P < 0.01). Encounters between banded males (13.2 sec, SE = 1.8, n = 10) did not differ from those between banded and unbanded males (12.5 sec, SE = 1.2, n = 19).
Most encounters resulted from a perching male engaging another male that flew in the vicinity. The resident male returned from con- specific encounters to the same perch or one within 5 m in 938% of 27 cases. The challenging male returned within a minute after the end of an encounter in 32% of 25 cases. This usually resulted in repeated encounters until only the resident returned. At the study site, territories were linear arrays of perches along the hedgerows. The distance be- tween the two most separate perches for the 11 focal male samples averaged 4.2 m (SE = 0.6). Focal males achieved exclusive use of their defended area for 95.1% of the observed period (n = 195 min).
Males did not selectively perch on larval hosts. Only 7.7% of 142 perches were on the potential host, black cherry, Prunus serotina Ehrh. (Rosaceae). Perches were often on nonhosts staghorn sumac (Rhus ty- phina L.; Anacardiaceae) (48.6%) and white ash (Fraxinus americana L.; Oleaceae) (31.0%). The remaining perches were on foliage of non- host trees such as maple and elm, but even raspberry bushes, grape vines, corn, and goldenrod were used. Likelihood of males to perch on sumac during its period of blooming (July 1 to July 10) was not greater than before or after its blooming period (55.0%, n = 40, x? = 0.9, P > 0.3). Perches were 1-7 m from the ground (Fig. 3) with a mean of 2.9 m (SE = 0.2, n = 142). The most frequent classes were 2 m (33.8%) and 1 m (21.8%).
Certain locations within the entire study area (Fig. 1) were favored by perching males (Table 1). The southeast (SE) sampling subunit was defended during all but one of the observation periods, and twice as many males were observed in that subunit than in the next most used subunit. Males showed varying site fidelity (Table 1). For 7 males observed on multiple days, a male restricted to a subunit on one day was located in the same subunit on the next day in 62% of 21 possible cases. However, males voluntarily abandoned their territories for short periods. During the observation periods, 27% of 11 focal males were lost when they flew away from the area they had been defending, but were seen in the same areas later the same day.
More females were observed in the SW subunit, which had low male activity. During the study, two unsuccessful courtships were observed. On 27 June at 1318 EDT in the SE unit, a courted fresh female landed on a branch tip with her wings dorsally appressed. The male flew near
VOLUME 47, NUMBER 1 Pil
60 ” a 50 2 © > 40 — 2} ® % re Qo 30 Oo , om 20 wo} = = tO 0
1 2 3 4 5 6 7 Perch height (m)
Fic. 3. Distribution of male perches by height in meters. Total sample size is 142 perches.
the female for 10 sec, landed on the same branch, and tried to copulate. The female flew about 1 m and landed on the underside of a leaf with wings dorsally appressed. The male followed the female, landed, and again tried to copulate, but flew off when he was not successful. At 1353 h on 8 July in the NE subunit, another courted fresh female perched on the underside of a leaf, and the male was unsuccessful in his attempts to copulate.
Both males and females fed avidly at the flowers of staghorn sumac. Starting at 1428 h on 1 July, two marked males fed together without aggressive encounters although they were so close that there was some physical contact at the flowers. One female laid an egg on black cherry at 1249 h on 1 July, another laid an egg on apple, Pyrus malus L. at
TABLE 1. Male behavior and location of individuals in regard to subunits of the study area. The first two parameters are for 12 0.25-h periods in each area. The last three parameters are for the entire study. Fidelity is the percent of resightings within the same unit. Total females include captures and observations.
SE NE SW % samples area defended 92 67 50 Total observed males 20 10 7 Initial male captures 11 2 13 % male fidelity to area 82 0 44
Total females 8 il 13
28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
1248 h on 2 July, and another laid two eggs on choke cherry, Prunus virginiana L. starting at 1354 h on 5 July. An additional seven eggs and small larvae were found on choke cherry, mostly in the SW subunit.
DISCUSSION
Males of Limenitis arthemis defended territories as mate-locating behavior in central New York. Males were localized during midday and early afternoon. Site fidelity from day to day was generally high. Male chases of other males served to secure nearly exclusive use of these sites for the resident male. As in other territorial butterflies (Davies 1978, Rutowski & Gilchrist 1988, Rosenberg & Enquist 1991), the res- ident generally won these encounters. Conspecific encounters lasted over twice as long as those with other butterflies. The duration of conspecific encounters was quite similar to that for L. weidemeyerii (Rosenberg & Enquist 1991), and seems more than adequate to deter- mine the species and sex of the intruder (Scott 1974). Territory turnover was somewhat higher than seen in territorial males of some species (Lederhouse 1982a) but similar to that seen in others (Lederer 1960, Alcock & O’Neill 1986, Rutowski et al. 1989, Lederhouse et al. 1992). Apparently, voluntary abandonment of defended areas was relatively frequent.
The area most regularly defended did not correspond to where most females were observed. The area with the greatest number of females had more trees including more host plants. Five females were observed in oviposition flight in that area. Areas defended by males in this study were independent of larval hosts as in Limenitis camella and L. populi (Lederer 1960) or adjacent to hosts as in L. camella, L. populi, and L. weidemeyerii (Lederer 1960, Rosenberg 1989, Rosenberg & Enquist 1991). Also, males did not change where they defended during the blooming period of sumac. This suggests that areas most regularly defended are flyways, but not necessarily concentrations of hosts or nectar plants. This is further supported by the preponderance of male perches on nonhosts. Although copulations were not observed during this study, two mating refusal interactions were seen in the early af- ternoon. Shull (1987) found mating pairs of L. a. astyanax at a similar time.
The probability that a marked male would be recaptured was lower than that reported for many territorial species but greater than that reported for patrolling species (Lederhouse 1982b). However, for those males that were recaptured once, the probability of further recaptures was similar to that for other territorial species, as was the number of times those males were seen in the study area. The average residency of territorial males in this study is quite similar to that for the black
VOLUME 47, NUMBER 1 29
swallowtail, Papilio polyxenes Fabr. (Papilionidae) from the same area (Lederhouse 1983). The two astyanax-like males of L. a. astyanax seen in the study area were observed over 16 and 18 day periods, two of the three longest. Central New York is north of the usual range of Battus philenor although late season strays are regularly seen there (Shapiro 1974). It is worth further study to determine whether the longevity of these two males was merely coincidental or related to their phenotype.
Although most individuals in this study were banded, their behavior encompassed published accounts for both L. a. astyanax and L. a. arthemis. Banded and unbanded males clearly recognized each other as competitors for the same territories. Encounter durations between unbanded and banded males did not differ from those between banded males, but both were significantly longer than encounters with other species. This is consistent with the apparent panmixia that occurs where the two subspecies come into contact (Platt & Brower 1968, Platt 1983). Although all females that were observed ovipositing were banded, they laid on hosts usually considered to be hosts of L. a. astyanax (Pyle 1981).
ACKNOWLEDGMENTS
I thank M. Ayres, J. Scott, and an anonymous reviewer for helpful criticism of the manuscript. F. Sperling and K. Mikkola translated the Lederer reference. Completion of this project was supported by National Science Foundation Award BSR 91-07139.
LITERATURE CITED
ALCOCK, J. 1983. Territoriality by hilltopping males of the great purple hairstreak, Atlides halesus (Lepidoptera: Lycaenidae): Convergent evolution with a pompilid wasp. Behav. Ecol. Sociobiol. 18:57-62.
1985. Hilltopping in the nymphalid butterfly Chlosyne californica (Lepidop- tera). Amer. Mid]. Nat. 113:69-75.
ALCOCK, J. & D. GWYNNE. 1988. The mating system of Vanessa kershawi: Males defend landmark territories as mate encounter sites. J. Res. Lepid. 26:116-124. [“1987.”]
ALCOCK, J. & K. M. O'NEILL. 1986. Density-dependent mating tactics in the grey hairstreak, Strymon melinus (Lepidoptera: Lycaenidae). J. Zool. 209:105-113.
BAKER, R. R. 1972. Territorial behaviour of the nymphalid butterflies, Aglais urticae (L.) and Inachis io (L.). J. Anim. Ecol. 41:453—469.
BITZER, R. J. & K. C. SHAw. 1980. Territorial behavior of the red admiral, Vanessa atalanta (L.) (Lepidoptera: Nymphalidae). J. Res. Lepid. 18:36-49.
1983. Territorial behavior in Nymphalis antiopa and Polygonia comma (Nym- phalidae). J. Lepid. Soc. 37:1-18.
CLARK, A.H. 1932. The butterflies of the District of Columbia and vicinity. Smithsonian Institution U.S. Natl. Museum Bulletin 157. 337 pp.
CODELLA, S. G. & R.C. LEDERHOUSE. 1990. The effect of wing orientation on aposematic signalling in the pipevine swallowtail butterfly, Battus philenor. Anim. Behav. 40: 404-406.
Davies, N. B. 1978. Territorial defense in the speckled wood butterfly (Parage aergeria): The resident always wins. Anim. Behav. 26:188-147.
DENNIS, R. L. H. 1982. Mate location strategy in the wall brown butterfly, Lasiommata megera L. (Lep: Satyridae). Wait or seek? Entomol. Rec. J. Var. 94:209-214.
30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
DouwEs, P. 1975. Territorial behaviour in Heodes virgaureae L. (Lep., Lycaenidae) with particular reference to visual stimuli. Norw. J. Entomol. 22:143-154.
EBNER, J. A. 1970. The butterflies of Wisconsin. Milwaukee Public Museum, Milwaukee, Wisconsin. 205 pp.
Harris, L. 1972. Butterflies of Georgia. Univ. Oklahoma Press, Norman, Oklahoma. 326 pp.
KNAPTON, R. W. 1985. Lek structure and territoriality in the chryxus arctic butterfly (Satyridae). Behav. Ecol. Sociobiol. 17:389-395.
LEDERER, G. 1960. Verhaltensweisen der Imagines und der Entwicklungsstadien von Limenitis camilla camilla L. (Lep. Nymphalidae). Z. Tierpsych. 17:521-546.
LEDERHOUSE, R. C. 1978. Territorial behavior and reproductive ecology of the black swallowtail, Papilio polyxenes asterius Stoll. Ph.D. Diss., Cornell University, Ithaca, New York. 155 pp. Diss. Abstr. Int. order No. 7902342.
1982a. Territorial defense and lek behavior of the black swallowtail butterfly,
Papilio polyxenes. Behav. Ecol. Sociobiol. 10:109-118.
1982b. Factors affecting equal catchability in two swallowtail butterflies, Papilio
polyxenes and P. glaucus. Ecol. Entomol. 7:379-383.
1983. Population structure, residency and weather related mortality in the black swallowtail butterfly, Papilio polyxenes. Oecologia 59:307-811.
LEDERHOUSE, R. C., S. G. CODELLA, D. W. GROSSMUELLER & A. D. MACCARONE. 1992. Host plant-based territoriality in the white peacock butterfly, Anartia jatrophae (Lepidoptera: Nymphalidae). J. Insect Behav. 5:721-728.
MANLY, B. F. J. 1969. Some properties of a method of estimating the size of mobile animal populations. Biometrika 56:407—410.
MANLY, B. F. J. & M. J. PARR. 1968. A new method of estimating population size, survivorship, and birth rate from capture-recapture data. Trans. Soc. Brit. Entomol. 18:81-89.
OPLER, P. A. & G. O. KRIZEK. 1984. Butterflies east of the great plains: An illustrated natural history. Johns Hopkins University Press, Baltimore, Maryland. 294 pp. PLATT, A. P. 1983. Evolution of North American admiral butterflies. Bull. Entomol.
Soc. Amer. 29(8):10-22.
PLATT, A. P. & L. P. BROWER. 1968. Mimetic versus disruptive coloration in intergrading populations of Limenitis arthemis and astyanax butterflies. Evolution 22:699-718.
PLATT, A. P., R. P. COPPINGER & L. P. BROWER. 1971. Demonstration of the selective advantage of mimetic Limenitis butterflies presented to caged avian predators. Evo- lution 25:692-701.
POWELL, J. A. 1968. A study of area occupation and mating behavior in Incisalia iroides (Lepidoptera: Lycaenidae). J. New York Entomol. Soc. 76:47-57.
PYLE, R. M. 1981. The Audubon Society field guide to North American butterflies. Knopf, New York. 916 pp.
ROSENBERG, R. H. 1989. Behavior of the territorial species Limenitis weidemeyerii (Nymphalidae) within temporary feeding areas. J. Lepid. Soc. 43:102—107.
ROSENBERG, R. H. & M. ENQuisT. 1991. Contest behaviour in Weidemeyer’s admiral butterfly Limenitis weidemeyerii (Nymphalidae): The effect of size and residency. Anim. Behav. 42:805-811.
RUTOWSKI, R. L. & G. W. GILCHRIST. 1988. Male mate-locating behavior in the desert hackberry butterfly, Asterocampa leilia (Nymphalidae). J. Res. Lepid. 26:1-12. “1987.” ]
RuTowskt, R. L., J. ALCOCK & M. Carey. 1989. Hilltopping in the pipevine swallowtail butterfly (Battus philenor). Ethology 82:244-254.
Scott, J. A. 1974. Mate-locating behavior of butterflies. Amer. Mid]. Nat. 91:103-117.
1974. Butterflies and skippers of New York State. Cornell Univ. Agric. Exper- iment Stat. Search 4(3):1-60.
SHIELDS, O. 1968. Hilltopping. J. Res. Lepid. 6:69-178. [~1967.”]
SHULL, E. M. 1987. The butterflies of Indiana. Indiana Acad. Sci., Bloomington, Indiana. 262 pp.
VOLUME 47, NUMBER I Bil
SILBERGLIED, R. E., A. AIELLO & D. M. WINDsor. 1980. Disruptive coloration in butterflies: Lack of support in Anartia fatima. Science 209:617-619.
WALDBAUER, G. P., J. G. STERNBURG & A. W. GHENT. 1988. Lakes Michigan and Huron limit gene flow between the subspecies of the butterfly Limenitis arthemis. Canad. J. Zool. 66:1790-1795.
WICKMAN, P.-O. 1985a. Territorial defence and mating success in males of the small heath butterfly, Coenonympha pamphilus L. (Lepidoptera: Satyridae). Anim. Behav. 33:1162-1168.
1985b. The influence of temperature on the territorial and mate locating be-
haviour of the small heath butterfly, Coenonympha pamphilus (L.) (Lepidoptera:
Satyridae). Behav. Ecol. Sociobiol. 16:233-238.
Received for publication 20 March 1992; revised and accepted 21 September 1992.
Journal of the Lepidopterists’ Society 47(1), 1993, 32-48
DEVELOPMENTAL CHANGES AND WEAR OF LARVAL MANDIBLES IN HETEROCAMPA GUTTIVITTA AND H. SUBROTATA (NOTODONTIDAE)
DAWN E. DOCKTER Center for Biodiversity, Illinois Natural History Survey, Champaign, Illinois 61820
ABSTRACT. Detailed descriptions and scanning electron micrographs of worn and unworn larval mandibles and physical measurements of cuticle loss owing to wear within each instar were made for Heterocampa guttivitta (Walker) and H. subrotata (Harvey) (Notodontidae). First instar mandibles of both species lack a retinaculum and are used to skeletonize leaf tissue. The mandibles of later instars have a retinaculum and are more robust. The latter are used to cut through the leaf blade. The retinaculum of H. subrotata is bifurcate as opposed to the simple retinaculum of H. guttivitta. The teeth and reti- naculum on the mandibles of instars two through five of both species are almost completely worn down during their short period of use; the length of mandibles is reduced by at least 20 percent during the final larval instar.
Additional key words: saddled prominent, systematics, morphology.
Mandibular characters of lepidopterous larvae have been used in many taxonomic keys (Gardner 1946a, 1946b, Beck 1960, Godfrey 1972, Brown & Dewhurst 1975) and in phylogenetic studies of various No- todontidae (Godfrey et al. 1989). However, relatively little research has focused on the functional morphology and feeding ecology of lepidop- terous larval mandibles (Tragardh 1913, Bernays & Janzen 1988), and no study has accounted for wear of these structures. Thus, the reliability of some mandibular characters may be questionable. The purpose of this study was to document the ontogenetic changes of larval mandibles of Heterocampa guttivitta (Walker) and H. subrotata (Harvey) (No- todontidae) and to determine the extent of wear on the mandibles within each larval stadium of both species.
Heterocampa guttivitta, the saddled prominent, is an occasional pest in hardwood forests in the northeastern part of the United States. In the past it has caused reduction in yields in the sugar maple and lumber industries. Heterocampa guttivitta has five larval instars and overwin- ters in the pupal stage. In the northeast it has one generation per year, with the moth emerging in May-June (Forbes 1948). In central Illinois there are two generations per year, with the moths emerging May- June and July—August, based on collection records in the Illinois Natural History Survey (G. L. Godfrey pers. comm., Illinois Natural History Survey, Champaign, Illinois). The larvae have been reported feeding on Acer (Aceraceae), Betula (Betulaceae), Carya (Juglandaceae), Cas- tanea dentata (Fagaceae), Corylus (Corylaceae), Fagus (Fagaceae), Hamamelis virginiana (Hamamelidaceae), Juglans (Juglandaceae), Malus pumila (Rosaceae), Ostrya virginiana (Corylaceae), Populus (Salicaceae), Prunus (Rosaceae), Pyrus (Rosaceae), Quercus (Fagaceae),
VOLUME 47, NUMBER 1 33
Rubus (Rosaceae), Spiraea (Rosaceae), Ulmus (Ulmaceae), and Vibur- num (Caprifoliaceae) (Tietz 1972).
Heterocampa subrotata is the smallest Heterocampa species in North America. The “evenly green form” or variety “celtiphaga” (Forbes 1948) was used in this study. Larval hosts have been reported as Acer, Betula, Carya, Cornus (Cornaceae) and Hamamelis virginiana (Teitz 1972). Heterocampa subrotata also has five larval instars and overwin- ters as a pupa. It appears to have two generations in central Illinois, with adults emerging in May-June and July-August.
METHODS AND TECHNIQUES
Collecting and Rearing
To ascertain the feeding histories of the larvae used in this study, larvae were reared from eggs deposited by wild females. Three Het- erocampa guttivitta and five H. subrotata females were collected at a UV-light trap in Trelease Woods, Champaign County, Illinois. One H. guttivitta was collected at a similar UV-light trap in Wolf Creek State Park, Shelby County, Illinois. All life stages of both Heterocampa species were maintained in an insectary and were exposed to ambient tem- perature and humidity during the summer of 1989 in Champaign, Illinois.
Voucher specimens of both study species are placed in the Illinois Natural History Survey Insect Collection, Champaign, Illinois. Voucher specimens include the pinned adult females from which eggs were obtained, as well as alcoholic specimens of the following: eggs just prior to hatch, intact larvae from each stadium, and head capsules from the exuviae of all five larval stadia. George L. Godfrey verified all species identifications.
Individual field-collected females were placed into one-ounce plastic diet cups with cardboard lids. Each cup contained a strip of brown paper towel which served as a resting spot for the moth and as a possible Oviposition site.
Four cohorts of Heterocampa guttivitta and five cohorts of H. sub- rotata were reared. Each cohort consisted of approximately 45 indi- viduals. To reduce mortality attributable to handling, fresh leaves and the diet cup containing unhatched eggs were placed into a rearing container. Rearing containers were plastic Solo (P550) cups with a depth of 5 cm, a top diameter of 8 cm, and a bottom diameter of 5 cm. The cups were fitted with clear plastic lids. Newly hatched first instar larvae were allowed to crawl onto the leaves. After one or two days of feeding, the leaves were cut so that each section had only one larva on it. Each leaf section with a single larva was placed in a separate rearing container with additional leaves. The addition of several leaves to a container
34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
helped minimize the loss of leaf moisture. The larvae were reared individually in the described containers until reaching the prepupal stage.
An unknown percentage of larvae of both Heterocampa species in this study were infected with a cytoplasmic polyhedrosis virus (CPV) (J. V. Maddox pers. comm., Illinois Natural History Survey). Individuals in six of the eight cohorts reared for this study displayed symptoms consistent with those typical of CPV infections.
The CPV infection did not produce a high mortality in the larval cultures and was not diagnosed until most of the larvae had reached late third instar. Larvae were discarded and not used in this study if they displayed any symptoms consistent with a CPV infection. No attempt was made to histologically determine if larvae used were in- fected with CPV. The assumption was made that a CPV infection that produced no physically visible symptoms would have little or no effect on mandibular morphology or on the general nature of how mandibles wear with use.
Heterocampa guttivitta was reared on white oak (Quercus alba L., Fagaceae) and Heterocampa subrotata was reared on northern hack- berry (Celtis occidentalis L., Ulmaceae). White oak foliage was re- placed every third day or earlier if the existing foliage appeared to be dry. Northern hackberry was replaced every day or every other day depending on foliage condition. Dark green, mature leaves of northern hackberry and white oak were collected fresh each morning.
Prepupae were placed in half-pint cardboard ice cream containers (with no more than three individuals of the same species per container) that contained an approximately 2-cm layer of sand covered with ap- proximately 4 cm of moist peat moss. Larval exuviae were carefully removed from the pupal chambers and stored in 70-percent ethanol until the mandibles could be dissected from the head capsule and measured.
Unworn mandibles were collected from newly molted larvae. It was difficult to obtain larvae that had not fed unless they were isolated from the foliage before ecdysis. This was done by carefully cutting a small section from the leaf that contained the silk mat and larva. The leaf section, silk mat, and any remaining exuvia were removed after the larva had completed molting and begun moving around the container. Larvae were not used if they had fed on the dry leaf sections after molting. However, a majority of the larvae did eat part of their exuviae. The assumption was made in this study that feeding on exuvia caused negligible wear; no effort was made to keep the larvae from feeding on it or to determine if any wear was produced by this type of feeding.
Newly molted larvae were held without foliage for 8-12 hours before
VOLUME 47, NUMBER 1 35
being preserved. Mandibles that were not allowed to harden in this manner were difficult to dissect without tearing or otherwise damaging them. Larvae were killed in boiling water and subsequently preserved in 70-percent ethanol until the head capsules could be measured and the mandibles dissected and measured. Both the left and right mandibles were removed from intact, preserved larvae as described by Godfrey (1972). If the mandibles were not completely sclerotized at the time of preservation, the abductor muscle was cut where it attached to the mandible, but the adductor muscle was removed with the mandible. Excess muscle tissue was removed before the mandibles were measured.
Mandibles from exuviae were used to represent the worn condition because they could be obtained in a non-destructive manner, and be- cause they potentially demonstrate maximum wear. Head capsules were collected from the rearing containers with a small camel’s-hair paint brush and placed directly into 70-percent ethanol until they could be measured and the mandibles could be removed and measured. Man- dibles were removed from molted head capsules by severing any cu- ticular attachments and lifting out the mandibles with a curved dis- secting tool. They were returned to 70-percent ethanol after being measured.
Scanning Electron Microscopy Preparation
Mandibles were washed in three changes of 70-percent ethanol and then sonicated in a 1:1] solution of Photo-Flo and 70-percent ethanol for 30 seconds. Seventy-percent ethanol was used in the sonicating fluid to keep the mandibles from floating or sticking to the sides of the container above the fluid line during sonication. Sonication times of over 30 seconds occasionally result in lost or damaged setae. After sonication, the mandibles were washed three to five times in 70-percent ethanol and were then dehydrated in a graded ethanol series: 85%, 95%, and three times in 100% (10 minutes in each concentration). The specimens were critical-point dried and attached to stubs using alu- minum tape. The tape worked very well for attaching the small samples, which had a tendency to sink into liquid adhesives. However, aluminum tape produces a light-colored background, which may interfere with the viewing and photographing of the specimen. The specimens were coated three times with gold-palladium. Each coating was for 30 seconds at 30 mA. An AMRAY 1830 scanning electron microscope operating at 10 kV was used to view the specimens.
Statistical Analysis
Six to fifteen randomly chosen pairs of mandibles from each species, representing each stage of development (lst to 5th instar) and both
36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
states of wear (unworn and worn) were measured in order to determine the extent of wear. Two measurements were taken of each mandible: the distance between the condyle and the adductor apodeme (CI) and the distance from the condyle to the tip of the second tooth (CT) (Figs. 1-2). Teeth were not discernible on worn mandibles from stadia three, four, and five. In those cases, the point most distad from a line con- necting the condyle and the adductor apodeme was used in place of the second tooth (Figs. 1-2). CT and CI were taken for both right (R) and left (L) mandibles to produce four groups of measurements (RCT, LCT, RCI, and LCI) for each pair of mandibles. CI measurements were taken to ensure that changes in the length of the mandible (CT) were due to wear and not due to changes that resulted from sclerotization.
All measurements were taken at the highest practical magnification for each group (e.g., all first instar RCI and LCI for H. subrotata were taken at 120x) through a dissecting stereomicroscope with an ocular micrometer that had been calibrated with a stage micrometer.
Means and standard deviations for RCI, LCI, RCT, and LCT were calculated for each state of wear in each instar. An unpaired t-test was used to test for differences between means of unworn and worn man- dibles. A paired t-test was used to test for differences between right and left mandibles from the same larva.
RESULTS Mandible Descriptions
Notodontid mandibles are hollow structures with the distal (cutting) edge heavily sclerotized. They appear dark caramel brown in color under a dissecting stereomicroscope. The proximal end is open. There are two points of articulation on the mandible (Figs. 3-5). The condyle (C) articulates with the subgena and the acetabulum (A) (=socket) articulates with the lateral part of the clypeus.
There are four surfaces on the mandible: lateral, dorsal, ventral, and oral. These surfaces are defined with respect to the orientation of the mandible to the caterpillar. There is a textured area on the lateral surface, which is located in a slight depression. Two setae (M1 and M2) (Beck 1960) can be found in this area (Figs. 4-5).
The oral (=inner or mesal) surface was the emphasis of this study. Leaf cutting occurs on the extreme distal edge of the mandible. This edge may be smooth or it may have a series of teeth (dentes). The teeth are numbered from the ventral to the dorsal surface with Arabic nu- merals (Fig. 3) (Godfrey 1972). A retinaculum (R) or inner ridge is usually present in notodontids (Fig. 3). The retinaculum is part of the heavily sclerotized distal end of the mandible.
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C\
2
Fics. 1-2. 1, Worn and 2, Unworn 8rd instar mandibles of Heterocampa guttivitta. Dorsal view of left mandible showing how measurements were taken for analysis of wear. Measurement CI was used to indicate absolute size of mandible. Measurement CT was used to measure wear-related changes.
Description of Heterocampa guttivitta Mandibles
The unworn first instar mandible is laterally flattened and has five distinct teeth. These teeth, with the exception of the fifth, are usually pointed. The third and fourth teeth have flanges on the bases of their ventrolateral edges (Fig. 8). The first instar mandible lacks a retinac- ulum (Figs. 6-8). The teeth are greatly shortened in worn first instar
Fics. 83-5. Unworn 8rd instar mandible of Heterocampa guttivitta. Three views of the left mandible. 3, Oral surface; 4, Dorsal surface; and 5, Lateral surface. A = ace- tabulum (socket), AdAp = adductor apodeme, C = condyle, DP = distal pits, M1 and M2 = setae, R = retinaculum, Tl, T2, T38, T4, and T5 = teeth.
38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fics. 6-9. Oral surfaces of 6, unworn and 7, worn left mandibles of Ist instar Heterocampa guttivitta. 8, A view of the proximal end of the left 1st instar Heterocampa guttivitta mandible shows the flanges (arrows) on ventral-lateral sides of teeth 3 and 4; 9, Oral surfaces of worn left mandibles of lst instar Heterocampa subrotata. Micron bar = 20 microns.
VOLUME 47, NUMBER 1 39
mandibles, however, they are usually still discernable. Usually, the cuticle on much of the oral surface appears rough (Fig. 7). This rough- ness is probably a result of wear on the cuticle.
Second instar mandibles have five truncate teeth (Fig. 10). Teeth one and two are not very distinct from one another and often appear fused. The fifth tooth has a series of ridges on its inner surface. There are two distal pits (Fig. 10) on the mandible’s oral surface that lie between teeth two and three, and three and four. The second instar mandible is laterally flattened, but the presence of a retinaculum makes it appear more robust than the first instar mandible. The retinaculum is a simple ridge with several dentes at the dorsal end. The retinaculum is worn smooth, often to the point that there is little or no concavity between it and the outer cutting edge (Fig. 11). The cutting edge wears to a - smooth, continuous edge. Instars three through five appear to wear in a manner similar to that of the second instar.
The third instar mandible has four truncate teeth (Fig. 12). The first tooth is broad and smooth and may be the result of the fusion of two teeth. The fourth tooth is crenulate. In this instar, there are distal pits between teeth one and two, two and three, and three and four. The distal end of the mandible is curved at approximately a 90 degree angle, causing the cutting edge to be projected mediad. This angle makes the third instar mandible appear more robust than the second instar man- dible. The retinaculum is a simple crenulate ridge.
The fourth and fifth instar mandibles are very similar to those of the third instar, except that the distal teeth are not as distinctly separate. The crenae on the retinaculum of the fifth instar mandible are quite small and, in some cases, the retinaculum appears almost smooth.
Description of Heterocampa subrotata Mandibles
The unworn and worn mandibles of H. subrotata appear very similar to those of H. guttivitta with the following exceptions. H. subrotata mandibles are smaller than those of H. guttivitta (Table 1) and the retinaculum is strickingly different. The retinaculum of the second instar mandible is quite small and has a “hook” at the end. The surface of the retinaculum is irregular. Occasionally, there are cone-shaped projections located on the retinaculum (Fig. 14). There are five, short, rather pointed teeth on the third instar mandible (Fig. 15). The dorsal end of the retinaculum has a series of cone-shaped projections. The arrangement of these projections makes the end of the retinaculum appear bifurcate. The height and number of these projections are vari- able. However, the location of each projection in relation to the other projections appears to be fixed. The retinaculum of the fourth instar mandible ends in a definite bifurcation (Fig. 16). The branches of the
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fics. 10-13. Oral surfaces of 10, unworn and 11, worn left mandibles of 2nd instar Heterocampa guttivitta. Distal pits (DP) are located between teeth two and three, and three and four of the unworn mandible. The concavity between the distal cutting edge and the retinaculum is greatly reduced in the worn mandible. Oral surface of 4th instar mandibles of 12, Heterocampa guttivitta (unworn) and 13, H. subrotata (worn). Micron bar = 100 microns.
VOLUME 47, NUMBER | 4]
Fics. 14-17. Distal end of unworn mandibles of 14, 2nd instar; 15, 3rd instar; 16, 4th instar; and 17, 5th instar Heterocampa subrotata. Micron bar = 100 microns.
retinaculum and the retinaculum for a short distance before the bifur- cation are crenate. Some of the crenae are fairly sharp. The fifth instar mandible (Fig. 17) differs from the fourth instar mandible in that its retinaculum is dentate its entire length and is more strongly bifurcate.
Right Mandible Versus Left Mandible
The results from the paired t-test showed that there was no statis- tically significant difference (P > 0.10) between RCT and LCT or between RCI and LCI for unworn or worn mandibles of H. guttivitta at the level of P = 0.10, except between the 5th instar RCI and LCI measurements for unworn (P = 0.0669) and worn (P = 0.0422) man- dibles. There was no statistically significant difference (P > 0.10) be- tween RCT and LCT or between RCI and LCI for unworn mandibles of H. subrotata. No significant difference (P > 0.10) for 6 out of 10 measurements was found between RCT and LCT or between RCI and LCI for worn mandibles of H. subrotata. The four exceptions were as follows: 1st instar RCT vs. LCT (P = 0.0137), 2nd instar RCI vs. LCI (P = 0.0891), 4th instar RCT vs. LCT (P = 0.0380), 5th instar RCI vs. LCI (P = 0.0294). No major morphological differences between the left and right mandibles were found (Dockter 1991). Therefore, only the
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
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VOLUME 47, NUMBER 1 43
TABLE 2. The width of left mandibles as measured from the condyle to the adductor apodeme (Fig. 1) for Heterocampa guttivitta and H. subrotata. The measurements (CI), in mm, of unworn and worn mandibles are combined for each instar of each species.
Species/instar Range Mean? SD n
H. guttivitta
1 0.108-0.120 0.1135 0.003 20 2 0.251-0.279 0.262 0.008 21 3 0.393-0.473 0.436 0.019 23 4 0.626-0.748 0.684 0.028 25 5 1.005-1.139 1.054 0.045 17 H. subrotata 1 0.097-0.114 0.104 0.005 18 2 0.182-0.222 0.203 0.010 24 3 0.285-0.348 0.318 0.017 27 4 0.444-0.531 0.493 0.023 27 5 0.687-0.792 0.730 0.026 23
2 All means for worn mandibles were not significantly different from means for unworn mandibles at the level of 0.05 unless otherwise noted. b Significantly different at 0.05 but not at 0.01 (P = 0.0140, P = 0.0413, P = 0.0474, respectively).
left mandible was used for quantitative and qualitative descriptions in this paper (Tables 1-2).
Mandible Wear
First instar mandibles of both Heterocampa species were reduced in length by at least 5.2 to 5.5% (Table 3). The amount of wear increased in the second instar to 13.5 to 16.4% (Table 3). Mandible lengths of the final larval instar mandible are reduced by 19.2 to 20.5% (Table 3). The LCT length for larval mandibles from newly molted larvae of H.
TABLE 3. The percent of unworn mandible length that is lost during a larval instar for Heterocampa guttivitta and H. subrotata. Means from Table 1 were used to calculate percent length lost.
Species/instar % length lost
H. guttivitta
Re) 16.4 18.1 16.7 20.5
op WN
H. subrotata
5.2 13.5 13.9 16.6 19.2
OR WN
44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
guttivitta was significantly longer (P < 0.01) than the LCT length of H. guttivitta mandibles taken from the exuviae (Table 1). Unworn LCT length of H. subrotata was also significantly longer (P < 0.01) than worn LCT length (Table 1). There was no statistically significant dif- ference (P < 0.01) between unworn and worn LCI in H. guttivitta or in H. subrotata (Table 2). A worn mandible surface appears strikingly different from that of an unworn surface. At relatively low magnifi- cations (x 300), an unworn surface appears smooth in contrast to the rough surface of a worn mandible. High magnification of a worn man- dible (Figs. 18-19) demonstrates that portions of the cuticle are flaked from the surface.
The absolute difference between the length (CT) of unworn and worn mandibles does not represent the actual amount of cuticle loss due to wear. The amounts of wear reported in this study are conservative because there is a bend between the proximal and distal ends of the mandible. It is not possible to compare amounts of wear between species or instars from the data collected in this study because the angle between the proximal and distal areas varies between instars, and especially between species. The difference between unworn LCT and worn LCT becomes more representative of actual cuticle loss as the angle between the proximal and distal areas increases.
DISCUSSION Developmental and Behavioral Changes Within Species
The change from a shovel-shaped first instar mandible that has a sharply serrate distal margin but lacks a retinaculum, to a second instar mandible that is more robust at the mid- and proximal areas and bears a retinaculum, is associated with a change in feeding habits. This change occurs in both Heterocampa guttivitta and H. subrotata.
The following description of feeding behavior is based on observations made in the laboratory during the course of the present study. First instar larvae are leaf skeletonizers: they feed on the lower (occasionally upper) epidermis of the leaf while leaving the rest of the leaf tissue intact. The first instar uses the sharp serrations on the mandible to break through the epidermis. The head is quickly and forcefully tilted slightly to one side. This motion is repeated at the same spot on the leaf until the mandible penetrates the epidermis. Once the epidermis is broken a mandible is slid through the leaf tissue and the mandibles are closed to break off a piece of the leaf. Second instar larvae begin feeding on the leaf edge and bite through its entire thickness. During the early part of the second instar, larvae avoid the major veins, but toward the end of this instar, they eat through almost all veins except the midvein.
VOLUME 47, NUMBER 1
Fics. 18-19. 18, Detail of distal end of worn 4th instar Heterocampa guttivitta left mandible. Note the difference between unworn (u) and worn (w) cuticle. 19, High magnification of worn area of same mandible. Micron bar = 100 microns.
46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
A discussion of the structural changes of the mouthparts (especially mandibles) and the behavioral differences associated with them for two other species of notodontids can be found in Godfrey (1991).
The teeth on the cutting edge become more truncate and less sharply defined with successive instars in both species studied. The amount of pressure needed to break through the leaf tissue is a function of the amount of force exerted per unit area. Pressure can be increased by reducing the area over which the force is exerted or by increasing the force. When the larvae are small, a sharp tooth may be important to increase pressure so that the leaf tissue can be penetrated. As the larvae get larger, the muscles are capable of producing more force, and the extra pressure generated by a sharper more pointed edge may become unnecessary.
Developmental and Behavioral Changes Between Species
The ontogenetic development of the retinaculum in H. subrotata is quite different from that in H. guttivitta. The retinaculum on the mandibles of H. subrotata has a more intricate pattern than the reti- naculum of H. guttivitta. H. subrotata feeds on a more succulent host (northern hackberry) than H. guttivitta (white oak). Bernays and Janzen (1988) found a correlation between host texture and mandible mor- phology. They found that sphingids, which tend to feed on rather succulent leaves, have mandibles that are “long, toothed, and ridged in a variety of complex ways,” whereas saturniids have “short and simple’? mandibles and feed on “old, tough tannin-rich leaves.” The present study tends to support the findings of Bernays and Janzen (1988).
However, more work needs to be done in this area. It would be interesting to determine if the larvae Nemoria arizonaria Grote (Ge- ometridae), which exhibit caterpillar morphs that feed on two very different tissues of the same host, also show a change in mandibular structure to best exploit the host tissue. Greene (1989), in his work with N. arizonaria, found that the catkin morph, which is a pollen feeder, has a smaller head and jaws (mandibles) than the twig morph, which feeds on “leathery” oak leaves. Bernays (1986) found that head mass and mandible mass were significantly greater in grass-feeding lepidop- terous larvae which feed on a tougher diet. One might assume that cuticle production increases when the size of the head and jaw (man- dible) increases. It may be that the same factors that control these changes in allometry may also trigger mandibular polymorphisms: if mandibular polymorphisms are in fact shown to exist.
An advantage of an intricate retinaculum is that it may allow the host tissue to be broken into smaller pieces than would be possible with a more simple retinaculum. Bernays and Janzen (1988) indicate that
VOLUME 47, NUMBER 1 AT
the smaller the pieces produced by the biting process, the more nutrients that could be extracted from a given amount of tissue, because there is no further mechanical breakdown of food in the gut. However, Bernays (1991) in response to Barbehenn’s (1989) study which found that bite size was not correlated with digestibility, concludes that han- dling time may be more important than bite size. Bernays (1991) states, “Tough leaves are efficiently handled by the snipping, scissor action, whereas the softer more flaccid leaves are more efficiently ingested by the tearing, crushing action’.
Mandibular Characters in Systematics
Godfrey et al. (1989) used the presence or absence of teeth on the mandible’s distal edge to make hypotheses about phylogenetic rela- tionships among notodontids. However, before systematic (phyloge- netic) decisions are made or characters are described with respect to mandibles, one must be certain that unworn mandibles are being ex- amined, especially when a character is easily altered by wear. Never- theless, for the purpose of identification of caterpillars, it is necessary to describe the most common form of the mandibles, which is often the worn condition in field-collected larvae.
ACKNOWLEDGMENTS
I thank G. L. Godfrey and W. G. Ruesink for their guidance and support. I also thank J. E. Appleby, M. Berenbaum, J. Nardi, J. Sternburg, D. W. Onstad and C. Renfrew for their contributions to my work. I wish to thank M. Gillott and L. Crane for advice on specimen preparation for the SEM work and their help with the SEM. I also acknowledge the Illinois Natural History Survey and the Center for Electron Microscopy at the Uni- versity of Illinois for the use of their SEMs and their facilities. This project was supported by the University of Illinois Research Board and the Joyce Foundation.
LITERATURE CITED
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BECK, H. 1960. Die Larval systematik der Eulen (Noctuidae). Abhandl. Larvalsyst. Ins. (Akademie-Verlag, Berlin) 4:1—406.
BERNAYS, E. A. 1991. Evolution of insect morphology in relation to plants. Phil. Trans. R. Soc. Lond. B 333:257-264.
1986. Diet-induced head allometry among foliage-chewing insects and its im- portance for graminivores. Science 231:495-497.
BERNAYS, E. A. & D. H. JANZEN. 1988. Saturniid and sphingid caterpillars: Two ways to eat leaves. Ecology 69:1153-1160.
BROWN, E. S. & C. F. DEwHursST. 1975. The genus Spodoptera (Lepidoptera, Noctuidae) in Africa and the Near East. Bull. Entomol. Res. 65:221-262.
DockTER, D. E. 1991. Developmental changes and wear of larval mandibles of Het- erocampa guttivitta (Walker) and Heterocampa subrotata (Harvey) (Notodontidae). M.S. Thesis, Univ. Illinois, Urbana-Champaign.
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1946b. On larvae of the Noctuidae (Lepidoptera)—II. Trans. Roy. Entomol. Soc. Lond. 97:237-252.
GopFREY, G. L. 1972. A review and reclassification of larvae of the subfamily Hadeninae (Lepidoptera, Noctuidae) of America north of Mexico. USDA, Tech. Bull. 1450. 265 pp.
1991. Functional morphology of the larval mouthparts of Notodontidae (Lep- idoptera): Contrasting the structural differences and feeding behaviors of Hetero- campa obliqua and Crinodes besckei. Entomol. Soc. Amer. Symposium Proc. In press.
GODFREY, G. L., J. S. MILLER & D. J. CARTER. 1989. Two mouthpart modifications in larval Notodontidae (Lepidoptera): Their taxonomic distributions and putative func- tions. J. New York Entomol. Soc. 97:455-470.
GREENE, I. 1989. A diet induced developmental polymorphism in a caterpillar. Science 243:643-645.
SNoDGRASS, R. E. 1935. Principles of insect morphology. McGraw-Hill, New York. 667 pp.
TrETZ, H. M. 1972. An index to the described life histories, early stages and hosts of the Microlepidoptera of the continental United States and Canada, 2 vol. Allyn Museum of Entomology, Sarasota, Florida. 1041 pp.
TRAGARDH, I. 1918. Contributions towards the comparative morphology of the trophi of the lepidopterous leaf miners. Ark. Zool. 8:1-49.
Received for publication 2 June 1992; revised and accepted 20 September 1992.
Journal of the Lepidopterists’ Society 47(1), 1998, 49-54
DESIGNATION OF A LECTOTYPE OF NISONIADES SOMNUS AND NOTES ON THE OCCURRENCE OF ERYNNIS ICELUS IN FLORIDA (HESPERIDAE)
JOHN V. CALHOUN!’ 1731 San Mateo Drive, Dunedin, Florida 34698
ABSTRACT. Nisoniades somnus Lintner was described in 1881 from one male and one female from “Indian River, Florida.’ Neither specimen was identified as the holotype, therefore a lectotype and paralectotype are hereby designated. Dubious reports of Erynnis icelus from Florida also are examined.
Additional key words: Erynnis brizo, type locality, paralectotype.
Over a century ago, J. A. Lintner described a distinctive Floridian skipper as Nisoniades somnus (Lintner 1881). This taxon currently is considered a subspecies of Erynnis brizo (Boisduval & LeConte) and is restricted to the Florida peninsula (Burns 1964). The description was based on one male and one female from “Indian River, Florida” (given ambiguously as ‘Florida’ by Miller and Brown [1981]) deposited in the collection of W. H. Edwards. The types were undoubtedly collected by Dr. William Wittfeld (1827-1913) and/or his daughter Annie M. Wittfeld (1865-88) of Georgiana, Brevard County, Florida, who were regular correspondents of Edwards and the source of his “Indian River” records. The Wittfelds began collecting Lepidoptera for Edwards in 1880 (dos Passos 1951), thus the specimens probably were captured during the spring of 1880 or 1881.
In his original description, Lintner compared somnus almost exclu- sively to Erynnis icelus (Scudder & Burgess), rather than E. brizo. As a result, subsequent authors (e.g., Edwards 1884, Skinner 1898, Dyar 1902, Smith 1891, 1908) associated somnus more closely with E. icelus, alluding to a relationship between the two. This perceived relationship is surprising considering that Lintner (1881) himself revealed in the same paper that males of both somnus and E. brizo lack hair tufts on the hind tibiae, a structure present in E. icelus. Blatchley (1902) sum- marized the general opinion regarding these taxa when he remarked that somnus was “‘closely allied” to E. icelus and ‘may be only a large southern form.”
For many years following its original description, somnus was known from very few localities and most authors (e.g., French 1885, Maynard 1891, Skinner 1898) continued to list this taxon only from the type locality. An exception was Scudder (1889) who listed “‘Thanaos brizo”’
' Research Associate, Florida State Collection of Arthropods, Gainesville, Florida.
50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
from Florida and included the additional locality of “Haulover.” This record was provided by E. A. Schwarz, probably as a result of his visits to Florida in 1875 and 1876 (Schwarz 1888). This reference is especially interesting because Haulover formerly existed in northern Brevard County, approximately 22 km north of Georgiana, where the type specimens of N. somnus probably originated. Schwarz obviously rec- ognized the similarity of his specimens to E. brizo and identified them as such. This was the first glimpse into the true relationship between these taxa.
Dyar (1905) was the first to openly suggest that somnus was “perhaps but a dark form of brizo”’ and noted the resemblance of their genitalia. This notion was supported by Skinner (1914) who also commented on the similarity of their genitalia. F. E. Watson (in Grossbeck 1917) more confidently submitted that somnus is “probably a subspecies of brizo.” Following the acceptance of somnus as a subspecies of E. brizo by Barnes and McDunnough (1917), this taxonomic status was generally adopted. However, Holland (1931) stated that he was “unable to agree with this opinion” and retained the mistaken belief that somnus was “much nearer to T. icelus.”
Lintner (1881) did not designate either of his specimens of Nisoniades somnus as the holotype. Miller and Brown (1981) were unaware of the location of Lintner’s syntypes although Skinner (1914) stated that they were deposited in the Carnegie Museum of Natural History, where they remain today. These specimens were figured by Holland (1931: plate 51, figs. 3-4) who identified each as “type.” Both specimens lack antennae (the male retains a portion of the left antenna) which were noticeably drawn onto the Holland figures. The specimens are in good condition, except the abdomen of the female is now detached and pinned with the specimen in a dry vial. The male specimen (Fig. 1) (left forewing length, base to apex = 15 mm) is hereby designated as the lectotype. It bears three labels: ““Nisoniades/Somnus, 6/Lintn./ TYPE.” in Lintner’s hand; “Collection/W. H. Edwards” printed; and “Butterfly Book/PI. 51 Fig. 8,” printed and handwritten. I have affixed a red label declaring the specimen as the lectotype. The female spec- imen (Fig. 2) (left forewing length, base to apex = 16 mm) is designated as a paralectotype. It also bears three labels: ““Nisoniades/ Somnus, @/Lintn./ TYPE.” in Lintner’s hand; “Collection/W. H. Edwards” printed; and “Butterfly Book/PI. 51 Fig. 4,” printed and handwritten. A red label has been affixed to indicate its status as paralectotype. The type locality is restricted to Georgiana, approximately 5 km south of the city of Merritt Island, Brevard County, Florida. An additional male specimen of E. b. somnus was figured by Holland (1898, 1931:plate 48, fig. 2). This specimen, from the W. H. Edwards collection, is labelled
VOLUME 47, NUMBER 1 ol
Mt, llzt akg oe zeelPe. j Se |\7YX¥ PE TN Collection | 3! , a
W. H. Edwards |
| t
Butterfly Book
PILLS YS -Fig.4e
Fics. 1-2. Nisoniades somnus Lintner. 1, Lectotype male; 2, Paralectotype female.
in Edwards’ hand as “somnus/é/Ind. Riv.” and is considered a topotype.
Unlike most of Edwards’ specimens, the types of N. somnus do not possess locality data. Edwards did not place labels on his individual specimens until he sold his collection to W. J. Holland in the late 1880's (Brown 1964). At that time, he prepared labels that typically included the name of the species, sex of the specimen and a brief (sometimes cryptic) mention of the location of capture. Edwards probably did not affix such labels to the N. somnus types because Lintner’s labels already were present.
The difficulty experienced by most nineteenth century lepidopterists in recognizing distinct differences between E. b. somnus and E. icelus contributed to confusion over the distribution of E. icelus that haunted the literature for 80 years. Edwards (1884) casually listed E. icelus from “Fla,” regardless of the fact that his closest record was from Illinois. Subsequent authors, including French (1885), Maynard (1891), Skinner (1898) and Holland (1898) followed Edwards and continued to include Florida within the range of E. icelus. Scudder (1889) implied a reluc- tance to accept Florida reports when he remarked that “Edwards also gives it from Florida.” Apparently, Scudder had not seen any specimens of E. icelus from Florida, nor had he received any such reports from his many correspondents. Blatchley (1902) reported that he collected “several” E. icelus (supposedly determined by H. Skinner) in the spring of 1889 at Ormond, Volusia County, Florida (he listed E. b. somnus
o2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
separately). Not until the treatises on the Hesperioidea by Lindsey (1921) and Lindsey et al. (1931) did the Floridian reports finally become unacceptable. The furthest south from which these authors reported E. icelus was North Carolina. However, the saga continued when Macy and Shepard (1941) resurrected the Floridian reports and Evans (1953) indicated that the British Museum (Natural History) contained E. icelus from Florida. Forbes (1960) also listed E. icelus from Florida, possibly on the authority of Evans. Burns (1964) examined the purported Flo- ridian specimen of E. icelus in the British Museum, a dateless male from the R. Oberthtr collection marked only as “Floride,’ and con- sidered it mislabelled. Burns added that “E. icelus has often been attributed to Florida, chiefly in older literature; the error seems to stem from Edwards. Many highly questionable locality records (and food- plant records as well) have been uncritically repeated, in literature bearing on the Erynnis, to the extent that nowadays they may appear to be reliable, when actually they are not.” Although Kimball (1965) included a contemporary record (1961) of E. icelus from the Florida panhandle (determined by W. T. M. Forbes as “apparently this’) he retorted “I am much in doubt as to whether this species is really native to Florida.”
The basis of the early reports of E. icelus in Florida probably can be traced to a small female specimen of E. b. somnus from the W. H. Edwards collection labelled ““Nisoniades/icelus(?)/Lintn./?/Ind. Riv.” in Edwards’ hand. The specimen was undoubtedly collected by the Wittfelds at Georgiana, Brevard County, Florida at about the same time the types of Nisoniades somnus were collected (ca. 1880). This supports Skinner (1914) who suggested that Floridian records of E. icelus may actually be E. b. somnus. Improperly identified skippers are epidemic within early collections and even remotely similar species were confused. This problem is exemplified by H. G. Dyar who de- termined as E. b. somnus a Mississippi specimen of Erynnis zarucco (Lucas) (Burns 1964). However, this inherent identification problem does not entirely solve the Floridian E. icelus dilemma.
Six male specimens of E. icelus, bearing handwritten and printed labels reading “Fla” from the W. J. Holland collection, are deposited in the Carnegie Museum of Natural History (identifications verified by genitalic examination). Three of these specimens also possess hand- written labels reading “Morrison,” apparently in reference to the nine- teenth century collector Herbert K. Morrison. Morrison collected in Florida in 1883, 1884 and 1885 (Essig 1931). Morrison also visited at least ten other states between 1874 and 1888 (Essig 1981), all of which possess valid records of E. icelus (Burns 1964). Morrison was a prolific collector and such zeal increases the potential for accidental mislabel-
VOLUME 47, NUMBER 1 53
ling. Nonetheless, the validity of these specimens is difficult to ascertain, especially since no similarly labelled specimens in the Carnegie Museum are thought to be mislabelled (J. E. Rawlins pers. comm.). These six specimens are probably the basis for Holland’s (1898, 1931) inclusion of Florida within the range of E. icelus.
There is a very remote possibility that E. icelus occurred (or occurs) in northern Florida, especially the panhandle where habitats of more northern affinities occur. However, valid specimens of this species are not known from south of northern Georgia (Burns 1964, Opler & Krizek 1984). Unless additional evidence is revealed, the six Floridian speci- mens of E. icelus will remain an enigma.
ACKNOWLEDGMENTS
Iam grateful to John E. Rawlins of the Carnegie Museum of Natural History for helpful information and the loan of specimens. Thanks are also extended to Timothy L. McCabe of the New York State Museum for his verification of J. A. Lintner’s handwriting. John M. Burns and an anonymous reviewer critically reviewed the manuscript and provided many helpful suggestions.
LITERATURE CITED
BARNES, W. & J. MCDUNNOUGH. 1917. Check list of the Lepidoptera of boreal America. Herald Press, Decatur, Illinois. 392 pp.
BLATCHLEY, W. S. 1902. A list of the butterflies taken in the vicinity of Ormond, Florida, in March and April, 1899, pp. 227-233. In Blatchley, W. S. (ed.), A nature wooing at Ormond By The Sea. The Nature Publ. Co., Indianapolis, Indiana. 245 pp.
BROWN, F. M. 1964. The types of the satyrid butterflies described by William Henry Edwards. Trans. Amer. Entomol. Soc. 90:323-413.
Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif. Publ. Entomol. 37:1-216.
Dos Passos, C. F. 1951. The entomological reminiscences of William Henry Edwards. J. New York Entomol. Soc. 59:129-186.
Dyar, H. G. 1902. A list of North American Lepidoptera and key to the literature of this order of insects. Bull. Smiths. Inst. No. 52. 723 pp.
1905. A review of the Hesperiidae of the United States. J. New York Entomol. Soc. 13:111-141.
EDWARDS, W.H. 1884. Revised catalogue of the diurnal Lepidoptera of America north of Mexico. Trans. Amer. Entomol. Soc. 11:245-337.
Essic, E.O. 1931. A history of entomology. The Macmillan Co., New York. 1029 pp.
EvANs, W. H. 1953. A catalogue of the American Hesperiidae, indicating the classifi- cation and nomenclature adopted in the British Museum (Natural History). Part III. British Museum (Natural History), London, England. 246 pp., pls. 26-53.
FORBES, W. T. M. 1960. Lepidoptera of New York and neighboring states. Part IV, Agaristidae through Nymphalidae including butterflies. Cornell Univ. Agric. Exp. Sta., Memoir 371. Ithaca, New York. 188 pp.
FRENCH, G. H. 1885. The butterflies of the eastern United States. J. B. Lippincott Co., Philadelphia, Pennsylvania. 402 pp.
GROSSBECK, J. A. 1917. In Watson, F. E. (ed.), Insects of Florida IV. Lepidoptera. Bull. Amer. Mus. Nat. Hist. 37(Article 1):1—47.
HOLLAND, W. J. 1898. The butterfly book. Doubleday, Page & Co., New York. 382 pp.
1931. The butterfly book, new and thoroughly revised edition. Doubleday &
Co., Inc., Garden City, New York. 424 pp.
o4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
KIMBALL, C. P. 1965. Arthropods of Florida and neighboring land areas. Vol. 1. Lep- idoptera of Florida. Div. of Plant Industry, Gainesville, Florida. 363 pp.
LINDSEY, A. W. 1921. The Hesperioidea of America north of Mexico. Univ. Iowa Studies Nat. Hist. (First Ser. No. 43) 9(4):1-114.
LINDsEY, A. W., E. L. BELL & R. C. WILLIAMS, JR. 1931. The Hesperioidea of North America. Denison Univ. Bull., J. Sci. Lab. 26:1-142.
LINTNER, J. A. 1881. On some species of Nisoniades. Papilio 1:69-74.
Macy, R. W. & H. H. SHEPARD. 1941. Butterflies. Univ. Minnesota Press, Minneapolis. 247 pp.
MAYNARD, C. J. 1891. A manual of North American butterflies. De Wolf, Fiske and Co., Boston, Massachusetts. 226 pp.
MILLER, L. D. & F. M. BRowN. 1981. A catalogue/checklist of the butterflies of America north of Mexico. Lepid. Soc. Memoir No. 2. 280 pp.
OPLER, P. A. & G. O. KRIZEK. 1984. Butterflies east of the Great Plains. The Johns Hopkins Univ. Press, Baltimore, Maryland. 294 pp.
SCHWARZ, E. A. 1888. The insect fauna of semitropical Florida with special regard to the Coleoptera. Entomol. Amer. 4:165-175.
SCUDDER, S. H. 1889. The butterflies of the eastern United States and Canada with special reference to New England. Vol. II. Lycaenidae, Papilionidae, Hesperidae [sic.]. Published by the author, Cambridge, Massachusetts:767-1774.
SKINNER, H. 1898. A synonymic catalogue of North American Rhopalocera. Amer. Entomol. Soc., Philadelphia, Pennsylvania. 100 pp.
1914. Studies in the genus Thanaos. Trans. Amer. Entomol. Soc. 40:195-221.
SMITH, J. B. 1891. List of the Lepidoptera of boreal America. Amer. Entomol. Soc., Philadelphia, Pennsylvania. 124 pp.
1903. Check list of the Lepidoptera of boreal America. Amer. Entomol. Soc.,
Philadelphia, Pennsylvania. 136 pp.
Received for publication 25 June 1992; revised and accepted 20 September 1992.
Journal of the Lepidopterists’ Society 47(1), 1993, 55-59
FIRST NORTH AMERICAN RECORDS OF EPINOTIA ABBREVIANA (TORTRICIDAE), A EUROPEAN PEST OF ULMUS SPECIES
Pi DANG
% The Canadian National Collection of Insects, Centre for Land and Biological Resources Research, Agriculture Canada, K. W. Neatby Building, Ottawa, Ontario K1A 0C6, Canada
ABSTRACT. Epinotia abbreviana (Fabricius), recently found established in New- foundland, Canada, is described and diagnosed, with illustrations of genitalia and wings.
Additional key words: diagnosis, distribution, host plant, genital structure, New-
foundland.
Epinotia abbreviana (Fabricius) is native to Europe. The larval stages of this species feed on various species of Ulmus (Ulmaceae). Two male specimens were reared from larvae collected in 1981 on elm in St. John’s, Newfoundland, Canada; in 1988 two females were reared from larvae collected on Ulmus rubra Muhl. in Bowring Park, St. John’s. The second collection indicates that this species has become established in the area.
Both series of specimens were collected by personnel of the Forest Insect and Disease Survey (FIDS) from Forestry Canada’s Newfound- land and Labrador Region. The species was identified by the author based on detailed examinations of all four specimens. Specimens col- lected from England (1 6) and Germany (1 2) also were examined to lend further support to the identification.
The description, illustrations, photographs, and review of biological aspects of this species provided in this article will help researchers to recognize and identify the pest. This information will be particularly useful for surveying and monitoring the species in St. John’s and neigh- boring areas. Descriptions and illustrations of various morphological aspects of the species also can be found in Bradley et al. (1979), Kuz- netsov (1978), Graaf Bentinck and Diakonoff (1968), Hannemann (1961), Benander (1950), Pierce and Metcalfe (1922), and Kennel (1921).
DIAGNOSTIC FEATURES
Description. Epinotia abbreviana is a variable species. The forewing of specimens collected in Newfoundland exhibits the two extremes of variation, which ranges from a pale form with distinct and contrasting markings to a dark form with an almost uniform dark gray-brown forewing and faint markings. Bradley et al. (1979) provided a series of wing illustrations showing the variability of this species in England. The ISCC-NBS (Inter-Society Color Council—National Bureau of Stan-
56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Fics. 1-2. Dorsal aspect of E. abbreviana adults; 1, form 1; 2, form 2.
dards) Color-Name Charts (Kelly & Judd 1955) are used in the following descriptions.
Head. Vertex, antenna medium brown to dark gray-brown. Frons white. Labial! palpus medium brown on apical third, other areas yellow-white to white.
Thorax. Notum yellow-brown to dark gray-brown. Pleural area yellow-white. Forewing 7.0-7.5 mm long, without costal fold. Form 1 (Fig. 1): Forewing mostly medium brown; area between medial fascia and basal patch yellow-white, extended across wing, shaped like “greater-than” or “less-than”’ sign; costal strigulae well defined, with alternating yellow-white and medium brown comma-shaped spots; area between postmedial and subterminal fasciae, and that between subterminal and terminal fasciae, narrow, silver- gray, extending from dorsal end of terminal margin to costal margin at two-thirds length from wing base, and from terminal margin at one-fourth length from wing apex to costal margin at three-fourths length from wing base, respectively; tornal area with silver-gray ocellus. Form 2 (Fig. 2): Forewing medium brown in most areas, fasciae obsolete; as with form 1, tornal ocellus and two oblique lines in areas between postmedial and terminal fasciae silver-gray, but faintly visible; costal strigulae dull, consisting of alternating yellow- brown and medium brown spots. Hindwing uniform gray-brown; fringe paler. Form 3: Similar to form 2, except much darker, dark gray-brown. Legs in all forms with fore- and midlegs medium brown, except basal and apical margins of femur, tibia, tarsomeres, and mid area of tibia white; hindleg yellow-white, except tarsomeres medium brown basally.
Abdomen. Medium brown to dark gray-brown dorsally.
Male genitalia (Figs. 3-4): Mesal surface of valva with numerous, short, stout setae along ventral margin of sacculus, and with slender, dorsally directed setae on remaining areas. Tegumen well developed, triangular. Uncus long and slender, gradually tapered apically, gently curved ventrally, with small, inconspicuous bifid apex. Socius well de- veloped, with dense, posterodorsally directed, long setae; apex, as seen laterally, bluntly convex. Aedeagus cylindrical; cornuti long, well sclerotized, distinctly curved apically, 10-11 in number.
Female genitalia (Figs. 5-6): Papillae anales foot-shaped, distinctly arched anterodor- sally, fringed laterally with long, hooked setae. Lamella postvaginalis small, densely spiculate. Colliculum long, cylindrical, slightly sclerotized, smooth, as long as non-scler- otized part of ductus bursae. Corpus bursae voluminous, potato-shaped, slightly wider than long, with 2 well-sclerotized, horn-shaped, anteromedially directed signa, one on each side; surface of corpus bursae finely reticulate.
Remarks. The male genitalia of Epinotia abbreviana are similar to those of E. sperana McDunnough, E. myricana McDunnough, E. eth- nica Heinrich, E. ulmicola Kuznetsov, E. solandriana (Linnaeus), and
o7
VOLUME 47, NUMBER 1
uncus and
Male genitalia of E. abbreviana; 3, lateral aspect of tegumen,
socius; 4, posteroventral aspect with both valvae spread.
Fics. 3-4.
Fics. 5-6. Female genitalia of E. abbreviana; 5, ventral aspect; 6, lateral aspect.
58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
E. trigonella (Linnaeus); the first three species are native to North America, the third is found in south of the Primorye Territory of Russia, and the last two are widespread in the Holarctic region. Specimens of E. ulmicola were not available for study. However, according to Kuz- netsov (1966), E. ulmicola is distinguished from E. abbreviana (sensu E. trimaculana Donovan) by the following characters: apex of uncus simple, bifid in E. abbreviana; distoventral angle of sacculus obtuse in mesal view, approximately 90° in E. abbreviana; and apex of socius pointed in lateral view, bluntly convex in E. abbreviana. The females of these two species are indistinguishable. All other species mentioned above have a long, fingerlike uncus with a distinctly bifid apex, and valvae with the ventral margin deeply and broadly emarginate at one- third the length from base (Fig. 4). Epinotia abbreviana is distinguished by the following characters: 1) costal fold of forewing absent, present in others; 2) uncus at least as long as aedeagus, shorter in others; and 3) apex of socius bluntly convex in lateral view, pointed in others. The female is characterized by the distinct, dorsoanteriorly arched papillae anales, which are fringed laterally with large, slender, hooked setae; by the lamella postvaginalis with an acute dorsolateral angle on each side; and by a pair of large, horn-shaped signa.
Material studied. CANADA: Newfoundland: St. John’s, em. 18, 21.VI.1981, elm, (FIDS), 2 6; Bowring Park, St. John’s, em. 6-7. VII.1988, Ulmus rubra, (FIDS), 2 9, all in CNC. ENGLAND: Abingdon, 30.V1.1924, (H. C. Hayward), 1 6, in USNM. GERMANY: Nieder- Weser, Bremen-Stadtwald, 5.VII.1941, Ulmus, (E. Jackh), 1 2, in USNM.
Distribution and biology. Epinotia abbreviana is widespread and distributed throughout Europe and Asia Minor. Early larval instars of this species feed inside the developing bud of species of Ulmus. A characteristic ring of small perforations appears on the leaf surface when the leaf becomes fully expanded in the spring; Bradley et al. (1979) provided excellent illustrations of leaves of Ulmus damaged by the larvae. Later larval instars become leaf tiers. Epinotia abbreviana is a potential pest of Ulmus species in Canada and the United States.
ACKNOWLEDGMENTS
I thank Kevin Pardy of the Newfoundland and Labrador Region, Forestry Canada for providing collection information and specimens of E. abbreviana collected in St. John’s, and R. W. Hodges of the United States National Museum of Natural History, Washitetey D.C., for the loan of specimens of E. abbreviana from Europe.
LITERATURE CITED
BRADLEY, J. D., W. G. TREMEWAN & A. SMITH. 1979. British Tortricoid moths. Tor- tricidae: Olethreutinae. The Ray Society, London. 320 pp.
VOLUME 47, NUMBER 1 59
BENANDER, P. 1950. Fiarilar. Lepidoptera IJ. Smafjarilar. Microlepidoptera. Andra familjegruppen Vecklarefjarilar. Tortricina. Svensk Insektfauna 10. 173 pp., 9 plates. GRAAF BENTINCK, G. A. & A. DIAKONOFF. 1968. De Nederlandse Bladrollers (Tortrici- dae). Een Geillustreerd Overzicht. Monogr. Nederland. Entomol. Vereen. 3:1-201.
HANNEMANN, H. J. 1961. Kleinschmetterlinge oder Microlepidoptera I. Die Wickler (s. str.) (Tortricidae). Die Tierwelt Deutsch. 48:1—233.
KELLY, K. L. & D. B. Jupp. 1955. The ISCC-NBS method of designing colors and a dictionary of color names. U.S. Department of Commerce, National Bureau of Stan- dards, Washington, D.C. Circular 553. 158 pp.; and ISCC-NBS color-name charts illustrated with centroid colors, a supplement to circular 553.
KENNEL, J. V. 1921. Die Palaearktischen Tortriciden. Eine monographische Darstellung. Zoologica 54:1-742, 24 plates.
KUZNETSOV, V. I. 1966. New species of leaf-rollers (Lepidoptera, Tortricidae) from south of the Primorye Territory. Trudy Zool. Inst. Leningrad 37:177-207. [In Russian. ]
KUZNETSOV, V. I. 1978. Family Tortricidae (Olethreutidae, Cochylidae)—Tortricid moths, pp. 193-686. In Medvedev, G. S. (ed.), Keys to the insects of the European part of the USSR. Vol. IV. Lepidoptera Part 1. Opred. Fauna SSSR 117:1-686. [Translated from Russian]. Amerind Publ. Co. Pvt., New Delhi. 1987. 991 pp.
PIERCE, F. N. & J. W. METCALFE. 1922. The genitalia of the group Tortricidae of the Lepidoptera of the British Islands. Oundle, Northants. 101 pp., 34 plates.
Received for publication 20 June 1992; revised and accepted 26 September 1992.
Journal of the Lepidopterists’ Society 47(1), 1998, 60-77
A REVISION OF THE SPECIES OF NEMATOCAMPA (GEOMETRIDAE: ENNOMINAE) OCCURRING IN THE UNITED STATES AND CANADA
DOUGLAS C. FERGUSON
Systematic Entomology Laboratory, Agricultural Research Service, USDA, % U.S. National Museum of Natural History, Washington, D.C. 20560
ABSTRACT. The three nearctic species of Nematocampa are revised, with special emphasis on the biology and complex nomenclatural history of the type species, N. resistaria (Herrich-Schaffer, 1855). Of the two names most commonly applied to this species, limbata Haworth, 1809, is a primary homonym, and filamentaria Guenée, 1857, is a junior synonym of resistaria. Nematocampa resistaria is transcontinental; N. breh- meata (Grossbeck, 1907) is limited to California; N. baggettaria, new species, is south- eastern; and N. expunctaria Grote, 1872, is synonymized under N. resistaria. The well- known larva of N. resistaria feeds on plants of at least 20 families, but the larvae of the others are unknown. About 20 additional species occur in the neotropics.
Additional key words: taxonomy, nomenclature, host plants.
Nematocampa Guenée is a New World genus of at least 23 species, a few of which apparently are undescribed. Although concentrated in the tropics and occurring southward to Argentina, three species are present in the temperate zone of the United States, and one reaches Canada. The neotropical species are diverse and may not all be con- generic. A few other neotropical Ennominae, perhaps most notably Melinodes conspicua Schaus from Brasil, have a reticulate wing pattern suggestive of that of Nematocampa but are not closely related.
Although the type species, N. resistaria (Herrich-Schaffer), has a large literature that goes back to 1809, was found and drawn by John Abbot perhaps even earlier (Abbot drawing copied by Guenée 1857:9, p. xlvi; 1858:pl. 2, fig. 3), and is widely known because of its distinctive appearance and unusual, filament-bearing larva, its nomenclature and taxonomy have not been interpreted correctly. It was described under eight names, two of which, N. limbata (Haworth) and N. filamentaria Guenée, have competed incorrectly for priority in all of the more recent literature. Despite that long history, another very distinct species of the southeastern U.S. remained undiscovered until the 1980’s and is de- scribed in this paper.
Originally I intended only to describe this new species, Nematocampa baggettaria, and perhaps verify the correct name for the type species. However, the paper assumed the proportions of a revision as more material and more literature were examined, and the full complexity of the problem unfolded. Every name referring to Nematocampa spe- cies in the fauna of America north of Mexico is here applied differently except that of N. brehmeata.
VOLUME 47, NUMBER 1 6l
Nematocampa Guenée
Nematocampa Guenée, 1857, in Boisduval and Guenée, Hist. Nat. des Insectes, Species général des Lépidoptéres 9:120. Type species: Nematocampa filamentaria Guenée, 1857, ibidem 9:121; 1858, ibidem Atlas:pl. 2, fig. 3; pl. 5, fig. 1, by monotypy. Nematocampa filamentaria Guenée is herein regarded as a junior subjective synonym of Microgonia resistaria Herrich-Schaffer, 1855, Sammlung Aussereuropdischer Schmetterlinge, p. 41, pl. [65], fig. 368.
Species of this genus may nearly always be recognized by the char- acteristic wing pattern. The dark medial line of the forewing is unusu- ally far out, partly touching or confluent with the postmedial line (Figs. 1-18), or missing (Figs. 19-23); in N. resistaria and brehmeata these lines touch near the inner margin and at vein M,, thereby making a closed cell between them near the middle of the wing. Neotropical species, however, do not have a well-developed closed cell, or it may not show in poorly marked specimens. The outer third of both wings has conspicuous areas of brown or purplish-brown shading in most species. The pale ground color, varying from whitish to deep orange yellow, is often striated transversely with multiple, fine, short streaks, and marked longitudinally with fine dark lines on the veins, giving a reticulated effect. Length of forewing: 6¢, 7-14 mm; 22, 7-16 mm.
Other features of Nematocampa are as follows: male antenna pris- matic, compressed, finely setose; female antenna filiform, finely setose; chaetosema small, with 5-6 bristles; front smooth; palpus short, only slightly surpassing front; male hindtibia with inner preapical spur cu- riously modified in most species (Figs. 24-26), elongated almost to end of tibia and claviform, enlarged distally to 2-4 times thickness of normal, linear spurs.
Male genitalia (Figs. 28-33) with end of gnathos laterally compressed in typical group, dorsoventrally compressed in some neotropical species; juxta large and elongated dorsally (toward uncus), where it becomes notched or bifurcated in all North American species and some neo- tropical ones; valve in all North American and some neotropical species divided into a long costal lobe and shorter, rounded, saccular lobe, as in Ennominae of the tribe Semiothisini; other neotropical species, pre- sumed to be more primitive ones in this respect, have valve undivided; most neotropical species, those with undivided valve, have a pair of short spinose processes (resembling a furca) arising from juxta, one on each side; in those with a bilobed valve, the spinose processes have degenerated to a pair of simple sclerites flanking juxta and apparently forming narrow bridges between juxta and transtilla. A long, hairy corema (Fig. 28) arises laterally from near the base of each valve in most species, but may be reduced or vestigial (Fig. 30). Female genitalia typically with a longitudinally ovate signum, with an indeterminate number of dentate processes radiating from its margin.
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Fics. 1-9. Nematocampa resistaria. 1, 6, Armdale, Halifax Co., Nova Scotia, 12 Aug. 1948. 2, 6, Seton Cr., Lillooet, British Columbia, 4 July 1991. 3, 6, Beaverton, 9 mi W of Portland, Oregon, 14 June 1963, C.W. Nelson. 4, 6, Oak Zone, 5 mi SW of Midway, Wasatch Co., Utah, 29 July 1971. 5, 6, Sycamore Landing, nr. Seneca, Montgomery Co., Maryland, 28 May 1977. 6, 6, McClellanville, Charleston Co., South Carolina, 20 May 1974, R. B. Dominick. 7, 4, Eagle L., Colorado Co., Texas, 27 April 1978, A. & M. E. Blanchard. 8, 2°, Baddeck Bridge, Victoria Co., Nova Scotia, 29 July 1970. 9, 2°, Sycamore Landing, Seneca, Montgomery Co., Maryland, 24 July 1976. Magnification: 2x.
Remarks. Nematocampa in the broad sense has three main species groups. Group 1—those with or without a modified hindtibial spur, but always with a bilobed valve, degenerate furca, strongly bifurcate juxta, and one cornutus; group | includes the three North American species treated in this revision and two closely related neotropical species
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Fics. 10-18. Nematocampa species. 10, N. resistaria °, Oak Zone, 5 mi SW of Midway, Wasatch Co., Utah, 29 July 1971. 11, N. resistaria 2 (white ground color), nr. Elsie, Clatsop Co., Oregon, 2 Sept. 1968, S. G. Jewett. 12, N. resistaria 2 (yellow ground color), same locality, 6 Sept. 1968, E. L. Griepentrog. 13, N. resistaria 2, Withlacoochee State Forest, Hernando Co., Florida, reared from larva on Myrica 7 May 1988, H. D. Baggett. 14, N. brehmeata 4, Mt. Shasta (city), Siskiyou Co., California, 1 Aug. 1990. 15, N. brehmeata 8, Anderson Springs, Lake Co., California, 3 July 1949, W. R. Bauer. 16, N. brehmeata 4, same data as for Fig. 15. 17, N. brehmeata 2, same data as for Fig. 15 but collected 26 July 1952. 18, N. brehmeata 2, San Antonio Cr., Sonoma Co., California, 21 July 1939, W. R. Bauer. Magnification: 2x.
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Fics. 19-23. N. baggettaria. 19, Holotype 6. 20, 6, 8 mi N of Sumatra, Apalachicola Natl. Forest, Liberty Co., Florida, 2 June 1990, H. D. Baggett. 21, 6, 4.2 mi NE of Abita Springs, St. Tammany Parish, Louisiana, 25 April 1984 (spring brood), V. A. Brou. 22, 2, Torreya State Park, Liberty Co., Florida, 19 August 1982, H. D. Baggett. 23, 2, Same data as for Fig. 21 but collected 14 April 1984 (spring brood). All illustrated specimens are in the collection of the USNM, and all were collected by the author unless otherwise indicated. Magnification: 2x.
that I could identify, namely N. evanidaria Schaus and N. arenosa Butler (plus other neotropical species apparently undescribed). Group 2—the remaining species of similar appearance and with similarly modified hindtibial spur, but with different male genitalia, as follows: valve entire, not two-lobed; juxta not or hardly bifurcate; furca fully developed; and aedeagus with one or two cornuti (N. completa Warren, N. angulifera Oberthir, N. reticulata Butler, N. decolorata Warren, and probably others). Group 3—a group of three or four species of different appearance, in part with a narrow, dark, outer marginal border only, without a modified hindtibial spur, with simpler male genitalia and reduced juxta, no remnants of a furca, numerous cornuti in the vesica, and many other differences (e.g., some with bipectinate male antennae). This group includes N. falsa Warren, N. confusa Warren, and possibly N. benescripta Warren and N. interrupta Warren, al- though the last two seem different again. There is little to tie group 3 to Nematocampa, and these species will almost certainly be removed to other genera after further study. Certain species of other ennomine groups, most notably Melinodes conspicua Schaus from Brasil, may have a pattern suggestive of Nematocampa but are probably unrelated.
When I prepared the Check List of North American Geometridae (Ferguson 1983), I left Nematocampa at the end of the Ennominae
65
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Fics. 24-27. Male right hindtibiae of Nematocampa species. 24, N. resistaria, Oke- fenokee Swamp, Georgia. 25, N. resistaria, near Elsie, Clatsop Co., Oregon. 26, N. brehmeata, Anderson Springs, Lake Co., California. 27, N. baggettaria. 4.2 mi NE of Abita Springs, St. Tammany Parish, Louisiana.
because, like other authors, I had not determined where it belongs. This has not changed because it would require comprehensive revisionary study of the neotropical Ennominae to determine the phylogenetic relationships of Nematocampa.
Key to Species of Nematocampa of the United States and Canada
Male with hindtibia enlarged and inner preapical spur curiously modified, long, clavate (Figs. 24-26); ground color of wings whitish or yellow; wing length generally more than 10 mm; widely distributed eee 2
— Male with hindtibia not enlarged and preapical spurs unmodified (Fig. 27); ground
color of wings orange brown to yellowish; moths small, wing length less than HOt. southeastern United States’. baggettaria 2. Females with ground color white or nearly so; males with clavate hindtibial spur (Figs. 24, 25) longer than first tarsal segment, conspicuously swollen to 3-4 times thickness of other preapical spur; transcontinental, widespread ..................... resistaria — Females with ground color yellow (a few Oregon females of resistaria with yellow
—
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Fics. 28-33. Male genitalia. 28, N. resistaria, McMinnville, Oregon. 29, Aedeagus of same specimen. 30, N. baggettaria, Jefferson Co., Florida. 31, Aedeagus of same specimen. 32, N. brehmeata, Anderson Springs, Lake Co., California. 33, Aedeagus of same specimen.
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ground color will key here); males with clavate hindtibial spur (Fig. 26) swollen only to twice thickness of other preapical spur; California ww. brehmeata
Nematocampa resistaria (Herrich-Schaffer), revised status
(Figs. 1-18, 24, 25, 28, 29, 34)
Phalaena limbata Haworth, 1809, Lepidoptera Brittanica, Pt. 2:346. (Publication dates for the four parts of the Lepidoptera Britannica were established by Griffin, 1932). HOMONYM
NOTE: Phalaena limbata Haworth is a junior primary homonym of Phalaena limbata Linnaeus, 1767, Systema Naturae, ed. 12, p. 873, which is the European species now known as Evergestis limbata (L.) (Crambidae). According to the International Code of Zoological Nomenclature (1985, Art. 59a), “a species-group name that is a junior primary homonym must be permanently rejected.” It is unlikely that the name limbata could be conserved because it has not been in continuous use for the past 50 years. Some authors have used the name filamentaria Guenée for this species.
Macaria limbata, Stephens 1829(2):155. Wood 1839, 1854:116, pl. 26, fig. 748n.
Ania limbata, Stephens [1831]:322. Hulst 1896:373. Dyar 1902 [1903]:338.
Nematocampa limbata, Barnes and McDunnough 1917:121. McDunnough 1938:169. Ferguson 1983:98.
Type locality: England (“Angliae rarissime’’). Presumed to be a false type locality.
Microgonia resistaria Herrich-Schaffer, 1855, Sammlung Aussereuropaischer Schmetter- linge, p. 41, pl. [65], fig. 368.
Nematocampa resistaria, Walker 1860:147.
Type locality: not given.
Microgonia vestitaria Herrich-Schaffer, 1855, ibidem, pp. 63, 82, pl. [65], fig. 368. Type locality: Brasil (considered to be an error; see below).
NOTE: Herrich-Schaffer proposed different species names, resistaria and vestitaria, on different pages of the same work in reference to the same figure. The names were published simultaneously, but resistaria has page priority. This synonymy was rec- ognized by Guenée (1857, pt. 9:121), who applied the name resistaria and listed vestitaria as a junior synonym. Walker (1860:147) did likewise, but Guenée qualifies as the first reviser. Packard (1876:471) synonymized both under N. filamentaria Guenée but attributed resistaria to Walker, 1860, overlooking Herrich-Schaffer’s prior publication. Although Guenée (1857), oblivious in this instance to the principle of priority, used the older name resistaria for what he thought was a South American variety of his new North American species, N. filamentaria, both Walker (1860) and Packard (1876) considered resistaria to be the North American species. The names resistaria and vestitaria of Herrich-Schaffer, filamentaria Guenée, and limbata Haworth continued to be treated as synonyms by most subsequent authors. Although the only type locality mentioned by Herrich-Schaffer is Brasil (for vesti- taria), his figure 368 clearly represents a male of the widespread North American species. I found no neotropical species or specimen that agrees with the figure and consider the locality to be an error. A similar error may be seen in figures 373 and 374 on the same plate (pl. [65]). The type locality for Gnophos armataria Herrich- Schaffer is given as Venezuela, although the specimens depicted are of the North American species treated in more recent literature as Priocycla or Cepphis armataria (Herrich-Schaffer).
Nematocampa filamentaria Guenée, 1857, Hist. Nat. des Insectes, Species Général des Leépidopteéres 9:121; pl. 5, fig. 1; pl. 2, fig. 3 (larva). Packard 1876:471, pl. 11, fig. 46; pl. 18, figs. 8, 8a. Capps 1943:147. Forbes 1948:110. Ferguson 1954:321.
Type locality: Canada, by present lectotype designation.
NOTE: This species was described from one male and three female specimens from “Amérique septentrionale.” The male and one female are in the United States Na- tional Museum (USNM); but the location of the second female, the syntype illustrated by Guenée (1857:pl. 5, fig. 1), has not been determined. Although this syntype appears to belong to the genus Nematocampa, it does not agree with any known species.
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Ordinarily, one would choose the illustrated syntype as the lectotype, but because the source and identity of the specimen shown by Guenée are in doubt, I hereby designate as lectotype the male in the USNM, which bears the word “Canada” on a small, round label, as well as the usual Guenée and Oberthtir labels and a Guenée type label. Although the right wings and abdomen are lost, the specimen clearly represents resistaria; all the more so because it is from a region where only that species is known to occur. The specimen reached the USNM with other Guenée types through the Oberthiir and Barnes collections.
Nematocampa expunctaria Grote, 1872, Canad. Entomol. 4:101. Capps 1943:147. Fer- guson 1983:98. REVISED SYNONYMY
Nematocampa limbata torm expunctaria, McDunnough 1938:169. Type locality: Alabama. [Holotype in Academy of Natural Sciences of Philadelphia. ]
Ania limbaria var. chagnoni Swett, 1913, Canad. Entomol. 45:76. Type locality: Isle Ste. Therese, St. Johns Co., Quebec. [Holotype in Museum of Comparative Zoology, Harvard University (MCZ).]
NOTE: The name was based on one male of a melanic form of N. resistaria.
Nematocampa limbata orfordensis Cassino and Swett, 1922, The Lepidopterist 3:156. McDunnough 1938:169. Ferguson 1983:98. REVISED SYNONYMY Type locality: Port Orford, Oregon. [Holotype male in MCZ.]
Eugonobapta brunneolineata Hulst, 1900, J. New York Entomol. Soc. 11:218. REVISED SYNONYMY
Ellopia brunneolineata, McDunnough 1938: 171
Nematocampa brunneolineata, Ferguson 1983:98 (as synonym of expunctaria) Type locality: Hastings, Florida. [Holotype in American Museum of Natural History, New York.]
Diagnosis. This widespread nearctic species is distinguished by the pale whitish ground color of nearly all females (with rare exceptions) as compared to the more yellowish males; the swollen male hindtibia and the large size of the modified male hindtibial spur (Figs. 24, 25), which is swollen to three times the thickness of the other spurs; the combination of long, nearly straight prongs on the bifurcate juxta and the large coremata of the male genitalia (Fig. 28); and the combination of large, spiny signum but otherwise unsclerotized bursa copulatrix in the female genitalia (Fig. 34).
Further description. Both sexes with scales of front, antennae, palpi, and body light brown, legs and vertex somewhat paler.
Male (Figs. 1-18): Outer margin of forewing variable from rounded to angulate, hindwing usually rounded; ground color pale to deep yellow, with brown or purplish- brown markings well developed (Fig. 2) to obsolescent (Fig. 6) and geographically variable. Hindtibia much enlarged, apparently containing a large hair pencil that seems never to be fully extruded in museum specimens; hindtibia also with curiously modified, clavate, inner preapical spur twice as long as, and distally widened to at least three times as wide as outer preapical spur; modified spur longer than metatarsus. Female (Figs. 8-13): Outer margins of both wings angulate, that of forewing quite strongly so and concave between terminus of M, and apex. Ground color whitish or cream colored, with reddish-brown markings much less variable geographically than those of male (variation discussed below).
Undersides of both sexes paler and and with markings less distinct than above. The reticulate nature of the wing pattern is more emphasized in females than in males because the longitudinal veins are more strongly outlined. Length of forewing: 46, 10-14 mm; 99, 12-14 mm (reared specimens may be smaller). .
Male genitalia (Figs. 28, 29). Similar to those of N. brehmeata but with saccular lobe of valve more narrowed distally, prong of bifurcate juxta nearly straight and somewhat divergent, not incurved as in brehmeata, and with gnathos more produced distally; differ
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39
36
Fics. 34-36. Female genitalia. 34, N. resistaria, Okefenokee Swamp, Georgia. 35, N. brehmeata, Anderson Springs, Lake Co., California. 36, N. baggettaria, no locality (old specimen without data in U.S. National Museum).
obviously from those of N. baggettaria in having large coremata, a longer, slightly curved saccular (ventral) lobe on valve, and prongs on bifurcate juxta that are shorter than juxtal neck or base from which they arise.
Female genitalia (Fig. 34). Characterized by combination of large stellate signum, about three times wider than narrowest constriction of ductus bursae and often larger than that of brehmeata (specimen illustrated is an exception); an ovoid corpus bursae, twice as long as wide and with little sclerotization of its surface; and an ostial funnel whose widest dimension is considerably less than width of seventh segment at that point. By comparison, N. brehmeata (Fig. 35) has a wider ostial funnel, nearly as wide as the segment; more sclerotization on corpus bursae between signum and ductus bursae; and smaller signum only twice as wide as ductus bursae. Nematocampa baggettaria (Fig. 36) has an almost globular corpus bursae, and small, differently shaped signum hardly wider than narrowest constriction of ductus bursae.
Early stages. The peculiar larva of N. resistaria, which has been given the common name of “the filament bearer,” was described and illustrated many times (e.g., Guenée 1857:pl. 2, fig. 3; Packard 1876:pl. 13, fig. 8; Peterson 1948:L56F; Furniss & Carolin 1977: fig. 122; Stehr 1987:504, fig. 26.281; Ives & Wong 1988:20, fig. G, p. 142, fig. E). Guenée’s illustration was copied from a John Abbot drawing, probably of a specimen from Georgia; Packard’s was a drawing of a larva from Salem, Mass.; Furniss and Carolin’s was from the western U.S.; and Ives and Wong’s were from the prairie provinces of Canada. These illustrations show considerable variation. Mosher (1917:41-—48, fig. 2E, H) concentrated mainly on the pupa and illustrated it, but she also briefly described the larva.
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The following is a description of the mature larva based mainly on photographs from T. L. McCabe of larvae from Pinebush, Albany County, New York, reared on Quercus ilicifolia (Fagaceae). Body cylindrical, of medium to somewhat stout form, with two unequal pairs of slender, tapered, fleshy, dorsal filaments arising from near the posterior margins of segments A2 and A3, the anterior pair being the longer and, when extended, nearly equal to the combined lengths of the first four body segments. Filaments somewhat extensile, straight when extended, bent and curled at the tips when retracted; pale basally, shading to dark purplish brown distally, but conspicuously white tipped, and with their surfaces appearing minutely tuberculate or pubescent. Correspondingly situ- ated, brown, knoblike prominences on Al, each with a strong apical seta, and between these and the filaments of A2, two pairs of smaller, wartlike humps on A2, the anterior pair whitish, the following pair smaller and brown. Segment A8 somewhat elevated and pointed dorsally, after which the dorsum slopes downward abruptly toward the posterior edge of the anal plate. Head light rust red, marbled with pale tan to pinkish brown. Body variegated light and dark brown, the pale ground color often with a pinkish or purplish tint, although thoracic segments are dark brown and A1-2 partly yellowish brown. Dorsum from first pair of filaments to anal plate mostly dark brown, flanked by a gray to brown subdorsal stripe. Superimposed upon this are two or three dark-brown, oblique, abdominal bands that arise in the dorsal region and incline gradually downward and forward to just above lateral fold where they terminate in vague, elongate, diamond-shaped or fusiform, brown patches. Venter with a confused pattern of mainly rust-brown shading on a paler ground. Thoracic legs dark brown; prolegs mottled brown with whitish anterior edging on anterior proleg continuous with whitish posterior border of anal plate in one larva but not the other.
Ives and Wong (1988) published two colored photographs, one showing the filaments curled and the other showing them straight. Their illustrations also show a variable and complex pattern involving an irregular, continuous whitish lateral stripe on the head and first two thoracic segments, and oblique whitish lateral stripes on segments A6 to A8, the last running down the lateral side of the proleg. These features vary somewhat from those of the New York larvae described above. Nematocampa resistaria feeds on plants of at least 20 families. Prentice et al. (1963:489) listed the following as hosts (number of collections in parentheses): Pseudotsuga menziesii (Mirb.) Franco. (242); Tsuga hetero- phylla (Rafn.) Sarg. (220); Abies balsamea (L.) Miller (Pinaceae) (66); Salix spp. (Sali- caceae) (20); Picea glauca (Moench) Voss (Pinaceae) (18); Betula papyrifera Marsh. (Betulaceae) (14); Larix laricina (DuRoi) K. Koch (Pinaceae) (12); Thuja plicata D. Don (Cupressaceae) (12); Picea engelmannii Parry (Pinaceae) (11); Ulmus americana L. (Ulmaceae) (8); Larix occidentalis Nutt. (6); Pinus monticola Doug]. (Pinaceae) (5); Fraxinus americana L. (Oleaceae) (5); and Acer rubrum L. (Aceraceae) (5); Tilia amer- icana L. (Tiliaceae) (4); Abies lasiocarpa (Hook.) Nutt. (Pinaceae) (3); Ostrya virginiana (Mill.) K. Koch (Betulaceae) (2); Acer negundo L. (Aceraceae) (2); Abies grandis (Doug. Lind.) (Pinaceae) (2); and Alnus rubra Bong. (Betulaceae) (2); Prentice et al. list 10 other host plants with only one collection for each, but only two additional plant families are represented, Rosaceae (Prunus virginiana L.) and Fagaceae (Quercus macrocarpa Michx. ). The above records probably are biased in a way that reflects more thorough sampling of economically important trees and little attention to shrubs.
The compilation of host information by Tietz (1972) gives references to 25 food plants, of which the following are plants of genera not included above: Aesculus hippocastanum L. (Hippocastanaceae); Carya sp. (Juglandaceae); Castanea dentata (Marsh.) Borkh. (Fa- gaceae); Corylus sp. (Betulaceae); Crataegus sp.; Pyrus ioensis (Wood) Bailey; Pyrus malus L.; Rosa sp.; Rubus allegheniensis Porter; Rubus idaeus L. var. strigosus (Michx.) Maxim.; Fragaria chiloensis (L.) (all Rosaceae); Humulus lupulus L. (Moraceae); Ribes americanum Mill.; Ribes lacustre (Pers.) Poir.; Ribes sativum Syme (Saxifragaceae); Robinia Pseudo-Acacia L. (Fabaceae); and Sedum sp. (Crassulaceae). It was also reared from Abies lasiocarpa (Hook.) Nutt. (Pinaceae) at Bonner, Montana (U.S. Forest Service); Liquidambar styraciflua L. (Hamamelidaceae) (Kimball, 1965:186); Myrica cerifera L. (Myricaceae) (by D. Baggett, in USNM); Myrica gale L. (USNM); “sweet fern” [Myrica asplenifolia L.] (Mosher, 1917:48); Quercus ilicifolia Wangenh. (Fagaceae) (T. L. McCabe
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pers. comm.); Vaccinium arboreum Marsh. (Ericaceae) (T. S. Dickel pers. comm.); Ce- anothus velutinus (Rhamnaceae) in Idaho (U.S. Forest Service); Amelanchier and Cra- taegus spp. (Rosaceae) (L. R. Rupert; pers. comm. reported by the author, 1954:321); cherry (USNM); and Gleditsia triacanthos L. (Fabaceae) (reared by the author).
Distribution. Nematocampa resistaria occurs across southern Canada from Nova Scotia to Vancouver Island, B.C., southward at least to Hernando Co., Florida, Mississippi, southern Louisiana, Harris, Jackson, Colorado, and Blanco counties, Texas, and in the West at least to Morgan Co., Colorado, Sanpete and Wasatch counties, Utah, and Josephine Co., Oregon. In mountainous or semi-arid regions it is a species of low elevations and riparian habitats. It is replaced in California by N. brehmeata.
Flight period. The species is single brooded in Canada and most of the northern U.S., flying mainly in July and August. It becomes double brooded in the middle states, emerging in May and again in late July (e.g., in Maryland), but it seems to be no more than double brooded in the South. Thirty specimens in the USNM from South Carolina, Georgia, northern Florida, and Texas were all collected in April and May. In Louisiana and Mississippi it was collected in April, May, June, and July, based on many records from year-round light-trapping (V. A. Brou and B. Mather collections). The flight period in the Northwest is also mainly in July and August, rarely in June, but with a few September and early October records for western Oregon.
Geographical variation. Nematocampa resistaria has two main geographical variants that were named as species or subspecies: N. expunctaria (=brunneolineata) in the Southeast, and subspecies orfordensis in the Pacific Northwest. Grote (1872, 1882), Capps (1943:147), and the present author (1983:98) regarded expunctaria as a distinct species in which the wing pattern of the male is reduced to little more than the antemedial and postmedial lines (Figs. 6, 7). Females remain unchanged. Capps, knowing only the type of expunctaria, further described what he thought were differences in the genitalia. A reevaluation of more and better material now indicates that expunctaria is the same species as resistaria, at best a weak subspecies occupying a narrow coastal zone from South Carolina to northern Florida and westward through southern Louisiana to Texas. Intermediate forms of every degree may be found. Where the range of the species continues westward into eastern Texas, the moths revert to the more normal, well-marked form of resistaria. The differences in genitalia mentioned by Capps prove to be of no consequence when more than one specimen is examined.
The form described as orfordensis is localized and possibly related to the cool maritime climate of coastal Oregon, Washington, and Vancouver Island. It is as large as N. breh- meata (wing length: 64, 13-14 mm; ¢¢, 15-16 mm) and sometimes, like that species, a deep shade of yellow. A size gradient is apparent between populations of the coastal region and the interior, those from eastern Oregon and Washington hardly differing from eastern specimens. Again, it is a poorly defined subspecies. However, one unusual feature of orfordensis is the occasional occurrence of yellow females colored like males (Fig. 12). Three of these in the USNM were collected near Elsie, Clatsop Co., Oregon on 5, 6 September 1963, 1968, and 1969. It is not a seasonal form because normal females with whitish ground color were collected with them. These are the only yellow females of N. resistaria that I have seen, although N. brehmeata always has yellow females.
Material examined. 244 adults, 7 genitalia slides.
Nematocampa brehmeata (Grossbeck) (Figs. 14-18, 26, 32, 38, 35) Ania brehmeata Grossbeck, 1907, Trans. Amer. Entomol. Soc. 33:343.
Nematocampa brehmeata, Barnes and McDunnough 1917:121. McDunnough 1938:169. Ferguson 1983:98.
Type locality: Cazadero, Sonoma County, California. [Holotype in AMNH.]
Diagnosis. This is a Californian species in which both sexes are yellow, showing somewhat of a reversal of the sexual dimorphism of resistaria
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because the females are often a deeper yellow than the males. The dark markings of the outer third of both wings in males are more broken up by encroachments of pale ground color and in females are lacking entirely. These yellow females, without dark submarginal markings, are very different in appearance from most females of resistaria. Di- agnostic differences in the hindtibial spurs were noted in the key. In the male genitalia, the prongs of the bifurcate juxta are incurved distally, not straight. This species occurs in northern and central California west of the Sierra Nevada. It appears to be the only species of Nematocampa in that area, although resistaria occurs in nearby Oregon.
Further description. Body, head, antennae, and legs similar to those of resistaria except that modified hindtibial spur of male is more slender, distally swollen to not more than twice thickness of the other preapical spur, and slightly shorter than metatarsus. Wing shape and pattern similar to those of resistaria except that the dark, purplish-brown shading in outer third of wings in males tends to appear reduced, narrowed, or otherwise broken up, usually faded, and further reduced or lost on the hindwing. Dark shading of outer third of wings lost entirely in the four females examined. Ground color varies from light yellow to intense orange yellow in both sexes, but males are mostly light (9 of 11), and females may more often be deep yellow (2 of 4). Length of forewing: 66, 12-14 mm; 99, 138-14 mm.
Male genitalia (Figs. 32, 33). Differ from those of resistaria in slightly wider saccular lobe of valve; shorter coremata, only about half as long; narrower overall shape of juxta, with its two bowed prongs diverging basally but curving toward each other distally; and two free sclerites laterad of base of juxtal prongs thicker, and also bowed, not straight like those of resistaria.
Female genitalia (Fig. 35). Ostial funnel very large, almost as wide across opening as width of seventh segment at that point; about twice as large as ostial funnel of resistaria. Bursa copulatrix quite heavily sclerotized in zone between signum and ductus bursae; integument in this area pleated or rugose in both species, but not sclerotized in resistaria. Ovipositor wider and less elongated than that of resistaria, the lobes (papillae anales) being shorter than ostial funnel is wide (longer than width of funnel in resistaria).
Early stages. Unknown.
Distribution. I saw specimens from the following localities in California: Santa Cruz, Santa Cruz Co.; Los Gatos, Santa Clara Co.; Lucas Valley, Marin Co.; San Antonio Creek, Sonoma Co.; Napa, Napa Co.; Lucerne and Anderson Springs, Cobb. Mt., Lake Co.; Michigan Bluff, Placer Co.; Nelson Creek and Mohawk, Plumas Co.; Laytonville, Men- docino Co.; Oroville, Butte Co.; Hat Creek, Shasta Co.; Mt. Shasta (city), Siskiyou Co., and Gasquet, Del Norte Co. I collected three males of this species in a riparian habitat on the outskirts of the town of Mt. Shasta, on the road to Lake Shasta. The site was in a moist stream bottom with abundant willows, dogwood, and alder, mixed conifers nearby, and a pond fringed with Typha marsh.
Flight period. 20 June—27 August.
Geographical variation. Compared to those from elsewhere, three males from Mt. Shasta are smaller (wing length: 12-18 mm), slightly paler yellow, and with all dark markings intensified, although specimens from Shasta and Del Norte counties are the usual orange-yellow and more nearly normal in size and markings. The Mt. Shasta specimens could be mistaken for N. resistaria, but in pattern and structure they are brehmeata.
Material examined. Thirty-five adults, 5 genitalia slides.
Remarks. The distinction between N. brehmeata and resistaria may not always be as clear as I have indicated, because a few specimens that appear intermediate have been taken near the contact zone. Two males from Mt. Shasta (Fig. 14) almost have the markings of resistaria, and the male from McMinnville, Oregon, whose genitalia are shown in Fig.
VOLUME 47, NUMBER 1 73
28, resembles a deep-yellow male of brehmeata, although its juxta is clearly that of resistaria. Also, the only truly yellow females of resistaria (Fig. 12) are from Oregon, but they are otherwise consistent with resistaria, not brehmeata.
Nematocampa baggettaria Ferguson, n. sp.
(Figs. 19-28, 27, 30, 31, 36, 37-39)
Diagnosis. This is the smallest and most distinct of the nearctic species, with the wing margins rounded, not angulate, wings mostly orange brown but with dark, purplish-brown shading in most of the outer third in females. The pattern is simplified, with two regular lines on the forewing, one on the hindwing, and almost no reticulation of lighter areas in either sex. It is the only species of Nematocampa in the U.S. without a swollen, clavate, hindtibial spur. The species is known from northern Florida and the central Gulf Coast region, and there is one record from Lumberton, North Carolina.
Further description. Head, antennae, and legs similar to those of resistaria except that the male hindtibia is not swollen and does not bear a specialized, large, clavate spur; all spurs are small and normal. Front in both sexes light yellowish brown, variably sprinkled with bright red-brown scales or with a diffuse red-brown border on each side. Male (Figs. 19-21): outer margin of wings rounded like those of southern resistaria or more so, and with apex less acute; wings almost uniformly ochreous orange brown, or variably tinged with purplish in outer third; forewing with antemedial a thin, dark, regular, convex line; postmedial line similar, curved subparallel to outer margin or nearly straight; hindwing with slightly curved postmedial line bisecting wing nearly in middle; small, rounded, dark discal dot on each wing; fringes dusky; wings faintly dusted with a few dark scales but without reticulation or any sign of a median band in median space. Underside darker, with markings reduced, although postmedial of hindwing may be closely preceded for most of its length by a faint, thinner, subparallel line. Spring brood (April) specimens with tendency to be slightly larger and paler than summer (June-August) specimens. Length of forewing: holotype, 8 mm; other 66, 7-8 mm. Female (Figs. 22, 23): Apex of forewing somewhat produced but with outer margins rounded like those of male, unlike females of other species. Ochreous orange-brown ground color of wings dusted with reddish-brown scales that outline veins of median space in some specimens, giving a suggestion of reticulate pattern seen in pale wing areas of other species. Forewing with antemedial line thicker than that of male and purplish; postmedial line curved or nearly straight, blackish, commonly indented at cubital fold; postmedial of hindwing slightly curved to nearly straight, bisecting wing near middle. Discal spots small but prominent. Outer third of both wings dark purplish brown except for orange-yellow patch toward apex of forewing and variable indications of same color in form of a diffuse, mesial, transverse band in outer third of hindwing; fringes concolorous. Underside similarly marked but with lines thickened and diffuse, and all paler areas more or less suffused with gray or purplish brown. Little seasonal variation apparent. Length of forewing: 7- 9 mm.
Male genitalia (Figs. 30, 31, 37, 38). Similar in general form to those of N. resistaria and brehmeata but with two conspicuous differences. Long, hairy coremata that arise near bases of saccular lobes in other species all but absent (vestiges remain), and prongs of bifurcate juxta are short in baggettaria. Prongs shorter than paired, longitudinally parallel sclerites lying off to each side of “neck” of juxta, whereas in resistaria prongs are much longer than these sclerites. Overall, genitalia smaller and more delicate, saccular lobe less produced, and vesica of aedeagus with smaller cornutus. Two specimens are illustrated to show variation. The prongs of the bifurcate juxta are asymmetrical in Fig.
74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
Pie NOE oa
= or
aha ore
SM SUT ate Pa NLR
rime
Fics. 37-39. Genitalia of N. baggettaria. 37, 6, Abita Springs, Louisiana. 38, Aedeagus of same specimen. 39, 2, Abita Springs, Louisiana.
37, which is probably not normal; and the aedeagus of the other specimen (Fig. 31) has a knoblike process on its proximal end, also abnormal.
Female genitalia (Figs. 36, 39). Main difference in female is in signum, which is small and of the simpler, two-pointed type found in some neotropical species. Signa of resistaria and brehmeata differ in being large, round or ovate disks bearing small surface points and fringed marginally with many more sclerotized points, which form a dentate margin on the signum that is widest toward its anterior end. Two specimens are illustrated, with the bursa copulatrix differently oriented.
Types. Holotype: 6, Torreya State Park, Liberty Co., Florida, H. D. Baggett; in U.S. National Museum of Natural History. Paratypes (19): Torreya State Park, Liberty Co.,
VOLUME 47, NUMBER 1 , 79
Florida, 17 July 1982 (1 4), H. D. Baggett; same locality and date (1 2), W. L. Adair; same locality, 4 July 1986, (1 4, 1 2); 27 June 1981 (1 4); 17 Aug. 1982 (1 ¢); 4 Sept. 1983 (1 ¢), H. D. Baggett. Eight mi N of Sumatra, Apalachicola Natl. Forest, Liberty Co., Florida, 10 May 1990 (1 2), 2 June 1990 (1 4), H. D. Baggett. Goose Pasture, Jefferson Co., Florida, 27 May 1989 (1 4), H. D. Baggett. Manatee Springs State Park, Levy Co., Florida, 16 July 1982 (1 2), H. D. Baggett. Cedar Key, Levy Co. (Hardwood Swamp, CR- 347, 5.4 mi N Jet. SR-24), Florida, 27 June 1987 (1 2), T. M. and L. Neal. 4.2 mi NE of Abita Springs, St. Tammany Parish, Louisiana, 25 Apr. 1984 (1 ¢), 7 May 1983 (1 3), 14 May 1984 (1 2), 25 May 1984 (1 2), 23 June 1983 (1 9), 16 July 1983 (1 4), 29 Sept. 1983 (1 6), V. A. Brou. Lumberton, North Carolina (on Interstate 95, at light), 21 Aug. 1987 (1 2), R. Gilmore. Paratypes deposited in U.S. National Museum, the Florida State Col- lection at Gainesville, and in the private collections of H. D. Baggett, V. A. Brou, and others.
Early stages. Unknown.
Distribution. Seen only from the Apalachicola National Forest and Torreya State Park, Liberty Co., Goose Pasture, Jefferson Co., and Manatee Springs and Cedar Key, Levy Co., Florida; Abita Springs, St. Tammany Parish, Louisiana; and Lumberton [Robeson Co.], North Carolina.
Flight period. Collected every month from April to September.
Material examined. Twenty-six specimens.
Remarks. This species is named for H. D. (Dave) Baggett of Palatka, Florida, who first brought it to my attention and who collected about half of the specimens seen.
ACKNOWLEDGMENTS
I thank H. D. Baggett of Palatka, Florida and V. A. Brou of Abita Springs, Louisiana, the lepidopterists who independently collected virtually all of the known material of the new species described above, and who brought it to my attention by sending specimens. I also acknowledge the assistance of those in charge of the Lepidoptera collections at the California Academy of Sciences, San Francisco, the University of California at Berkeley and Davis, Oregon State University at Corvallis, and the Florida State Collection of Arthropods at Gainesville for the opportunity to examine material in those collections; and I am particularly indebted to J. D. Lattin and J. C. Miller of the Department of Entomology, Oregon State University, for arranging travel support that made visits pos- sible on two occasions. Thus I studied the material at Corvallis and in the State Collection at Salem, Oregon, following which I returned to the region yet a third time in 1990 and was able to collect and see the habitats of both N. brehmeata and N. resistaria orfordensis. I thank the following for their reviews of the paper: John G. Franclemont, Department of Entomology, Cornell University, Ithaca, New York; Ronald W. Hodges and Paul M. Marsh, same address as the author; and Timothy L. McCabe, New York State Museum, Albany. McCabe also contributed biological information. The drawings are by Kellie L. Marsh, and the photographs are by the author.
LITERATURE CITED
BARNES, W. & J. H. MCDUNNOUGH. 1917. Check list of the Lepidoptera of boreal America. Herald Press, Decatur, Illinois. 392 pp.
Capps, H. W. 1943. Some American geometrid moths of the subfamily Ennominae heretofore associated with or closely related to Ellopia Treitschke. Proc. U.S. Natl. Mus. 93:115-151, pls. 1-10.
CASSINO, S. E. & L. W. SwETT. 1922. Some new geometrids. The Lepidopterist 3:155- 158.
Dyar, H.G. 1902 [1903]. A list of North American Lepidoptera and key to the literature of this order of insects. Bull. U.S. Natl. Mus. 52. xix + 723 pp.
FERGUSON, D. C. 1954. The Lepidoptera of Nova Scotia, part 1, macrolepidoptera. Proc. Nova Scotian Inst. Sci. 23:i-iv (unnumbered), 161-375.
76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
1983. Geometridae, pp. 68-107. In Hodges, R. W. et al. (eds.), Check list of the Lepidoptera of America north of Mexico. E. W. Classey Ltd. and The Wedge Entomological Research Foundation, London. xxiv + 284 pp.
ForBES, W. T. M. 1948. The Lepidoptera of New York and neighboring states, Pt. 2. Cornell Univ. Agr. Exp. Sta. Mem. 274:1-263.
FURNISS, R. L. & V. M. CAROLIN. 1977. Western forest insects. U.S. Dept. Agr. For. Serv. Misc. Publ. 1339. Washington, D.C. vii + 657 pp., illus.
GRIFFIN, F. J. 1932. Note on Haworth’s ‘Lepidoptera Britannica’ etc., 1803-1828. Ann. Mag. Nat. Hist. 9(10th ser.):531-532.
GROTE, A. R. 1872. Descriptions of Lepidoptera from Alabama. Canad. Entomol. 4:101- 102.
1882. North American Geometridae. Canad. Entomol. 14:106-111.
GUENEE, A. 1857. Uranides et Phalénites (pt. 1), pp. lvi + 1-514 and pls. 1-12 in Atlas (1958). Vol. 9. In Boisduval J. A. & A. Guenée (eds.), Histoire naturelle des insectes, species général des lépidoptéres. Libraire Encyclopédique de Roret, Paris.
Haworth, A. H. 1809. Lepidoptera Brittanica ..., etc., pt. 2. R. Taylor, London. Pp. 137-376.
HERRICH-SCHAFFER, G. A. W. 1850-1858. Sammlung neuer oder wenig bekannter aussereuropdischer Schmetterlinge. Vol. 1, series 1. G. J. Manz, Regensburg. 84 pp., 96 pls.
HuLst, G. D. 1896. Classification of the Geometrina of North America with descriptions of new genera and species. Amer. Entomol. Soc. Trans. 23:245-386.
INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE. 1985. International Trust for Zoological Nomenclature in association with the British Museum (Nat. Hist.), London. xx + 338 pp.
Ives, W. G. H. & H. R. Wonc. 1988. Tree and shrub insects of the prairie provinces. Information Rept. NOR-X-292, Northern Forestry Centre, Edmonton, Alberta. xii + 32” pp:, 117 col. pls.
KIMBALL, C. P. 1965. Lepidoptera of Florida. Arthropods of Florida and neighboring land areas. Vol. 1. Div. of Plant Industry, Florida Dept. Agric., Gainesville. 365 pp., illus. ;
LINNAEUS, C. 1767. Systema Naturae..., etc. Vol. 1(2), 12th Edition. Laurentii Salvii, Holmiae. Pp 533-1327. .
McDUNNOUGH, J. H. 1938. Check list of the Lepidoptera of Canada and the United States of America, pt. 1, macrolepidoptera. South California Acad. Sci. Mem. 1. 272 pp:
MOosHER, E. 1917. Pupae of some Maine species of Notodontoidea. Maine Agr. Exp. Sta. Bull. 259. Univ. Maine, Orono. 84 pp.
PACKARD, A. S. 1876. A monograph of the geometrid moths or Phalaenidae of the United States. Vol. 10 in F. V. Hayden, Rept. U.S. Geol. Surv. Terr. U.S. Govt. Printing Office, Washington. 607 pp.
PETERSON, A. 1948. Larvae of insects. Pt. 1, Lepidoptera and plant infesting Hyme- noptera. Printed for the author by Edwards Bros., Ann Arbor, Michigan. 315 pp.
PRENTICE, R. M. (compiler). 1963. Forest Lepidoptera of Canada recorded by the Forest Insect Survey. Vol. 3. Canada Dept. Forestry Publ. 1018. Pp. 281-543.
STEHR, F. W. 1987. Immature insects. Kendall/Hunt Publ. Co., Dubuque, Iowa. xiv + 754 pp., illus.
STEPHENS, J. F. 1829. A systematic catalogue of British insects ..., etc. Pt. 2. Baldwin and Cradock, London. 388 pp.
1829-{1831]. Illustrations of British entomology; or, a synopsis of indigenous insects ..., etc. Vol. 8. Baldwin and Cradock, London. 333 pp. + 5 unnumbered pp., pls. 25-381.
SWETT, L. W. 1913. Geometrid notes—New varieties. Canad. Entomol. 45:75-76.
TiETZ, H. M. 1972. An index to the described life histories, early stages and hosts of the macrolepidoptera of the continental United States and Canada. Allyn Mus. En- tomol., Sarasota, Florida. iv + 1041 pp. in 2 vols.
VOLUME 47, NUMBER 1 Bath
WALKER, F. 1860. List of the specimens of lepidopterous insects in the collection of the British Museum, pt. 20. Trustees of the British Museum, London. Pp. 1-498. Woop, W. 18389. Index Entomologicus ..., etc. Publ. by the author. Covent Garden,
London. 267 pp., 54 pls. 1854. Index Entomologicus. New edition revised by J. O. Westwood. G. Willis,
Covent Garden, London. 298 pp., 59 pls.
Received for publication 1 October 1992; accepted 4 October 1992.
GENERAL NOTES
Journal of the Lepidopterists’ Society 47(1), 1998, 78-79
DIMORPHISM IN THE FEMALE OF BATTUS ZETIDES (PAPILIONIDAE) ON HISPANIOLA
Additional key words: endemism, Antilles, systematics, variation.
N. D. Riley (1975) recorded Battus zetides Munroe (Papilionidae) as poorly known; in fact, based on the few specimens then extant, he incorrectly illustrated the species as lacking tails. Subsequent reports indicate that B. zetides can be found in isolated enclaves of upland mesic broadleaf deciduous forest, occurring on both the north and south paleoislands of Hispaniola (Gali & Schwartz 1983, Schwartz 1989, Johnson & Matusik 1988). Both Riley (1975) and Schwartz (1989) report the sexes of B. zetides as alike on both wing surfaces, as typical of most Battus species (D’Abrera 1981).
In 1988, we detailed location and habitat of the remote “Las Abejas’ forest, Parque Nacional Sierra de Bahoruco [sic] (Pedernales Province, Dominican Republic) (Johnson & Matusik 1988). This upland forest, visited by us each year from 1981-1991, harbored a prolific population of B. zetides (Gali & Schwartz 1988) but is now suffering severe deforestation (Johnson & Matusik 1988, Johnson 1989).
The purpose of this note is to document a striking dimorphic female form of B. zetides occurring at Las Abejas. Contrasting the “ochre-yellow” or “yellows and oranges’ gen- erally attributed to wingbands of B. zetides (Riley 1975, Schwartz 1989; Fig. 1A), bands in this new form are mostly white, occasionally mottled pale yellow in distal areas of cells M, to lA + 2A (Fig. 1B). This whitened condition, extending the silverlike appearance of the ventral hindwings (Fig. 1B, right), creates an identification problem in the field because such individuals resemble no other papilionid species known from the neotropics. To our knowledge, striking dimorphism in females of Battus has not been previously reported.
Ten specimens of the whitened female form have been collected at Las Abejas since 1986 (Specimen Data below) but, because our collections of B. zetides have been widely disseminated in public and private collections since 1981, it is difficult to quantify fre- quency of occurrence. Although females of B. zetides are generally less vagile than males and seldom venture from the forest canopy, females are probably more readily collected at Las Abejas than at other Hispaniolan locales since, as reported before (Johnson & Matusik 1988), steep ravine edges surrounding “Lower Abejas” allow for fortuitous col- lecting of the bottomland canopy. No doubt this unique collecting situation accounts for discovery of the form, which probably occurs in all populations of the species. A recon- structed estimate of frequency, based simply on recollection of collecting conditions for B. zetides at Las Abejas on a “day to day” basis, suggests females (usually immediately released) constituted about 5% of our catch. With this in mind, the ten known specimens of the whitened female probably represented about 1% of the females taken by us at the site.
It is important to note the striking appearance of this female form in the field. Field sightings have figured importantly in the historical documentation of certain rare, or seldom-collected, Antillean butterflies (Schwartz 1989, Brown & Heineman 1972). Schwartz (1989) has noted the soaring flight of B. zetides and that the butterfly is seen much more often than collected. The peculiar whitened female form of B. zetides should be antic- ipated by collectors and not misconstrued in flight as possibly representing an unknown neotropical swallowtail.
Specimen data (numbers parenthetical). All “Las Abejas forest” [detailed above], D. Matusik collector: 1-9 August 1991 (6) David Matusik Collection (DMC), 30 July 1990 (1) American Museum of Natural History (AMNH) (Fig. 1B), 29 June 1989 (2) (DMC), 5 July 1986 (1) (DMC).
VOLUME 47, NUMBER 1 . 79
Fic. 1. Battus zetides. A, Typical female (“Lower Abejas,” sensu Johnson & Matusik [1988], 31 July 1990, AMNH) upper surface left, under surface right. B, Whitened female form (same data, except 30 July 1990, AMNH) same views.
LITERATURE CITED
Brown, F. M. & B. HEINEMAN. 1972. Jamaica and its butterflies. E. W. Classey, Hampton, Middlesex. 478 pp.
D’ABRERA, B. 1981. Butterflies of the Neotropical Realm. Part 1. Papilionidae and Pieridae. Landsdowne Editions, East Melbourne. 172 pp.
GALI, F. & A. SCHWARTZ. 1983. Battus zetides in the Dominican Republic. J. Lepid. Soc. 37:171-174.
JOHNSON, K. 1989. Letter to the Editor [concerning habitat destruction in the Parque Nacional Sierra de Bahoruco]. Lepidopterists News 43:89.
JOHNSON, K. & D. MATusIK. 1988. Five new species and one new subspecies of butterflies from the Sierra de Baoruco of Hispaniola. Ann. Carnegie Mus. 57:221-254.
RILEy, N. D. 1975. A field guide to the butterflies of the West Indies. New York Times Book Company, Garden City. 224 pp.
SCHWARTZ, A. 1989. The butterflies of Hispaniola, Univ. Florida Press, Gainesville. 580 pp.
KURT JOHNSON, Department of Entomology, American Museum of Natural History, Central Park West at 79th St., New York, New York 10024; AND DaviD MATUSIK, Department of Entomology, Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605.
Received for publication 24 December 1990; revised and accepted 26 September 1992.
Journal of the Lepidopterists’ Society 47(1), 1998, 80-82
THE AHRENHOLZ TECHNIQUE FOR ATTRACTING TROPICAL SKIPPERS (HESPERIIDAE)
Additional key words: Pyrrhopyginae, Pyrginae, Hesperiinae, army ants, behavior.
Skippers (Hesperiidae) constitute a large component of tropical butterfly communities. In South America, sites in mature lowland tropical forest may contain several hundred species. Although some species, especially those favoring clearings and heavily disturbed forests, may be locally common, most forest skippers occur at low population densities, as is generally true for tropical butterflies (Ebert 1969). Their swift, erratic flight is difficult to follow, and oftentimes, they are extremely wary. Thus, collecting and photographing forest skippers is difficult.
Various methods overcome the problem of sampling species that are otherwise rare and wary. Some butterflies are attracted to traps baited with rotting animals, excrement, decaying fruit, or pheromones. Others seek mud or sweaty clothes, presumably attracted to the sodium chloride. Malaise and light traps also may sample otherwise scarce species, and colored pieces of cloth, paper, or plastic can be effective in drawing high-flying species to the ground. Some butterflies, such as Ithomiinae and Danainae, seek pyrroli- zidine alkaloids, whether in the nectar of Asteraceae or in decaying heliotrope plants (Boraginaceae).
Most skippers are not attracted to bait traps, show no interest in colored rags or plastics, and generally are not collected in Malaise and light traps. Only a small percentage of skippers sip moisture at mud, and most of these are males. Flowering shrubs or trees, which are heavily frequented by both sexes, are very scarce or patchy in South American rain forests and usually occur 20-45 m above the ground in the canopy.
It has been observed repeatedly, however, that many hesperiid species visit fresh bird droppings in the interior of the forest (mostly upon leaves or the ground) or on rocks, stones, or sand on river banks. Further, a number of otherwise scarce skippers congregate in the neighborhood of army-ant (Formicidae: Ecitoninae) swarms in southeastern Peru (Lamas 1983), where they feed on antbird (Formicariidae) droppings. Taking advantage of these peculiar congregations, first described by Zikan (1929) for Brazilian skippers, we have collected some species that are poorly represented in collections. Unfortunately, the thick underbrush and wariness of these skippers still makes it difficult to approach them.
The purpose of this note is to report a new method, which we name the “Ahrenholz Technique,’ for attracting skippers in Neotropical rain forests. It was devised and suc- cessfully used by our good friend David Ahrenholz in Rond6énia, Brazil and eastern Ecuador. After he described the method to us in 1990, we experimented with it at Pakitza, Manu National Park; at Tambopata, Tambopata-Candamo Reserved Zone; and at Pampas del Heath National Sanctuary, in southeastern Peru (see Erwin 1985, 1991, Lamas 1985, Lamas et al. 1991 for location and description of the two former sites). We summarize our observations and those of Ahrenholz.
The Ahrenholz Technique
We placed small (ca. 1 cm?), approximately square, pieces of toilet or tissue paper, wetted with saliva, on the uppersides of exposed, broad leaves, or on cleared patches of ground. These pieces of paper have a rough resemblance to fresh bird droppings and attracted many species of skippers, Nymphalidae (Caeruleuptychia, Pyrrhogyra, Adelpha, Nessaea, Catonephele, Marpesia, Memphis, Heliconius, Forbestra), Riodinidae (Euse- lasia, Ancyluris, Thisbe), and Pieridae (Dismorphia). Other insects, including small flies, wasps, and orthopterans, visited the paper too. We also placed similar pieces of wet paper on sand at river banks, where only skippers and a few flies were attracted. We experi- mented with white, sky blue, light green, and light pink colored pieces of paper, without perceiving any differences in attractiveness. The exact shape of the paper did not seem to matter much either.
VOLUME 47, NUMBER 1 81
Fic. 1. Astraptes fulgerator (Walch) (Pyrginae), on right, and Aides duma argyrina Cowan (Hesperiinae), attracted using the Ahrenholz technique, at Fazenda Rancho Gran- de, Rondénia, Brazil. Photograph courtesy of D. Ahrenholz.
A skipper, on discovering a piece of paper, landed quickly upon it, extended its proboscis and probed the wet paper for several seconds to a few minutes (Fig. 1). It remained wary, but was relatively easy to approach because we placed the paper in an exposed site. If the butterfly was collected, we replaced the paper, which usually fell into the net or on the ground, and wetted it again because dry paper was less attractive to the skippers. Although butterflies investigated paper wetted with rain water, they flew off after ex- tending their proboscides. Other liquids, such as urine and sweetened soft drinks, did not work as well as saliva. Inside the forest, the paper attracted butterflies even if no ants were present, although with less success. Along river banks, they were effective whenever skipper butterflies were in the area.
Although skippers may use olfactory and/or auditory cues generated by the ants, their prey, and/or the antbirds to find ant swarms, they appear to locate bird droppings, or their “mimics” (the pieces of paper) visually, as shown at the river banks, where no ant swarms have been observed.
We list the skipper genera collected at Pakitza and Tambopata in October 1991, and at Pampas del Heath in June 1992, using the Ahrenholz Technique: A) Inside the forest. Pyrrhopyginae: Jemadia; Pyrginae: Phocides, Phanus, Udranomia, Epargyreus, Au- giades, Aguna, Polythrix, Chrysoplectrum, Urbanus, Astraptes, Dyscophellus, Tele- miades, Pachyneuria, Clito, Zera, Quadrus, Gindanes, Milanion, Anastrus, Antigonus, Aethilla and Achlyodes; Hesperiinae: Vidius, Vettius, Justinia, Ebusus, Tigasis, Thoon, Talides, Tisias, Carystus, Carystoides, Perichares, Orses, Lycas, Metron, Phemiades, Panoquina, Oxynthes, Niconiades, Aides, Saliana, Thracides, Aroma and Pyrrhopygop-
82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
sis. B) At river banks. Pyrrhopyginae: Pyrrhopyge, Elbella, and Jemadia; Pyrginae: Phocides, Proteides, Epargyreus, Polygonus, Antigonus, Anastrus, Ebrietas, Campto- pleura and Cycloglypha; Hesperiinae: Metron, Lindra and Panoquina.
We are grateful to Dave Ahrenholz for allowing us to report his innovative technique and results. We thank the staff at Pakitza, Tambopata, and Pampas del Heath for sup- porting our research. This note is contribution no. 51, Biological Diversity in Latin America (BIOLAT) Project, Smithsonian Institution, and contribution no. 712, Departamento de Zoologia, Universidade Federal do Parana.
LITERATURE CITED
EBERT, H. 1969. On the frequency of butterflies in eastern Brazil, with a list of the butterfly fauna of Pocos de Caldas, Minas Gerais. J. Lepid. Soc. 23, Suppl. 3, 48 pp.
ERWIN, T. L. 1985. Tambopata Reserved Zone, Madre de Dios, Peru: History and description of the Reserve. Rev. Per. Entomol. 27:1-8.
1991. Natural history of the carabid beetles at the BIOLAT Biological Station, Rio Manu, Pakitza, Peru. Rev. Per. Entomol. 33:1-85.
Lamas, G. 19838. Mariposas atraidas por hormigas legionarias en la Reserva de Tam- bopata, Peru. Rev. Soc. Mex. Lepid. 8(2):49-51.
LAMaS, G. 1985. Los Papilionoidea (Lepidoptera) de la Zona Reservada de Tambopata, Madre de Dios, Peru. I: Papilionidae, Pieridae y Nymphalidae (en parte). Rev. Per. Entomol. 27:59-78.
Lamas, G., R. K. ROBBINS & D. J. HARvEy. 1991. A preliminary butterfly fauna of Pakitza, Parque Nacional del Manu, Peru, with an estimate of its species richness. Publ. Mus. Hist. Nat. UNMSM (A) 40:1-19.
ZIKAN, J. F. 1929. Myrmekophilie bei Hesperiden? Entomol. Rundschau 46(7):27-28.
GERARDO LAMAS, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Apartado 14-0434, Lima-14, Peru; OLAF H. H. MIELKE, Departamento de Zoologia, Universidade Federal do Paranda, Caixa Postal 19020, 81.531 Curitiba, PR, Brazil; AND ROBERT K. ROBBINS, Department of Entomology, NHB Stop 127, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, USA.
Received for publication 4 September 1992; revised and accepted 4 October 1992.
Journal of the Lepidopterists’ Society 47(1), 1998, 83-84
OBITUARY
DONALD PAUL FRECHIN (1918-1991)
Don Frechin, a Charter Member of the Lepidopterists’ Society and well-known Wash- ington state entomologist, passed away on 10 December 1991, at the age of 73. Don was born in Garden City, Kansas, on 6 January 1918; he moved with his parents to Bremerton, Washington, when he was 9 months old. He was an avid bug collector throughout his boyhood in western Washington. Following in his father’s footsteps, he began working at the Navy shipyards in Bremerton in 1940. He married Gudrun (Gudy) Loyland in 1942 and raised seven children. In 1959, he took a job with the Boeing Company near Seattle, commuting by ferry for five years across Puget Sound. In 1964 the family moved to their north Seattle residence at 1745 Northeast 102nd Street, where Don lived out his life.
Don was an active collector throughout his life and exchanged specimens and infor- mation with many members of the Society. He was an ardent general collector, picking up examples of virtually all orders that could be pinned. Among the Lepidoptera, his passions included butterflies and larger moths, especially tiger moths and hepialids. Rear- ing insects was one of his favorite pastimes—he always had livestock around of sundry swallowtails, saturniids, and arctiids. For a time, he was involved with handpairing and hybridization of arctiids. Later in life he focused his attentions on tiger beetles, amassing a collection of nearly 8000 specimens.
Anyone who knew Don could not help but be impressed by his enthusiasm for, and knowledge of, Washington’s insect fauna. As a graduate student studying the biosyste- matics of ghost moths, I sought out Don after seeing many of his hepialid specimens in major collections. Upon contacting Don, I found him to be the most knowledgeable Lepidopterist in North America on the habits of this seldom encountered family of Lepidoptera. I will never forget the time he (at the age of 64) took me to a trail at Steven’s Pass in the northern Cascades, where he thought I might see Gazoryctra roseicaput. It was a cool, blustery September evening, clouds had engulfed the entire pass reducing visibility to little more than 30 feet. By dusk, when the moths were supposed to fly, the
84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY
temperature had dropped to 48°F (=9°C). I was sure no moths would be on the wing in such weather and suggested to Don that our time would be better spent at a lower, warmer elevation ... perhaps in a restaurant with beverages on tap. He smiled and countered with a “wait and see” look. Sure enough, the moths appeared, when and where Don suggested—they disappeared almost as quickly, just thirty minutes later.
The size and nature of Don's collection was ever-changing as he often traded or sold parts of it. Eugene Munroe, acting on behalf of the Canadian National Collection (CNC), purchased most of Don’s moths in the late 1950's, including Edward C. Johnston’s im- portant collection which Don recently had acquired. In the mid-1980’s, I purchased Don’s synoptic collection of Washington state Macrolepidoptera; this collection of 700 specimens is housed at the University of Connecticut. More than 39,000 specimens remain at his home in Seattle; the bulk of these are 22,000 Coleoptera and 14,500 Lepidoptera (mostly papered or unspread).
At least two species and one genus of insects were named after Don. Munroe described the pyralids Pogonogenys frechini and the genus Frechinia. Sanford Leffler described the tiger beetle, Cicindela bellissima frechini, endemic to the Olympic Peninsula, after him.
Don is survived by his wife, six children, and four grandchildren. Part of his legacy will be the countless collections he made of Washington state invertebrates, especially from the biologically unique areas that he so often visited, such as the Rocky Prairie near Tenino, the Olympic Peninsula, and the arctic-alpine areas of the northern Cascades. His absence from the Society will be keenly felt.
Davip L. WAGNER, Department of Ecology and Evolutionary Biology, U-Box 48, University of Connecticut, Storrs, Connecticut 06269-30438.
Journal of the Lepidopterists’ Society 47(1), 1993, 85
MANUSCRIPT REVIEWERS, 1992
The merit of a scientific journal depends on the quality of its reviewers as well as of its authors, but the former are usually unknown to readers. The Journal relied on the expertise of 68 reviewers last year to provide 93 evaluations of manuscripts. It is with much gratitude that the Journal acknowledges the services of the people listed below from whom manuscript reviews were received in 1992.
Elizabeth A. Bell, Santa Cruz, CA *Elizabeth A. Bernays, Tempe, AZ *Lincoln P. Brower, Gainesville, FL
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MS John M. Burns, Washington, DC
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* Reviewed two or more manuscripts.
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Journal of the Lepidopterists’ Society 47(1), 1993, 86
ANNOUNCEMENT PUBLICATIONS OF THE LEPIDOPTERISTS’ SOCIETY
THE LEPIDOPTERISTS SOCIETY COMMEMORATIVE VOLUME, 1945-1978. Published 1977. 374 pages. A 25-year review of the Society’s or- ganization, personnel, and activities, with biographical sketches. Includes a 25-year cumulative index of the Journal of the Lepi- dopterists’ Society by author, subject, and taxon. Members and subscribers: $8.00. Non-members: $12.00.
MEMOIRS OF THE LEPIDOPTERISTS’ SOCIETY
Memoir No. 1. A Synonymic List of the Nearctic Rhopalocera by Cyril F. dos Passos. Published 1964. 145 pages. Includes references to original descriptions and synonymies of North American but- terflies and skippers. OUT-OF-PRINT.
Memoir No. 2. Catalogue/Checklist of the Butterflies of America North of Mexico by Lee D. Miller and F. Martin Brown. Published 1981. 280 pages. Includes references to original descriptions, syn- onymies, and locations of type specimens. Members and subscrib- ers: $12.00 cloth, $7.00 paper. Non-members: $19.00 cloth, $10.50 paper. |
Memoir No. 3. Supplement to the Catalogue/Checklist of the Butterflies of America North of Mexico by Clifford D. Ferris (editor). Published 1989. Includes general notes plus corrections
and additions to the original Memoir No. 2. Members and sub- scribers: $6.00. Non-members: $10.00.
Memoir No. 4. Foodplants of World Saturniidae by Steven Stone. Published 1991. A listing of foodplants for more than 500 species of saturniid moths worldwide. Members and subscribers: $7.20. Non-members: $12.00.
1992 MEMBERSHIP DIRECTORY (currently to October 1992). Published 1992. 78 pages. Biennial directory of members of the Lepidop- terists’ Society, with members’ geographic and taxonomic interests. $5.00. Not available for commercial use.
Send orders to: Ron Leuschner, Publications Coordinator 1900 John Street Manhattan Beach, California 90266-2608 U.S.A.
Date of Issue (Vol. 47, No. 1): 24 February 1993
EDITORIAL STAFF OF THE. JOURNAL
JOHN W. Brown, Editor Entomology Department San Diego Natural History Museum P.O. Box 1390 San Diego, California 92112 U.S.A.
Associate Editors: M. DEANE Bowers (USA), BOYCE A. DRUMMOND (USA), LAWRENCE F. GALL (USA), _ GERARDO LAMAS (Peru), ROBERT C. LEDERHOUSE (USA), ROBERT K. ROBBINS (USA), CHRISTER WIKLUND (Sweden)
NOTICE TO CONTRIBUTORS
Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Technical Comments, Book Reviews, Obituaries, Feature Photographs, and Cover Illustrations. Reviews should treat books published within the past two years. Obituaries must be authorized by the President of the Society. Re- quirements for Feature Photographs and Cover Illustrations are stated on page 111 in Volume 44(2). Journal submissions should be sent to the editor at the above address. _ Short manuscripts concerning new state records, current events, and notices should be
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Literature Cited: References in the text of Articles, Profiles, General Notes, and Technical Comments should be given as Sheppard (1959) or (Sheppard 1959, 196la, 1961b) and listed alphabetically under the heading LITERATURE CITED, in the following format without underlining:
SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 209 pp.
196la. Some contributions to population genetics resulting from the study of
the Lepidoptera. Adv. Genet. 10:165-216.
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CONTENTS
PRESIDENTIAL ADDRESS, 1992: MEGATRENDS AND THE LEPI- DOPTERISTS SOCIETY. Floyd W. Preston
THE NATURE OF ANT ATTENDANCE AND THE SURVIVAL OF LARVAL ICARICIA ACMON (LYCAENIDAE). Merrill A: Peterson
BISTON BETULARIA (GEOMETRIDAE), THE PEPPERED MOTH, IN WIRRAL, ENGLAND: AN EXPERIMENT IN ASSEMBLING. Cyril A. Clarke, Frieda M. M. Clarke and Bruce Grant
TERRITORIALITY ALONG FLYWAYS AS MATE-LOCATING BEHAVIOR IN MALE LIMENITIS ARTHEMIS (NYMPHALIDAE). Robert C. Lederhousé 2 0) se
DEVELOPMENTAL CHANGES AND WEAR OF LARVAL MANDIBLES IN HETEROCAMPA GUTTIVITTA AND H. SUBROTATA (NOTODONTIDAE). Dawn E. Dockter
DESIGNATION OF A LECTOTYPE OF NISONIADES SOMNUS AND NOTES ON THE OCCURRENCE OF ERYNNIS ICELUS IN FLORIDA (HES- PERIIDAE). John V.-Calhoun. 2. 1 3 :
First NORTH AMERICAN RECORDS OF EPINOTIA ABBREVIANA (TORTRICIDAE), A EUROPEAN PEST OF ULMUS SPECIES. P. TS DGN8 ce
A REVISION OF THE SPECIES OF NEMATOCAMPA (GEOMETRIDAE: ENNNOMINAE) OCCURRING IN THE UNITED STATES AND CAN- ADA. (Douglas C.. Ferguson 0
GENERAL NOTES
Dimorphism in the female of Battus zetides (Papilionidae) on Hispaniola. Kurt Johnson and David Matusik’) 0 ee ee
The Ahrenholz technique for attracting tropical skippers (Hesperi- idae). Gerardo Lamas, Olaf H. H. Mielke and Robert K. Robbins
OBITUARY Donald Paul Frechin (1918-1991). David L. Wagner 0
MANUSCRIPT REVIEWERS; 199200 ee
ANNOUNCEMENT
PUBLICATIONS OF THE LEPIDOPTERISTS SOCIETY
THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.
22
49
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60
i a 4 Volume 47 LOE ran a Number 2
ISSN 0024-0966
JOURNAL
of the
LEPIDOPTERISTS’ SOCIETY
Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS
16 June 1993 De Arie re,
THE LEPIDOPTERISTS’ SOCIETY
EXECUTIVE COUNCIL
Ray E. STANFORD, President HIROSHI INOUE, Vice President FLoyD W. PRESTON, Immediate Past President IAN KITCHING, Vice President M. DEANE BOWERS, Vice President ROBERT J. BORTH, Treasurer
WILLIAM D. WINTER, Secretary
Members at large:
Karolis Bagdonas Charles V. Covell, Jr. Eric H. Metzler Steven J. Cary Linda S. Fink Robert K. Robbins Stephanie S. McKown Scott E. Miller J. Benjamin Ziegler
EDITORIAL BOARD
PAUL A. OPLER (Chairman), FREDERICK W. STEHR (Member at large) JOHN W. BROWN (Journal), WILLIAM E. MILLER (Memoirs) STEPHANIE S. MCKOWN (News)
HONORARY LIFE MEMBERS OF THE SOCIETY
CHARLES L. REMINGTON (1966), F. MARTIN BROWN (1973), E. G. MUNROE (1978), ZDRAVKO LORKOVIC (1980), IAN F. B. COMMON (1987), JOHN G. FRANCLEMONT (1988), LINCOLN P. BROWER (1990), DOUGLAS C. FERGUSON (1990),
HON. MIRIAM ROTHSCHILD (1991), CLAUDE LEMAIRE (1992)
The object of the Lepidopterists’ Society, which was formed in May 1947 and for- mally constituted in December 1950, is “to promote the science of lepidopterology in all its branches, ... . to issue a periodical and other publications on Lepidoptera, to facilitate the exchange of specimens and ideas by both the professional worker and the amateur in the field; to secure cooperation in all measures’ directed towards these aims.
Membership in the Society is open to all persons interested in the study of Lepi- doptera. All members receive the Journal and the News of the Lepidopterists Society. Institutions may subscribe to the Journal but may not become members. Prospective members should send to the Treasurer full dues for the current year, together with their full name, address, and special lepidopterological interests. In alternate years a list of members of the Society is issued, with addresses and special interests. There are four numbers in each volume of the Journal, scheduled for February, May; August and November, and six numbers of the News each year.
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Send remittances, payable to The Lepidopterists’ Society, to: Robert J. Borth, Treasurer, 6926 North Belmont Lane, Fox Point, Wisconsin 53217, U.S.A.; and address changes to: Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, California 90007-4057 U.S.A. For information about the Society, contact: William D. Winter, Sec- retary, 257 Common St., Dedham, Massachusetts 02026-4020, U.S.A. (617-326-2634). To order back issues of the Journal, News, and Memoirs, write for availability and prices to the Publications Manager: Ronald Leuschner, 1900 John St., Manhattan Beach, Cali- fornia 90266-2608, U.S.A.
Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for $40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los Angeles, California 90007-4057. Second-class postage paid at Los Angeles, California and additional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, California 90007- 4057.
Cover illustration: A silverspot butterfly (Speyeria sp.) resting beneath the leaf of a fern. Original drawing by Martie Clemons, Dudek and Associates, Inc., 605 Third Street, Encinitas, California 92024.
JOURNAL OF
Tue LeEpiIporpreRIstTs’ SOCIETY
Volume 47 1993 Number 2
Journal of the Lepidopterists’ Society 47(2), 1993, 87-105
BIOLOGY AND POPULATION DYNAMICS OF PLACIDULA EURYANASSA, A RELICT ITHOMIINE BUTTERFLY (NYMPHALIDAE: ITHOMIINAE)
ANDRE VICTOR LUCCI FREITAS
Curso de Pés-Graduagao em Ecologia, Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, C.P. 6109, 13081 Campinas, Sao Paulo, Brazil
ABSTRACT. Placidula euryanassa is a primitive ithomiine restricted to the Atlantic coast of South America. Females lay eggs in clusters on Brugmansia suaveolens (Sola- naceae). The larvae are gregarious, passing through five instars. Pupation occurs off the host plant; adults emerge after 8 to 14 days. In the laboratory the sex ratio is statistically equal to unity; in field captures the sex ratio is male biased. Individual mobility is low and the population size varies greatly during the year. Adults show a type II survival curve. Many biological features indicate that Placidula is relatively more r-selected than most Ithomiinae.
Additional key words: immatures, r-strategist, mark-recapture, Solanaceae.
Placidula euryanassa (Felder & Felder) is a member of the subfamily Ithomiinae. The systematic position of the species is uncertain (Brown 1987, Motta 1989). The genus Placidula is