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Eggs and Rapagula of Ululodes and Ascaloptynx (Neuroptera

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Eggs and Rapagula of Ululodes and Ascaloptynx (Neuroptera PSYCHE Vol. 79 March-June, 1972 No. I-2 EGGS AND RAPAGULA OF ULULODE8 AND ztSGztLOPTYNX (NEUROPTERA: ASCALAPHIDAE): A COMPARATIVE STUDY* BY CHARLES S. HENRY Biological Laboratories Harvard University, Cambridge, Mass. o2138 I. INTRODUCTION Ascalaphidae is a fairly large amily o planipennian Neuroptera, encompassing perhaps 300 species in 65 genera, yet relatively little is known of the biology ot its various representatives. Although several systematic monographs concerned exclusively with the family have been published (Lefebvre, I842; Rambur, I842; McLachlan, I87I; Van der Weele, I9O8; Navels, I913), attempts to discern the true phylogenetic relationships among the various genera and other taxa must necessarily await studies o lie-history and behavior. In t:act, definite larva-adult associations are lacking for a great majority of ascalaphid species. Larvae have been reliably identified only (or the genera ztscalaphus Fabricius, Helicomitus McLachlan, Pseudo- ptynx Weele, Suhpalacsa Lefebvre, and Ululodes Currie, all be- longing to the "split-eyed" subfamily Ascalaphinae; o the "entire- eyed" subfamily Neuroptynginae, only zlscaloptynx Banks has been associated with a larval type (MacLeod, 97o: p. 55). I have succeeded in rearing an additional species o Ululodes t:rom egg to adult: Ululodes mexicana (McLachlan), common in northern Mexico and southern Arizona and New Mexico. I have also been able to compare the (orm and habits of the various lie stages of this species with those ot a sympatric neuroptyngine species, ./lscalo- ptynx furciger (McLa.chlan). The identity of this last-named species is inferred, but based upon various types of evidence: (I) third- instar larvae, raised from field-collected eggs, were positively identi- fied by Dr. Ellis G. MacLeod (University of Illinois) as those of Ascaloptynx; (2) the only species of the genus Ascaloptynx--or, for that matter, of subfamily Neuroptynginaemever taken at the *Manuscript received by the editor August 1, 1972. 2 Psyche [March-June collecting sites in question is [urciger; (3) males and females o dscaloptynx [urciger were captured in fair abundance in the same areas and at the same time of year that eggs and young larvae were collected; (4) dissections of gravid d. [urciger emales revealed eggs and egg-attendants (see text) that were in every respect identical to field-laid eggs that yielded dscaloptynx larvae. The first student of ascalaphid life-history was. the Reverend L. Guilding, whose 827 account of the vario,us life stages Ululodes macleayana (Guilding)* rom St. Vincent's Island in the West Indies remains one o.f the best such studies, ever published. In de- scribing the eggs o.f this species, he noted the presence of curious rod-shaped structures encircling the twiff beneath the. egg-mass. He called these egg-attendants "repagula,"** meaning "barriers," be- cause he believed that they prevented ants and other predators from approaching and destroying the eggs and newly hatched larvae; con- versely, he felt that the repagula discouraged the larvae from leaving their twig "until they have acquired strength to resist... [their] enemies." McClendon (9o) also noted repagula accompanying the eggs of Ululodes hyalina (Latreille) [believed by MacLeod (964) to be Ululodes senex (Burmeister) ] rom Texas; he con- firmed Guilding's description of these structures and co.ncluded from his dissections of gravid female ascalaphids (unfortunately unillus- trated) that certain ovarioles ("tubercles") were specialized for the production of repagula. Thus it was shown that repagula were in act abortive eggs. Seventy years later, New (1971), apparently overlooking the papers of Guilding and McClendon, re-iotroduced the concept of ovariolar dimorphism in his cursory study o the eggs and repagula of eight species of Ascalaphidae rom central Brazil. New's paper is important, nonetheless, for two reasons: first, it proved that repagula-formation is not unique to the genus Ululodes but is also found in the closely-allied genera Colobopterus Rambur, Ascalorphne Banks and Cordulecerus Rambur, and in the neuro- ptyngine genera Byas Rambur and Episperches Gerstaecker secondly, it sug,ested that the repagula of neuroptyngine species are funda- mentally different in orm from those of ascalaphine species the former type more nearly resembling fertile eggs than the latter type. The functional significance of repagula has not been investigated. Most authors (Irnms, 1957, for example) have. accepted Guilding's "ant guard" view. New (1971) intimates that repagula and eggs *Guilding's name for this species was dscalaphus macIeayanus. **singular repagulum. 1972] Henry Ululodes and AscMoptynx 3 alike o Ululodes were carried off by ants in Brazil, but admits that more controlled experimentation is needed to assess the effects of the repagula on egg or larval survival. Through my studies o the lie-histories of Ululodes mexicana and Ascaloptynx furciger, I have been placed in a unique position to verify or disprove experi- mentally the ant-guard hypothesis, as it applies to these representa- tive species of two taxonomically distant genera. Especially important is the fact that Ululodes is an ascalaphine genus while Ascaloptynx is a neuroptyngine one, thus encouraging direct comparison o the unctional signicance of the morphologically disparate repagula char- acteristic of the members of these subfamilies. In the remainder o this paper, I intend (I) to describe the eggs and repagula o Ululodes mexicana and Ascaloptynx furciger, (2) to outline the procedures and results of several simple experiments undertaken to test the protective importance of the repagula o bo,th species, and (3) to discuss the unctional significance o the ascalaphine type of egg and repagula as compared to that of the neuroptyngine type. II. ACKNOWLEDGEMENTS Special thanks are extended to Professor Frank /I. Carpenter, my advisor and friend for the past five years. All aspects o the work in this paper draw heavily rom his awesome knowledge o insect behavior, morphology, evolution and taxonomy, especially o the order Neuroptera. I am also in his debt or his help in guiding this study into publishable orm. My wie, Nancy FitzGerald, should also be cited as a primary source of inspiration for this project. Her shrewd suggestions and sharp observations in the ield are apparent to. me. again and again as ! review my notebooks nd ind her influence in their pages. In addition, warm thanks are extended to: Pro.essor Ellis G. MacLeod (University o.f Illinois), for his generous willingness to transmit his intimate knowledge of the habits of all sorts of neuropterous insects to a neophyte like myself, and for confirming the identity of Ascaloptynx larvae; Professo.r Edward O. Wilson (Harvard Biological Laboratories), for enthusiastically supporting my research effort and for determining the genera, and species of ants used in the ant-guard experiments; Mr. Vincent Roth of the South- western Research Station, for his assistance with my field work in Arizona; Dr. Lauren Anderson (University of California at River- side), for his advice on securing newly-hatched ascalaphids; and Mr. Robert E. Silberglied (Harvard Biological Laboratories), t;or his photographic help and biological acumen. Psyche [March-June Figure 1. Typical sites for oviposition, Ululodes mexicana (l-a) and Ascaloptynx [urciger (l-b). Year-round financial support for my studies was for three years provided by the National Science Foundation, in the form of a Pre-Doctoral Fellowship (September, 968 to September, I97). A final fourth year of research was completed with the aid of a Rich- mond Fellowship (97I-'72) offered through Harvard's Department of Biology. Unusual expenses associated with travel to the field and with publication of this paper were generously advanced by the Committee on Evolutionary Biology, Harvard University (NSF Grant GB 279, Reed Rollins, Harvard University, Principal Investigator). III. METHODS AND MATERIALS Eggs o both Ululodes mexicana and Ascaloptynx furciger were collected during August and September in the southeastern part o Arizona, within a 25-mile radius o the Southwestern Research Station o the American Museum o Natural History (SWRS). All egg-masses, 25 of Ululodes and 35 of Ascaloptynx, were taken on shrubs in the arroyos and canyons of the Chiricahua and Peloncillo Mountains, at elevations of 4700 to 570o feet (5oo-8oo meters). Several egg-masses were left in situ and visited twice daily; others 1972] Henry Ululodes and Ascalottynx 5 were placed twigs and all in 2.5 cm X 9.0 cm glass screw-top vials and transported carefully to the research station laboratory. There, the egg-bearing twigs were taped in their "natural" positions, covered with large glass jars for observation and situated in a southwest window shielded by heavy curtains orm the vicissitudes of indoor artificial light. Complete records from oviposition through hatching to descent of larvae were obtained for seven egg-masses of Ululodeg and eight of Ascaloptynx. Additional egg-clumps of both species of Ascalaphidae were exposed to several natural and exotic insect preda- tors (mostly ants) under various experimental conditions, to ascertain the effectiveness of the repagula. In each experiment, for each ascalaphid species, two normal egg-masses and one egg-mass stripped of its repagula were mounted (on their respective twigs) in the vicinity of foraging ants, in such a manner that the bottom-most egg of each clump was positioned 16 cm above the ground. The behavioral details
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