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The numbers of Psyche issued during the past year were mailed on the following dates: Vol. 77, no. 4, December, 1970: October 21, 1971 Vol. 78, no. 1-2, March-June, 1971: December 30, 1971 Vol. 78, no. 3, September, 1971: February 23, 1972 PSYCHE A JOURNAL OF ENTOMOLOGY Vol. 78 March-June, 1971 No. i-2 CONTENTS The Reproductive Pattern of Dinoponera grandls Roger (Hymenoptera, Ponerinae) with Notes on the Ethology of the Species. Caryl P. Haskins and Paul A. Zahl 1 Fine Structure of the Thread Connections in the Orb Web of Araneus diadematus. Robert R. Jackson 12 Micratopus Casey in the United States (Coleoptera: Carabidae: Bembidiinae) . Thomas C. Barr, Jr. 32 Colonization of the Northeastern United States by Two Palearctic Moths (Lepidoptera : Tortricidae) . Jerry A. Powell and John M. Burns 38 A New Scenopinidae (Diptera) from Bermuda. L. P. Kelsey 49 Differentiation of the Carabid Antenna Cleaner. T. F. Hlavac 51 Notes of the Phasmatodea of the West Indies: Two New Genera. Carl Farr Moxey 67 The Male Genitalia of Blattaria. VI. Blaberidae: Oxyhaloinae. Louis M. Roth 84 Studies on the Biology of the Chrysopidae II. The Feeding Behavior of the Adult of Chrysopa carnea (Neuroptera) . Joseph K. Sheldon and Ellis G. MacLeod 107 "“S f I .! CAMBRIDGE ENTOMOLOGICAL CLUB Officers for 1970-71 President R. E. Silberglied, Harvard University Vice-President C. F. Money, Ilarvard University Secretary . C. S. H enry, Harvard University Treasurer F. M. Carpenter, Harvard University Executive Committee A. F. Newton, Jr., Harvard University T. F. H lavac, Harvard University EDITORIAL BOARD OF PSYCHE F. M. Carpenter (Editor), Fisher Professor of Natural History , Harvard University P. J. Darlington, Jr., Alexander Agassiz Professor of Zoology , H award U niversity W. L. Brown, Jr., Professor of Entomology , Cornell University; Associate in Entomology , Museum of Comparative Zoology E. 0. Wilson, Professor of Zoology , Harvard University H. W. Levi, Professor of Biology and Curator of Arachnology , Museum of Comparative Zoology FI. E. Evans, Alexander Agassiz Professor of Zoology , Harvard U niversity PSYCHE is published quarterly by the Cambridge Entomological Civil), the issues appearing in March, June, September and December. Subscription price, per year, payable in advance: $4.50 to Club members, $6.00 to all other subscribers. Single copies, $2.00. Checks and remittances should be addressed to Treasurer, Cambridge Ento- mological Club, 16 Divinity Avenue, Cambridge, Mass. 02138. Orders for missing numbers, notices of change of address, etc., should be sent to the Editorial Office of Psyche, 16 Divinity Ave., Cambridge, Mass. 02138. For previous volumes, see notice on inside back cover. IMPORTANT NOTICE TO CONTRIBUTORS Manuscripts intended for publication should be addressed to Professor E. M. Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass. 02138. Authors contributing articles over 4 printed pages in length may be required to bear a part of the extra expense, for additional pages. This expense will be that of typesetting only, which is about $13.50 per page. The actual cost of preparing cuts for all illustrations must be borne by contributors: the cost for full page plates from line drawings is ordinarily $12.00 each, and the full page half-tones, $20.00 each; smaller sizes in proportion. AUTHOR’S SEPARATES Reprints of articles may be secured by authors, if they are ordered at the time proofs are received for corrections. A statement of their cost will be furnished by the Editor on application. The December, 1970, Psyche (Vol. 77, no. 3) was mailed Octo- ber 21, 1971. The Lexington Press, Inc., Lexington, Massachusetts PSYCHE Vol. 78 March- June, 1971 No. 1-2 THE REPRODUCTIVE PATTERN OF DINOPONERA GRANDIS ROGER (HYMENOPTERA, PONERINAE) WITH NOTES ON THE ETHOLOGY OF THE SPECIES By Caryl P. Haskins1 and Paul A. Zahl2 Introduction The recent stimulating suggestions of Hamilton (1964#; 1964^; 1970) that social behavior in the Hymenoptera may have evolved at least in part as a consequence of particular conditions favoring kinship selection offered by the haplodiploid pattern of sex deter- mination in the higher Hymenoptera have, among other things, given special significance to the detailed study of breeding patterns in social members of that order. In this connection, we have for several years been investigating the breeding patterns, the effects of excessive inbreeding, and the modes of formation of new colonies, in a number of primitive ants. In this context, species in which either sex lacks functional wings at maturity take on special interest. Outside the Dorylinae and scattered groups of socially parasitic ants in other subfamilies, such forms, in which a typical mating-cum- dispersion flight is evidently impossible, are rather rare among the higher Formicidae. It is notable, however, and may be of a yet unidentified evolutionary significance, that marked brachyptery and even apterv in females are unusually evident in the two most generalized subfamilies of ants, the Myrmeciinae and the Ponerinae. Within the single genus Myrmecia, for example, forms in which the reproductive and colony-founding female is subapterous or even wingless and exhibits radically reduced thoracic musculature are by no means uncommon. Among the Ponerinae, as Wheeler pointed out some years ago (1933), wingless eratogynes replace the normal female forms in several genera, such as Acanthostichus , Eusphinctus , M egaponcm, Onychomyrmex, and Pletroctena. These ergatogynes, ’Carnegie Institution of Washington, Washington, D. C. ’National Geographic Society, Washington, D. C. Manuscript received by the editor May 5, 1971. T 2 Psyche [March-June Figure 1. Worker cocoon and pupae (upper), male cocoon and pupa (lower) Dinoponrra grandis, reared respectively in colony fragments A and B, described ( X 2). 1971] Haskins Zahl — Dinoponera 3 to be sure, are still morphologically distinguishable from the workers. In Leptogenys, sens, str ., however, in some species of Rhytidoponera , and in Diacamma , Streblognathus, and Dinoponera , no caste mor- phologically distinguishable from the worker has ever been reported, though normal males, in some cases evidently well adapted to secure outbreeding within the species, are the rule. A number of years ago Wheeler and Chapman (1922) described a male of a Philippine species of Diacamma in copula with an individual morphologically indistinguishable from a typical worker, suggesting the lack even of an identifiable ergatogyne in this species, the “workers” differing only in the presence or absence of a functional spermatheca and perhaps in the degree of ovariole development — a situation well known in several species of Rhytidoponera (Haskins and Whelden, 1965). It became of interest, therefore, to learn whether such workerlike individuals form the normal reproductive caste in Dino- ponera. That this situation, if real, could typify a rather ancient evolutionary condition is hinted by earlier findings of F. M. Car- penter. Carpenter suggested some years ago (1930) that a fairly close fossil relative of both Dinoponera and Streblognathus may be Archiponera wheeleri, described by him in 1930 from the Mio- cene Florissant shales of Colorado. The absence of described mor- phologically differentiable females in either Dinoponera or Stre- blognathus (1929; 1930) gave special emphasis to a search for such a caste among the fossils of Archiponera. No examples were dis- covered, though typical winged males were described. The observations to be presented confirm the production of workers by one or more wild-collected females of Dinoponera grandis, indis- tinguishable from workers in external morphology, in the artificial nest. M aterial The monotypic ponerine genus Dinoponera has been known since 1830, when its single species, D. grandis was described by Guerin from Para and Bahia, Brazil (1830). Carpenter noted (1930) that apparent morphological affinities of both it and the South African monotypic form Streblognathus aethiopicus to fossils of the Miocene Archiponera wheeleri in the Florissant shales could sug- gest that the two modern species are ancient relicts of an archaic ponerine complex which originally had a much wider distribution. The range of D. grandis given by Carlos Emery (1911) is “Middle American tropics as far as Paraguay,” and collecting localities for various described subspecies recorded up to that time 4 Psyche [March-June include Sao Paulo, Missiones, Espiritu Santo, Matto Grosso, Bahia, and Para in Brazil, as well as “Perou.” For the present study, the authors selected a single, restricted population of the typical form, not far from Para, which they have intermittently had under inde- pendent field observation since 1937. Some 30 “workers,” together with more than a dozen cocoons and larvae, were taken from a typical colony by one of us (PAZ) in December, 1969, and brought to Washington, D. C., where they were housed in observation nests and the recorded observations made over a period of somewhat more than a year.3 Methods and Observations The colony was approximately evenly divided, and the fractions housed in glass-and-plastic earth-containing Lubbock nests, 45.7 cm X 28.5 cm, and 3.0 cm in depth. Two of these were stacked in each of 2 aquaria of dimensions 61 cm X 29.0 cm X 22.5 cm, to serve as foraging arenas. These aquaria were covered at all times with 2 glass plates, with an aperture of 1.0 cm, through which was inserted the stem of a Weston Mirroband recording thermometer. Room temperature was kept constant at 750 F. Since the entrances to the Lubbock nests were kept open at all times, and soil was excavated and carried into the arena fairly continually by the ants, humidity usually approached saturation, A. Breeding Pattern The eggs of D. grandis are comparatively large (approximately 2.5 mm in length) and unusually elongate. They cohere in packets, usually of approximately 6 to 15 ova, and are assiduously tended by the workers. Indeed, the nurses spend much time in the nest at rest with such packets held in the mandibles. Shortly before hatching, single eggs are detached from the packet, licked and tended indi- vidually, and commonly deposited separately on the nest floor. Imme- diately after hatching the larvae are separately attended and fre- quently carried about. We believe (though it is not yet proved) that for the first, and possibly the second, instar they are fed in- gluvially by the nurses. Older larvae are given partially dissected arthropod prey in typical ponerine fashion, the fresh prey being commonly deposited on the ventral surface. The larvae develop rapidly through this stage. When about to spin, they are temporarily covered with earth in the typical ponerine manner. The cocoons of 3The authors wish to express their great appreciation to the National Geographic Society for its support of certain aspects of the 1969 field work in Brazil. 1971] Haskins & Zahl — Dinoponera 5 Figure 2. Workers (left); worker cocoon; male pupa; male cocoon Dinoponera grandis (X 2). 6 Psyche [March-June Date 12/6/69 2/7/70 2/15/70 2/27/70 3/3/70 3/18/70 3/20/70 6/11/70 7/15/70 8/28/70 9/4/70 9/16/70 9/20/70 9/25/70 10/8/70 10/10/70 10/11/70 10/12/70 10/14/70 10/15/70 11/3/70 11/29/70 12/10/70 12/29/70 12/30/70 1/2/71 1/6/71 1/6/71 1/7/71 1/9/71 1/10/71 1/13/71 1/27/71 1/28/71 2/2/71 2/4/71 2/5/71 l Records of Brood Rearing in Colony Group A Brood and Description First egg seen. Two half grown larvae; 7 small larvae. Two large larvae; 5-7 small larvae; about 6 eggs. Four large larvae; 5 medium larvae; 4 small larvae; eggs. Four large larvae; 4 medium larvae; eggs. Two cocoons; 4 large larvae; numerous medium-small larvae. Three cocoons; 4 large larvae; 1 small larva. Three cocoons; 3 medium larvae; eggs. Two cocoons opened. One contained a young, unpigmented, normal worker pupa, the second a semipupa. A further cocoon opened, revealing a partially pigmented, normal worker pupa. Three cocoons; 0 larvae; packet of about 10 eggs. First larva in new group hatched. Two additional larvae hatched. Fourth and fifth larvae hatched. Remaining pupa in cocoon died and extracted by workers. Nearly mature, normal worker pupa. First new cocoon spun. Two large larvae; 2 medium larvae; eggs. Second larva buried for spinning. Second cocoon spun. Meconium appeared in first cocoon. Meconium appeared in second cocoon. Third cocoon spun. Fourth larva buried for spinning. (The nest was inadvertently disturbed a few hours later. This larva was then prematurely disinterred by the workers and perished.) First cocoon hatched, eclosing nearly pigmented, normal young worker. Pupa in a second cocoon died and was extracted. A nearly mature worker. First larva hatched from new egg group. One large larva; 1 medium; 1 small. New larva hatched (1 large; 1 medium; 1 small). One large larva; 2 medium; 1 small. First larva banked for spinning (A.M.). First cocoon spun (P.M.). Fifth larva hatched. Sixth larva hatched. Meconium in first cocoon. Seventh larva hatched (1 cocoon; 2 medium, 4 small larvae). Second larva banked for spinning (P.M.). Second cocoon spun (2 large, 3 medium, 1 small larva; no eggs). Meconium in second cocoon. First new egg. Third larva banked for spinning. Second egg. Third cocoon spun. 1971] Haskins & Zahl — Dinoponera 7 2/8/71 2/1 Q/71 2/11/71 2/16/71 2/17/71 2/21/71 2/22/71 2/25/71 3/6/71 3/14/71 3/19/71 4/13/71- 4/14/71 One cocoon destroyed by nurses. Pupa not recovered. (Two (Two cocoons; 3 large larvae.) Fourth larva banked for spinning. Fourth cocoon spun (3 cocoons; 2 large larvae). Fifth larva banked for spinning. Fifth cocoon spun (4 cocoons; 1 large larva; no eggs). Sixth larva banked for spinning. Sixth cocoon spun (5 cocoons; 1 egg). Second cocoon rejected from nest. Opened, disclosing normal, apparently healthy worker pupa, entirely unpigmented. (Four cocoons, eggs.) Third cocoon rejected from nest. Opened, disclosing perfect, ap- parently vital worker pupa, with eyes fully pigmented. Fourth cocoon rejected from nest. Opened, disclosing perfect, apparently vital worker pupa, eyes fully pigmented, and body pigmentation well advanced. Fifth cocoon rejected from nest. Opened, disclosing perfect worker, with eyes fully pigmented, and body pigmentation well advanced. Sixth cocoon hatched, eclosing perfect worker of adult pigmen- tation. workers are large and robust (approximately 23.0 mm X 8.5 mm) and formed of a tough, dark brown silk. In eclosions of the imago that we have witnessed, attendant workers have assisted in opening the cocoon at the anterior pole, but it is possible that isolated pupae can emerge unassisted, as in some other Ponerinae. In the artificial nest, young workers have been almost fully pigmented at eclosion. It is likely that this is also the case under natural conditions, a situa- tion typical of some other members of the Tribe Ponerini. The two fragments of the collected colony were kept separate throughout the observations, and separate records of brood rearing were maintained. That for Group A is indicated in Table I. Thus from brood of this group, originally comprising 10 wild- collected “workers,” 15 cocoons of worker size and form were matured. The contents of 1 1 were definitely identified as worker pupae or adults. In the remaining 4 the cocoons were workerlike in form, but the contents could not be verified because of their early death or premature examination. The developmental periods recorded for 8 larvae followed from hatching to cocoon spinning were 24, 25, 27, 29, 31, 43, 44, and 44 days. The interval between the covering of a spinning larva with soil and the cleaning of the completed cocoon was 1 day for each of 6 individuals. The interval between the completion of the cocoon and the appearance of the meconial spot in 4 individuals was 4, 3, 4, and 5 days. Periods 8 Psyche [March-June recorded from cocoon spinning to pupal maturity for 2 individuals were respectively 50-51 and 52 days. As suggested by the data of Table 1, some ecological deficiency (perhaps too low a temperature) may have been responsible for the premature abandonment of all but 2 of the cocoons by the nurses, and may have prolonged times to eclosion abnormally in those that hatched. It was clearly evident that at least 1 adult fertile brood female, morphologically indistinguishable from a worker but capable of pro- ducing female progeny, had been included in the original wild col- lection The history of brood production in the second group of workers was quite different. Here, despite intensive care, egg production was poor, and the brood total low Only 2 cocoons were produced, both of male size and form (at about 20.0 mm X 6.0 mm, fairly reliably distinguishable by inspection from those of workers). One of these was opened artificially, disclosing a nearly mature male pupa. The second eclosed naturally, revealing a perfect male. No worker brood was produced. It thus appears that a fertilized ergatogyne was lacking in the second fraction of the colony B. Mating Pattern In Dinoponera , as in Streblognathus and Archponera, the males are decidedly smaller than the workers (see Figures 1, 2, and 3). They are much more lightly pigmented at maturity, and are relatively fragile. The compound eyes are large and the ocelli unusually prominent, conspicuously reflecting lew incident light. The wings are well developed, and adults once emerged from the nest fly actively. Within the parent colony, however, they are surprisingly inert. At least until full maturity they assume the pupal posture when disturbed and are carried by the workers as though they were brood, as the authors have frequently observed both under natural conditions and in the artificial nest. In the latter situation, males are often held in the mandibles of immobile workers as though they were brood, even when the colony is slightly stimulated. The males may well be night fliers of somewhat restricted range. We have not yet witnessed mating flights under natural conditions, nor deter- mined the precise mode of formation of new colonies. Note on the Ethology of D. grandis Ever since Henry Walter Bates (1892) almost eighty years ago described columns of D. grandis “marching through jungle thickts” the implication has been widely assumed and reiterated that the 1971] Haskins & Zahl — Dinoponera 9 Figure 3. Worker (left) ; male (right) of Dinoponera grandis, showing disparity in size (X 2). 10 Psyche [March-June species is a typical column termite-raider, foraging in the general pattern of T ermitopone in the New World or Megaponera in the Old. This conception has most recently been alluded to by Sudd (1967). All our observations, however, including those made both in the artificial nest and under natural conditions, seem contrary to this. Foraging workers of Dinoponera may indeed follow one another in tenuous, ill-defined columns. But all those that we have observed under natural conditions have been extremely loose forma- tions — so diffuse as hardly to merit the name. Moreover, we have never seen termite raiding under natural conditions. In the arti- ficial nest, the species proved a general and uncritical feeder on a wide range of arthropod prey, including the larvae and pupae of other ants when offered. Workers of Termes flavipes , when pre- sented in debris outside the nest, were indeed sought out, captured, and carried in: but with no detectably greater readiness than other insect prey. If Dinoponera is specialized to termite feeding at all, it is to a very slight degree. As with other members of the Ponerini, sugary substances are readily accepted — and, indeed, probably re- quired — • by the adults. Summary The failure to discover a morphologically distinct female caste among members of the archaic ponerine genera Dinoponera or Streblognathus, or in the fossil genus Archiponera, has long led to the suspicion that, as in Diacamma and species of Leptogenys and Rhytidoponera , such a caste may in fact be lacking and may be re- placed by a reproductive form morphologically very similar if not identical to the worker but physiologically and structurally capable of fertilization and the production of worker brood. This suspicion has now been experimentally verified in Dinoponera grandis in the artificial nest. Notes are appended on certain features of the breeding pattern and ethology of the species. References Bates, H. W. 1892. The Naturalist on the River Amazon. Ed. by Clodd, London. Carpenter, F. M. 1929. A Fossil Ant from the Lower Eocene (Wilcox) of Tennessee. J. Wash. Acad. Sci., 19: 300-301. Carpenter, F. M. 1930. The Fossil Ants of North America. Bull. Mus. Comp. Zool., Harvard Univ., LXX (1) : 27-29. Emery, Carlos. 1911. Genera Insectorum, 118me Fasicule (Hymenopt^ra) . 1971] Haskins & Zahl — Dinoponera 1 1 Guerin. In Duperrey, Voy. Coquille, Zool. Vol. 2 (2), p. 206 (1830). Cited in Emery, 1911. Hamilton, W. D. 1964a. The Genetical Evolution of Social Behavior. I. Jour. Theoretical Biology, 7: 1-16. Hamilton, W. D. 1964£. The Genetical Evolution of Social Behavior. II. Jour. Theoretical Biology, 7: 17-52. Hamilton, W. D. 1970. The Evolution of Social Behavior, as Illustrated by Social Insects. Unpublished essay. Haskins, C. P., and R. M. Whelden. 1965. Queenlessness, Worker Sibship, and Colony versus Population Structure in the Formicid Genus Rhyti- doponera. Psyche: 92: 87-112. Sudd, J. H. 1967. An Introduction to the Study of Ants. St. Martin’s Press, New York City, p. 78. Wheeler, W. M. 1933. Colony Founding Among Ants. Harvard Uni- versity Press, Cambridge, Mass. Wheeler, W. M., and J. W. Chapman. 1922. The Mating of Diacamma. Psyche , 29: 203-211. FINE STRUCTURE OF THREAD CONNECTIONS IN THE ORB WEB OF ARANEUS DIADEMATUS 1 By Robert R. Jackson2 Division of Research North Carolina Department of Mental Health Raleigh, North Carolina INTRODUCTION Web-building spiders are valuable subjects for the study of be- havior since the spider provides a record of much of its behavior through its web. Descriptions of the gross structure of the web and the behavior of the spider during its construction can be found in Savory (1952), Jacobi-Kleemann (1953), and Witt, Reed, and Peakall (1968). However, the gross structure is only part of the spider’s creation. The threads from which the web is constructed are only several microns thick. Consequently, the unaided human eye can determine basically only the position of the threads. Another part of the spider’s creation is visible only with magnifi- cation. This is the fine structure of the threads where they connect to one another. Rapidly produced and possessing remarkable strength ( DeWilde, 1943), thread connections are one of the more interesting accomplishments of the spider. However, little attention has been given to the fine structure of the web. Lehmensick and Kullmann (1957) and Friedrich and Langer (1969) examined spider silk by electron microscopy, but did not look at points of connection between threads. McCook (1889) and Nielsen (1931) have drawings of attachment disks for drag-lines made under low magnification. De- Wilde (1943) has a photomicrograph of a radius to frame connec- tion, and Comstock (1948) has a drawing from a photomicrograph of a spiral to radius connection. Robbins (Savory, 1952) has a photomicrograph of a drag-line attachment disk and one of sticky spiral to radius connections, both at low magnification. Also, there is a photomicrograph of a spiral to radius connection from the hori- This work was carried out in the laboratories of the North Carolina Department of Mental Health and was supported in part by a grant from Hoffman-LaRoche, Inc., Nutley, New Jersey, and by Grant Number GB- 6246X1 from the National Science Foundation to Peter N. Witt. The assist- ance of Dr. Peter N. Witt during all stages and Dr. Charles Walcott during the preparation of the manuscript of this work is gratefully acknowledged. 2Mailing address: Robert R. Jackson, Department of Zoology, University of California, Berkeley, California 94720. 12 1971] Jackson — Web of Araneus 13 zontal web of Cyrtophora citricola in Kullmann’s (1957) study of this peculiar spider. However, we have been able to find no detailed descriptions of thread connection fine structure. In the study reported in this paper, we investigated the fine structure of most of the types of thread connections which are found in the web of Aranens diadematus Clerck. Within the web there may be between 1000 and 1500 locations at which one thread is fastened to another thread (Fig. 1). Mooring threads fasten the web to some non-thread structure and may be continuous with some of the frame threads (Witt, et al. , 1968). There are frame thread to frame thread connections, forming Y’s with the stem of each Y being a mooring thread. Each radial thread is fastened to a frame thread. Occasionally, a radius will be in the form of a Y-structure at its peripheral end; the stem of the Y connects with the hub; and each arm, or secondary frame thread (Peters, 1939), fastens to either a frame thread or a radial thread. However, the majority of the connections within the web are the viscid or sticky spiral to radius connections in the trapping zone. Most of these are at points at which the sticky spiral thread extends from both sides of the radius. Occasionally, especially in the lower portion of the web, the sticky spiral thread meets the radius but does not continue on the other side. In the strengthening zone (Savory, 1952) or notched zone (McCook, 1889), non-sticky spiral threads are fastened to radial threads. The strengthening zone surrounds the apparently disorderly network of threads in the hub (McCook, 1889). Non-sticky spiral thread to radial thread connections are also present in the provisional spiral, a structure which is removed as the sticky spiral is added. Each of these types of connections will be described except for those at which a sticky spiral meets the radius but does not continue on the other side. METHODS All thread connections surveyed in this study came from webs of laboratory reared adult female cross spiders ( Araneus diadematus Clerck). Each spider was kept in a 51 x 51 X 9 cm aluminum frame with removable glass doors and was provided with one house fly twice per week, water daily, and controlled conditions of light and temperature (Witt, et al ., 1968). The age (3 to 4 months since emergence from the egg cocoon), weight (88 mg to 134 mg), and time between food consumption and web construction ( 1 to 8 days, with the majority at 1 day) were relatively constant for spiders used in this study; and within these ranges for these factors, no cor- 15 1971] Jackson — Web of Araneus relation was apparent between these factors and structural features of the connections. The day before the connections in a spider’s web were to be looked at, the glass doors were removed, and the old web was destroyed. The new web, built by the spider early the next morning, was treated as in Witt et al . (1968) except that the threads were not coated in any way. Afterwards several thread connections from the web were examined. In the web, threads are stretched tight into straight lines. When placed on a glass slide, threads with sticky globules often remained straight; but the globules were usually destroyed. Threads without globules curled and twisted when placed on a slide. To obtain a less distorted connection to study under a microscope, Permount mounting medium (Fisher Scientific Company) was streaked into a circle on a glass microscope slide, forming a basin. The slide was then placed against the web so that the threads surrounding the connection would stick to the Permount. Thus, the connection was suspended above the slide by the ridges of the circle of Permount. The threads were then burned loose from the outer edges of the circle by a surgical cauter, and the slide was taken away. The slide was always made on the same day that the web was built. Within a week of preparation of the slide, the connection was viewed under a Leitz Labolux D microscope; and negatives were made on Panatomic X sheet film using a Brinkmann Mark V camera with various objective lenses on the microscope. That structural Fig. 1. The vertical orb web of Araneus dladematus with labeled struc- tures. The scale, consisting of a weight suspended by cotton threads 20 mm apart, indicates size and vertical direction. Abbreviations: SS, loop of sticky spiral; SS-R, sticky spiral to radius connection; NS-R, non-sticky spiral to radius connection; R-F, radius to frame connection; FY, frame Y-structure; RY, radial Y-structure. Fig. 2a-d. Examples of sticky spiral (SS) to radius connections. In these photomicrographs, SS’s run vertical ; radii, horizontal. Radius is fastened to frame to right of page. Up and down on page corresponds to up and down, with respect to gravity, in web. a. Zero junction: points where SS joins radius on one side is directly across from where it joins at other side. Note sleeves on radius at both frame and hub side of SS. b. Smooth non-zero junction: there is space, smooth in appearance, along radius be- tween points at which SS joins each side of radius. Idiosyncrasies such as the doughnut-shaped structure in this SS-R made generalization about structure difficult, c. Non-zero junction with rough appearance, d. Note sleeves (S) on radius at frame side of SS and on SS, loose strands (L) on radius, and globules (G) on SS. 6 Psyche [March-June variations of the connection were not due to the amount of time which elapsed between slide preparation and exposure of the negative was shown by comparing photomicrographs with the slides after more time had elapsed. After several weeks to a year, essentially no changes in structure were discovered except for frequent disappear- ance and fusion of globules on the sticky spiral. Various measurements and counts were made from the negatives and prints. Since i [i was close to the limit of resolution on the photomicrograph, measurements in this range should be considered approximations. The thickness of a thread associated with a con- nection was usually measured at a distance of 150 /x from where the thread joined the other thread. In most cases, threads were fairly uniform in thickness at this distance. Closer to the other thread, thickness had a greater tendency to vary, and measurements were complicated by features of the connection. When the thickness of a thread at 150 /x was obviously not typical for that thread, as for example, when a sticky spiral had a globule on it at this point, the thickness was measured somewhat further from the connection. When strands were counted at a connection, “separate strands” were defined as strands with a visible space between them in the photomicrograph. However, sometimes an estimation of the number was used because some strands were hidden by others, and some were out of focus. The reader may refer when necessary to the following list, in alphabetical order, of abbreviations which will be used in this paper; FY, frame Y-structure; NS, loop of non-sticky spiral from the strengthening zone; NS-R, non-sticky spiral to radius connection from the strengthening zone; PS, loop of provisional spiral; PS-R, provisional spiral to radius connection; R-F, radius to frame connec- tion; RY, radial Y-structure; SS, loop of sticky spiral; SS-R, sticky spiral to radius connection. RESULTS The microscope reveals a wealth of structural detail at thread connections from which many component features can be abstracted, and the complexity and variability of which make description difficult. Some features of the structures will be pointed out in the following sections; and sample photomicrographs will be presented. Various descriptive statistics (percentages, means, coefficients of variation, and ranges) will be used to provide an impression of the prevalence of certain features and the central tendency and variability of meas- 1971] Jackson — W eb of Araneus 17 Table 1 Frequency of some structural features of sticky spiral to radius connections (SS-R’s). For each feature, 49 SS-R’s from 15 webs built by 8 different spiders were observed. Note that each SS-R had a sleeve on the radius at the frame side of the junction and that SS’S never had loose strands. Feature Percent of SS-R’s having the feature Zero Junction 22.4 Sleeve on Radius at Frame Side of Junction 100 Sleeve on Radius at Hub Side of Junction 46.9 Sleeve on SS 55-1 SS and Radius Both Single Stranded at Junction 36.7 Loose Strands Along Radius 35A Loose Strands Along SS 0 urements made on various features. We will draw attention to some of the questions suggested by these observations, and possible directions for further research will be indicated with hypotheses con- cerning function and origin of the structures. Sticky Spiral to Radius Connections: In reading the following section, the readers may refer to Table 1 and Table 2 for data pertaining to SS-R’s. We will define a “junction” as the space along a thread at which another thread is fastened to it. By “connection” we will mean i8 Psyche [March-June the junction plus the threads in the immediate vicinity of the junction. The point at which the SS joined one side of the radius was either directly across from where it joined the other side (zero-junction) (Fig. 2a) ; or there were varying amounts of space (the junction) along the radius between these points (non-zero junction) (Fig. 2b), causing the SS to have a disconnected appearance. In contrast to the SS’s, radii always appeared as continuous, nearly straight lines at SS-R’s. Junctions were smooth (Fig. 2b) or rough (Fig. 2c), with many gradations between. The smoother junctions were more prevalent. It would be interesting to know in what way, if any, the length and roughness of the junction are related to the functioning of the web. For example, do these features affect the strength of the con- nection or the transmission of vibrations along the radius? The radius at the frame side of all SS-R’s and, less frequently, the radius at the hub side of a SS was rougher in appearance and thicker adjacent to the junction than the same thread further away from the junction. These areas will be referred to as sleeves (Fig. 2d). For any given junction, the sleeve on the radius at the frame side was at least 25 jjl longer than the one on the radius at the hub side; and usually it was longer than the one on the SS. Perhaps threads are strengthened by sleeves, and the prevalence and length of sleeves are related to the magnitude of the tensions they must withstand in the web. Measurements of these tensions would be useful. At some SS-R’s, the SS joined the radius as a single strand at both sides of the junction (Fig. 2b and 2d). Other SS-R’s consisted of various numbers of strands (Fig. 2a, 2c, and 2e), often interconnected and variable in thickness. Perhaps the presence of many strands at a junction functions in the dispersal of stresses or to increase elasticity. If the presence or absence of non-zero junctions, sleeves on the radii at the hub side of junctions, sleeves on SS’s, and many strands at the connection improve the web in some way, why do some SS-R’s have these characteristics while others do not? Many SS-R’s had idiosyncrasies, the description of which will not be attempted here. The doughnut shaped structure in Fig 2b is an example. It does not seem likely that these structures have a specific function since any one of them occurred at only a single SS-R. They are probably artifacts of the forces acting on the connection during its formation. Thread connections, especially SS-R’s, are produced at a very rapid rate. The spider produces over a thousand Descriptive statistics for some structural features of sticky spiral to radius connections (SS-R’s). SS-R’s were rather variable with respect to these features, as indicated by the coefficients of variation. Note the differences in sleeve length on the radius at the different sides of the junction and the differences in thread thickness for the radius and SS. Thread thickness was measured at both sides of the junction. 1971] Jackson — Web of A raneus 19 •§ 2 E -2 3 32 Z 'a, GO W jo E 3 z -s c a .2 .2 C3 CJ .« ® £ 8 > U M-4 o 3 c .2 o -a +3 P* § o and FY’s. Feature Means Coefficient of Variation Range Length of Sleeve 74.8 {X 95 22 jti-180 ii Number of Strands at Junction 15-6 60 9-32 Thickness of Stem of Y 10.4 n 24 7 /a- 1 4 P Thickness of Single Stranded Arm of Y 00 bo “S 19 7 fi-IOfi 1971] Jackson — Web of Araneus 27 Fig. 5. Thread connections from hub. a. Notice loose thread (A), con- nections resembling NS-R’s (B) (Fig. 3), and compound connections (C). b. Junctions of unusual shapes. Also note globules (G), which resemble those on SS’s in Fig. 2. Fig. 6. Typical radius to frame connection. Radius, consisting of nu- merous strands, runs vertical; frame, horizontal. Frame can be seen to have sleeves at one side of radius and to consist of two strands at other side. Fig. 7. Typical frame Y-structure. Multi-strand frame fastened to single strand frame. 28 Psyche [March-June Fig. 8. Radial Y structure. Sleeves can be seen on stem and one arm of Y. Note that beyond sleeve on stem, radius is split into several strands. Fig. 9. a. Drag-line attachment disk with numerous strands of thread. b„ Higher magnification. Note globules (G). Fig. 10. Scanning electron micrograph of anterior spinneret showing nu- merous spools which open from piriform glands. Arrow indicates thread between 0.5 (J- — A ft in diameter, coming from spool .1200X- Photomicro- graph courtesy of Rainer Foelix, N. C. Department of Mental Health, Raleigh, N. C. 1971] Jackson — TVeb of Araneus 29 disks except that each thread is fastened to another thread rather than to a non-thread substrate. However, the number of strands (over 100, counting only those which touch the drag-line) at drag- line attachment disks (Fig. 9a) greatly exceeded the number at any connection from the web. Also, globules which, with the exception of one PS-R, were never observed on the minute strands of con- nections from the web, were found on some 0.5 /x — 1 [i thick strands at drag-line attachment disks (Fig. 9b). Are they involved in fastening the disks? What glands produce them? FORMATION OF JUNCTIONS How are threads fastened together at thread connections? By what mechanism is the junction formed? Within the glands, the silk is fluid. Eisner and Peakall (priv. comm.) have photomicrographs, taken under polarized light, of silk being pulled from the ampullate gland which indicate that the silk is already highly ordered at the spigot. Apparently the transforma- tion into a solid is at or previous to this point. Wilson (1969) hypothesized that the control valve, located inside the spinneret and behind the spigot, is the site of the transformation for the ampullate gland. One hypothesis for junction formation is that there is a cementing substance which fastens threads together. As the spinnerets touch the old threads, to which the new thread will be fastened, the cementing substance, from another gland, is added to the solid silk coming from the ampullate gland. If this hypothesis is true, then what is the cementing substance? Should we expect each type of connection to be equipped with the same type of cement? At SS-R’s, the cement is apparently not simply the viscid material from the aggregate glands. This conclusion is supported by two observations: 1) When SS-R’s were immersed in water, the globules on the SS were washed off; but the junction remained secure. 2) Artificial sticky spiral to frame connections and SS-R’s were pro- duced by placing a frame or radius over a Permount basin, then placing a SS from the same web across it. Never securely fastened, the SS at these connections could always be moved in any direction on the frame or radius. SS’s which cross but are not fastened to radii are common in the horizontal web of Uloborus diversus (Eber- hard, 1969), and we found them in webs of A. diadematus after the spider had ben administered a central stimulant (dextro-ampheta- 30 Psyche [March-June mine sulfate, ioo mg/kg). However, they do not normally occur in the vertical web of A. diadematus. Possibly the material from the piriform glands is a cement. Does it cement the drag-line to the substrate through fusing to both, or are the threads from the piriform glands themselves cemented at one end to the dragline and at the other to the substrate? Stretching may be responsible for solidification of fluid silk through causing filaments to slide past one another, assuming positions which allow for a greater degree of cross linkage (Lucas and Rudall, 1968). Another hypothesis of junction formation is that fluid silk for the new thread fuses with the old thread as it solidifies. As the spinnerets sit on the old thread, some fluid silk departs from the spigot of the ampullate gland (and the spigots and spools of any other gland involved). Solidification does not occur because there is insufficient stretching of the material. When the spinnerets are lifted and sufficient stretching occurs, the new silk solidifies and is fused to the old thread. More information will be necessary before we can choose between these hypotheses. Perhaps different mechanisms occur at different types of connections. Also, it may become necessary to somehow modify or combine these hypotheses. SUMMARY AND CONCLUSIONS The fine structure at thread connections in the web of Araneus diadematus was examined by photomicroscopy. Characteristic struc- tures were discovered for each connection type including presence of sleeves on threads, various lengths of junctions, various numbers of strands, and different distributions of globules. Possible functions and origins of structures were discussed. There was a discussion of two hypotheses related to connection formation: a) threads are cemented together, and b) new thread, while in a semi-solid state, fuses to old thread. This survey is only a modest beginning in the direction of study- ing the fine structure of thread connections. With a more extensive and systematic investigation, rigorous statistical comparisons could be made of different types of connections and of the same type of connection from different regions of the web. Structure could be compared for spiders of different age, weight, and species. Viewing the fine structure of connections has led to many interesting ques- tions. Some were mentioned in this paper; others have probably occurred to the reader. Hopefully, these will be pursued further in future studies. 1971] Jackson — Web of Araneus 31 References Cited Cowstock, J. H. 1940. The Spider Book. Comstock, Ithaca, N. Y. DeWilde, J. 1943. Some physical properties of the spinning threads of Aranea diademata. Arch, neerl. Physiol. 27: 118-132. Eberhard, W. 1969. Computer simulator of orb-web construction. Amer. Zool. 9: 229-238. Friedrich, V. L. and R. M. Lancer. 1969. Fine structure of cribellate spider silk. Amer. Zool. 9: 91-96. Jacobi-Kleemann, M. 1953. Uber die Lokomotion der Kreuzspinne Aranea diadema beim Netzbau (nach Filmanalysen) . Z. vergl. Physiol., 34: 606-654. Kullmann, E. 1957. Beobachtung des Netzbaues und Beitrage zur Biologie von Cyrtophora citricola Forskal (Araneae, Araneidae). Zool. Jahrb. Abt. Syst. 86: 181-220. Lehmensick, R. and E. Kullmann. 1957. Uber den Feinbau der Spinnfaden. In F. S. Sjostrand and J. Rhodin (ed.), Electron microscopy: Proc. of the Stockholm Con- ference, Sept. 1956. Academic Press, Inc., New York. Lucas, F. and K. M. Rudall. 1968. Extracellular fibrous proteins: The silks. Comprehensive Bio- chem. 26B: 475-558. McCook, H. 1889. American Spiders and their Spinningnvork. Vol. I, McCook, Academy of Natural Science, Philadelphia. Nielsen, R. 1931. The Biology of Spiders. Vol. II, Levin and Munksgaard, Copen- hagen. Peakall, D. B. 1969. Synthesis of silk, mechanism and location. Amer. Zool. 9: 71-80. Peters, H. M. 1939. Uber das Kreuzspinnennetz und seine Probleme. Naturwissen- schaften, 47: 776-786. Savory, T. H. 1952. The Spider’s IV eb. Frederick Warne and Co., Ltd., London. Wilson, R. S. 1969. Control of drag-line spinning in certain spiders. Amer. Zool. 9: 103-111. Witt, P. N., C. F. Reed and D. B. Peakall. 1968. A Spider’s Web. Springer Verlag, Berlin. MICRATOPUS CASEY IN THE UNITED STATES (COLEOPTERA: CARABIDAE : BEMBIDIINAE) By Thomas C. Barr, Jr. University of Kentucky, Lexington The late Rene Jeannel (e.g., 1946, p. 331) divided the bem- bidiine carabids, which he regarded as a subfamily Bembidiitae, into four tribes: Anillini, Limnastini, Tachyini, and Bembidiini. He included Micratopus Casey (1914, p. 42) and Limnastis Mots- chulsky in the Limnastini, although the former genus had previously formed the type of Casey’s tribe Micratopini. Ball (i960) retained Micratopini as a tribe of the Carabidae, but his “Bembidiini” is equivalent to the “Anillini”, “Tachyini”, and “Bembidiini” of Jeannel. Ball’s classificatory scheme thus implies that Limnastis and Micratopus are phylogenetically so remote that they merit a hier- archical status equivalent to that of all other bembidiines. Prior to his death Dr. Jeannel {in litt . ) indicated to me that he had used the name Limnastini rather than Casey’s older name Micratopini because he had not examined sufficient material of Micratopus and preferred the name of the tribe to be based on a well-known genus. After comparison of several species of Micratopus with several Old World species of Limnastis sent to me by Jeannel I see no reason not to apply the law of priority and accept Casey’s tribal name Micratopini, with Limnastini a synonym. However, Micratopini would become at best a subtribe in Bembidiini s. lat. in the classification of Ball (i960). Parenthetically it should be noted that Horologion speokoites Valentine (1932), an eyeless carabid known only from the unique holotype male taken in a West Virginia cave, has many features in common with the more primitive bembidiines (cf. Barr, 1969, p. 87). In my opinion it should be included within the subfamily Bembidiinae (or tribe Bembidiini s. lat.) close to and perhaps hier- archically coequal with the Anillini, rather than be relegated to the Psydrini, as was suggested by Valentine (1932) and actually done by Ball (i960). Micratopus Casey Casey, 1914, p. 42. Type species, M. fusciceps Casey, by original designa- tion. There appears to be a single species of Micratopus in the south- eastern United States, M. aenescens (LeConte). The species of the 32 1971] Barr — Micratopus 33 genus are distributed from the Gulf Coastal Plain and Interior Low Plateaus of the United States through Central America and the West Indies at least as far south as Panama and Trinidad. Collections are erratic; the largest series have come from berlesates of forest floor litter in swampy areas, or from black light traps near marshes and ponds. In Florida Mr. Harrison R. Steeves, Jr., obtained numerous M. aenescens from several localities by extracting them from leaf litter at the margins of lakes and swamps. In Putnam County, Tennessee, I found that I could regularly collect a few specimens from the rotting interior of an old chestnut log which lay on a small island in a wooded swamp. I took a single specimen by sifting debris in the damp, cool, sinkhole at the entrance to Bull Cave, Great Smoky Mountains National Park, Blount County, Tennessee. Casey (1914) states that his type series of M. jus deeps was taken by sifting leaf litter in a ravine near Vicksburg, Mississippi. The beetles are very active and crawl rapidly away when disturbed, but do not attempt flight. I have found them crawling out of the top of open Berlese funnels as often as falling through into the collecting dish. From all indications Micratopus spp. may be very abundant in favorable localities, but special techniques — litter processing or use of black light traps in swampy areas — seem indicated to obtain large series. These techniques have been seldom Fig. 1. Micratopus aenescens (LeConte), Putnam County, Tennessee. Length 2.4 mm. 34 Psyche [March-June employed by the average carabid collector, which probably explains the apparent rarity of species of this genus. Micratopus aenescens (LeConte) Figures i, 2 Blemus aenescens LeConte, 1848, p. 473. Type locality, “Georgia”, probably Habersham County. Type in Harvard Museum of Comparative Zoology. Tachys aenescens: Hayward, 1900, p. 197. Micratopus aenescens: Casey, 1914, p. 43. Micratopus fusciceps Casey, 1914, p. 43. Type locality, Vicksburg. Missis- sippi. Type in United States National Museum, new synonymy. Length 2. 1-2.5 mm- Dorsum of head dark chestnut; clypeus, labrum, antennae, pronotum and elytra brownish-piceous ; maxillary and labial palps, femora, coxae, and underparts of tibiae and tarsi flavotestaceous. Microsculpture of pronotum and elytra a very fine, anastomosing meshwork which imparts an opalescent cast; micro- sculpture of head isodiametric, open; integument more or less pubescent, least so on dorsum of head. Head with sides convergent, a little longer than wide, dorsum with minute punctures sparsely scattered and visible only at high magnification; labrum deeply emarginate, lateral lobes folded down over and partly obscuring mandibles, anterior margin bearing only four setae; mandibles short, thick, bifid at tip, without a seta in scrobe; mentum edentate, submentum with four (prebasilar) setae; maxillary and labial palps about as in Limnastis , penultimate seg- ments inflated, heavily pubescent, last segments minute, hyaline, glabrous; clypeus with a seta at each side, clypeolabral suture arcuate, convex forward ; frontal grooves shallow and very short, convergent ; eyes large and freely convex; only one pair of supraorbital setae. Pronotum about seven-tenths as long as wide, sides rounded and very feebly sinuate before the rounded, obtuse hind angles; base oblique behind hind angles, so that base is feebly “lobed” ; marginal gutter rather broad, hind angles not carinate ; disc sparsely pubescent. Elytra broad, depressed, subparallel, humeri prominent and apexes broadly truncate, length almost 1.5 times combined width; longitu- dinal striae moderately deep, distinct, interstriae each with a row of short pubescence; a single discal puncture on third stria near apex; umbilicate series with whips in punctures 1, 6, and 8. Antenna about six-tenths as long as body, segments II and III subequal and about six-tenths as long as segment IV; all segments more or less pubescent, but becoming more so from scape to segment 1971] Barr — Micratopus 35 Fig. 2. Aedeagus of Micratopus aenescens (LeConte), left lateral view, composite sketch from four preparations. Paramere shown detached. Length 0.30 mm., height 0.16 mm. IV. Male tarsi not differentiated from female; last abdominal sternite with only two marginal setae in both sexes. Protibia obliquely truncate near apex. Aedeagus (Figure 2) highly modified, trapezoidal, short, thick, and laterally compressed, about 0.16 X 0.30 mm. long; right side with a loosely attached lamina projecting a short distance upward and forward and articulating with a single slender (left?) paramere bearing two seatae at its apex; basal orifice a simple slot with no evident lobes; aedeagus walls thin, hyaline, with a thin sclerotized strip antero-dorsally and another, vertical strip at apex on left side ; internal sac with six rather complex sclerites : 1 ) basoventral : scroll-shaped, bifurcated, with minute spines internally; (2) dorsal: a slender, horizontally oriented, arcuate spicule; 3) basal: a slender, vertically oriented, arcuate spicule; 4) first medial: broad, bilobed, with a dorsal hook and ventral projection; 5) second medial: slender, bilobed, dorsal lobe bifurcate; and 6) apical: a blade-like hook. Measurements of a male from Gaits Landing, Cherokee County, Georgia (T. Barr, private collection) : total length 2.44 mm., head 0.48 mm. long X 0.44 mm. wide, pronotum 0.50 mm. long X 0.68 mm. wide, elytra 1.46 mm. long X 1.01 mm. wide (com- bined), antenna 1.46 mm. long, aedeagus 0.30 mm. long. 36 Psyche [March-June Geographic Distribution : — I have seen this species from the fol- lowing localities; abbreviations are USNM: United States National Museum; MCZ: Museum of Comparative Zoology, Harvard Uni- versity; and TCB: my private collection. Alabama: Selma (USNM). Arkansas: Carlisle (MCZ); “Ark.” (Hayward, MCZ). Florida: Enterprise (USNM); Tampa (USNM) ; numerous specimens in Berlese samples from Alachua, Baker, Dade, Highlands, and Seminole counties (TCB). Georgia: “Georgia”, probably Habersham County (LeConte, MCZ); Galt’s Landing, Cherokee County (TCB). Indiana: 5 miles west of Hardinsburg (USNM). Louisiana: Bay Sara (USNM); Houma (MCZ); Opelousas (Hayward, MCZ); Tal- lulah (USNM). Mississippi: Vicksburg (Casey, USNM). south CAROLINA: “S.C.” (MCZ). TENNESSEE: Bugger Swamp, near Cookeville, Putnam County (TCB) ; Bull Cave, Blount County (TCB). The range of M. aenescens is thus outlined as predominantly the Gulf Coastal Plain, with significant penetration northward into the Interior Low Plateaus (to Hardinsburg, Indiana). Leng (1920, p. 54) lists the species from “N.C.”, and it must certainly occur in Kentucky and possibly in southeastern Virginia. The apparent absence of Micratopus from southeastern Texas is probably a col- lecting accident. Discussion : — The species of Micratopus are differentiated with some difficulty, primarily because the range of variation within a species is difficult to determine with scattered material. Most of them, like M. aenescens , are probably rather widely distributed. The aedeagi resemble that of aenescens but are not easy to extract from dried specimens and are usually difficult or impossible to interpret unless several preparations are made. The accompanying figure of the aedeagus of M. aenescens (Fig. 2) was made with camera lucida but represents a composite of four aedeagi extracted from specimens at the same locality. I believe that the six minute sclerites of the internal sac, described in some detail for aenescens , have homologues in the Mexican and Central American species as well as those of the West Indies, and may prove to be of value in a diagnostic sense when the genus as a whole is investigated. The complex aedeagal structure presumably represents an aberrant and peculiar specialization. As such, it is of little help in determining the position of Micratopus within a bembidiine classificatory scheme. The external resemblances to Limnastis, however, leave little doubt that the two genera are closely related. Common features include 1971] Bar r — Micratopus 37 the deeply cleft labrum, the single pair of supraorbital punctures, the form of the maxillary and labial palps, the loosely aggregated umbilicate series of 9 punctures, the subtruncate elytral apexes, the absence of both the apical recurrent groove and a twisted epipleural fold, and the testaceous color. In view of the radically modified aedeagus of Micratopus , however, I prefer to retain both genera. Literature Cited Ball, George E. 1960. Carabidae, pp. 55-181, in Arnett, Ross H., Jr.: The beetles of the United States. Washington: Catholic Univ. America Press. Barr, Thomas C., Jr. 1969. Evolution of the Carabidae (Coleoptera) in the southern Ap- palachians, pp. 67-92, in Holt, Perry C., ed.: The distributional history of the biota of the southern Appalachians. Virginia Poly- tech. Inst., Blacksburg, Res. Mon. 1, 295 pp. Casey, Thomas L. 1914. Memoirs on the Coleoptera, V. Lancaster, Pennsylvania. Hayward, Roland 1900. A study of the species of Tachys of boreal America. Trans. American Entomol. Soc., 2 6 (1899): 191-238, pi. 6. Jeannel, Rene 1946. Coleopteres carabiques de la region malgache (premiere partie). Mus. Nat. Hist. Nat., Paris, 372 pp. Le Conte, John L. 1848. A descriptive catalogue of the geodephagous Coleoptera inhabit- ing the United States east of the Rocky Mountains. Ann. Lyc. Nat. Hist. New York, 4: 173-233, 334-474 (pagination error). Leng, Charles W. 1920. Catalogue of the Coleoptera of America, north of Mexico. Mount Vernon, New York: 470 pp. Valentine, J. Manson 1932. Horologion , a new genus of cave bettles (fam. Carabidae). Ann. Entomol. Soc. America, 25: pp. 1-8, 2 pis. COLONIZATION OF THE NORTHEASTERN UNITED STATES BY TWO PALEARCTIC MOTHS (LEPIDOPTERA: TORTRICIDAE) By Jerry A. Powell1 and John M. Burns2 Thirty years ago, two Palearctic tortricids, Croesia forskaleana (Linnaeus) and Clepsis unifasciana (Duponchel), were recorded in the Nearctic from Long Island, New York (Klots 1941). Almost no information on their progress has been published, but specimens subsequently collected on another island and at several points on the mainland indicate that these immigrants are spreading. Because rapid range changes are potentially useful in analyzing microevolu- tion (see, e.g., Baker and Stebbins 1965; Burns 1966) — the more so as they are accurately documented — • we attempt here to put the scene together, to interest other workers in future developments, and to encourage accelerated deposition of specimens in institutional collections. Although numerous lepidopteran and other insect species have reached the United States from the Old World and become estab- lished (e.g., Popham and Hall 1958), many came so early, or were so unspectacular after arriving, that their American history is obscure. The rare detailed accounts of spread deal with species such as the gypsy moth, Porthetria dispar (Linnaeus) (Corliss 1952), and the spotted alfalfa aphid, Therioaphis maculata (Buckton) (Smith 1959), which are conspicuous or economically important (or both). A few introductions of Microlepidoptera have been fairly well chronicled, such as that of the tortricid Cnephasia longana (Haworth) on the Pacific Coast (Powell 1 964*2) ; but most have not. At worst, as in California populations of the fungus-eating Oinophila v-flava (Haworth), a long gap in the temporal record, together with complex ecological and distributional data from the present, make an unequivocal choice between native and alien status impossible (Powell 1964^; Lawrence and Powell 1969). The crucifer-eating diamondback moth, Plutella maculipennis (Curtis), and various cosmopolitan household pests like the Indian meal moth, Plodia inter punctella (Hiibner), and the clothes moth, Tineola biselliella (Hummel), have been in North America so long that neither their beachheads nor their invasion patterns are known. Even 1University of California, Berkeley. 2Museum of Comparative Zoology, Harvard University. Manuscript received by the editor April 29, 1971. 38 1971] Powell & Burns — - Palearctic Moths 39 Fig. 1. Croesia forskaleana, male, dorsal view. New Market, Middle- sex County, New Jersey, VI-17-1968. Fig. 2. Croesia forskaleana, female, dorsal view. Palearctic. Fig. 3. Clepsis unifasciana, male, dorsal view. Palearctic. Fig. 4. Clepsis unifasciana, female, dorsal view. Palearctic. some rather recently introduced species have been so overlooked or ignored that only a vague rendering of their Nearctic history is possible. Three parallel cases involve Palearctic moths, the tortricids Acleris variegana ( Schiffermiiller) and Batodes angustiorana (Haworth) (Powell 1964#) and the oecophorid Borkhausenia (Batia) lunaris (Haworth) (Powell 1964c) : each inhabits parts of the Pacific Northwest and central coastal California without revealing whether it entered North America once and in the north, or once but to the south, or twice. Often there are several records of an alien species from about the time it was first noticed and a relative or absolute dearth of them for a long period thereafter. Even in so well sampled a group as butter- flies, the Palearctic hesperiid Thymelicus lineola (Ochsenheimer) — > which can quickly attain enormous population densities in newly colonized areas — was discovered at one locality in North America in 1910 and observed there for five years but was rarely recorded from anywhere for the next thirty years or more, in which it must have spread far in all directions from its origin (Burns 1966). For the two tortricids treated here, records are also intermittent in time and space. But the gaps are comparatively short and may not seriously hinder efforts at reconstructing New World movements of these moths. 40 Psyche [March-Junc Croesia forskaleana (Linnaeus) Phalaena Tortrix forskaleana Linne, 1758, Syst. Natur., ed. 10, p. 531. Argyrotoxa forskaleana: Klots, 1941, Bull. Brooklyn Entomol. Soc., 36: 126; Beckwith, 1962, Sci. Tree Topics, 2(9): 15. Croesia forskaleana: Obraztsov, 1956, Tijdschr. v. Entomol., 99: 127 (synonymy) ; MacKay, 1962, Canadian Entomol., Suppl. 28: 10 (larva). Diagnosis. — The ochreous yellow forewings of this species differ from the lemon yellow ones of native Nearctic species of Croesia . The delicate, reddish reticulation and the transverse blackish mark — which varies from a thin, median line, outwardly angulate in the cell, to a broad blotch on the mid dorsum — also distinguish C. forskaleana (figs. I and 2). Croesia semipurpurana (Kearfott) and C. curvalana (Kearfott) are native species that sometimes show blackish dorsal marking, but in them it is expressed as a broad blotch in the tornal region. All native species of Croesia have curving bands of shining rosaceous or lavender across the forewing, while C. forskaleana does not. Male and female genitalia of C. forskaleana are figured by Pierce and Metcalfe (1922). Geographic distribution. — Croesia forskaleana ranges widely in the Old World, from Great Britain and central and southern con- tinental Europe to the Caucasus (Meyrick 1895; Obraztsov 1956). For the United States, Klots (1941) gave a 1939 record from western Long Island, New York, and cited W. T. M. Forbes as authority for specimens dating back to 1934 from the northeastern tip of that island. Our data, which include an earlier record (1932) for the latter area, suggest a sequence of more or less steady spread. Croesia forskaleana was established in eastern Long Island by the early 1930’s and at the western end of the island and adjacent main- land New York (Westchester County) by the late 1930’s. After two more decades — in which there were no mainland records beyond the New York City area — • it was collected in westcentral Connecti- cut (1959), central Connecticut (1962), and east coastal and central New Jersey (1962) (fig. 5). Together, these records point to a multidirectional expansion of the moth on the mainland. Croesia forskaleana was found on a second island, Martha’s Vine- yard, Massachusetts, in 1944. It probably got there at about that time because it did not turn up in the course of a long survey of the Lepidoptera of Nantucket and Martha’s Vineyard islands (Jones and Kimball 1943), which was “completed” in 1942. Several points attest to the exhaustiveness of that survey: “. . . the Marthas Vineyard records are based [principally] upon the observations and 1971] Powell & Burns — Palearctic Moths 4i Fig. 5. Spatial distribution of Croesia forskaleana in North America. Earliest known year of occurrence at each locality is given. (The map shows the northeastern United States from Massachusetts to New Jersey.) collections of the author [F. M. Jones], beginning in 1913, con- tinued in 1917, 1921, 1925, and thereafter each year from 1927 to 1942. For twelve consecutive years a light-trap was operated all night, on all nights not too stormy, and usually from late June to early or mid-September” (Jones and Kimball 1943: 25). It was F. M. Jones himself who collected this species on Martha’s Vineyard, not only in 1944 but also in 1949. Negative evidence that we have accumulated from adjacent parts of the mainland suggests that Martha’s Vineyard was colonized via air movement directly from Long Island — • an overwater distance of as little as 60 miles. Biology. — Various European workers have stated that larvae of C. forskaleana feed on maple (Acer). Ford (1949: 56, 214, 218) specifically gave A. campestre and A. pseudoplatanus as foodplants 42 Psyche [March-June in England. Beckwith (1962) recorded a native American maple, A. saccharum, as a host in southwestern Connecticut; and MacKay (1962) examined larvae collected from maple on Long Island, New York. The larvae are generally described as leaf rollers. Besides Acer, Kennel (1910: 170) mentioned Rosa centifolia as a foodplant of C. forskaleana; but, because a related moth of similar appearance — C. bergmanniana (Linnaeus) — is known to be a Rosa feeder, this record may stem from a misidentification. Taking Kennel almost verbatim (but without citing him at that point), Swatschek (1958: 72) uncritically repeated Kennel’s Rosa centi- folia record. MacKay (1962: 10), citing Swatschek (1958), then simply included “Rosa” in a list of three C. forskaleana foodplants. To Rosa and maple, MacKay — citing Ford (1949) — -added “syca- more,” which in North America is usually taken to mean PI at anus, but which Ford and other Englishmen habitually apply to Acer pseudoplatanus! The literature would appear to be evolving in the direction of indiscriminate polyphagy. In Europe larvae are said to occur in May and June, pupae in June and July, and adults from the end of June to mid August (Meyrick 1895; Kennel 1910; Ford 1949). Our early and late records for colonizing adults are June 1 and August 1, with most records falling between June 19 and July 17. Apparently no detailed studies have been made, but it is likely that winter is passed by eggs in diapause, as is true of Croesia albi- co/nana (Clemens) (Powell 1964#). This supposition is supported by the fact that females taken in Connecticut and New York have dirt particles in the ovipositor setae. Presumably these particles are spread onto the eggs at oviposition. This behavior, which is accompanied by well developed structural modification in certain cnephasiine Tortricinae, is also thought to occur in a few other Tortricini — such as Acleris foliana (Walsingham) — all of which hibernate in the egg stage (Powell 1964#). Debris transfer during oviposition has not been recorded for Croesia. Material examined. — Connecticut: Middlesex Co., Middle- town, 2$ VII-7 and 8-62, 1$ VIII-1-62, 4d\ 2$ VII-15 to 19-63 (J. M. Burns). New Haven Co., Waterbury, 1 cf VI-15-59 (C. W. O’Brien). Massachusetts: Dukes Co., Martha’s Vineyard, 1 cf VII-23-44, 2 cf VI-29 and 30-49 (F. M. Jones), new jersey: Burlington Co., Ft. Dix, near Wrightstown, 1 cf VI-24 to VII-1-62 (light trap). Middlesex Co., New Market, 1 cf VI-17-68 (black- light trap). Monmouth Co., Ft. Monmouth, near Eatontown, 1 cf VII-9 to 15-62 (light trap), new York: Nassau Co., Sea 1971] Powell & Burns — Palearctic Moths 43 Fig. 6. Spatial distribution of Clcpsis unifasciana in North America. Earliest known year of occurrence at each locality is given. (The map shows the northeastern United States from Massachusetts to New Jersey.) Cliff, 2? VII-5-66 (A. Diakonoff and J. Powell); Valley Stream, 2cf VII-1-39 (A. G. Richards, Jr.). Orange Co., West Point, 1 cf VI-16-64 (B. Mather). Suffolk Co., Eastport, 1 cf VI-1-41 (D. Raynor); Mattituck, 1 cf VI-16-32, 1$ VI-30-37, 1 cf VII-8-46 (R. Latham); Orient, 1 cf VI-15-34, I? VII-2-35, 1 cf VI-18-36, 1 cf VII-30-36, 1$ VII-6-47 (R. Latham); Tucka- hoe, 1 cf VII-11-39 (collector unknown). Westchester Co., Mt. Vernon, 1 cf VII-20-38 (collector unknown); Ft. Slocum, New Rochelle, 20cf, 1$ VI-25 to VII-2-62 (light trap); Pelham, 3 cf , 4$ VII-17-50, 28 cf, 25$ VI-19 to VII-22-54, 1 cf VII-f-59 (A. B. Klots). County undetermined. Little Neck, VI-23-57* 1 cf VI- 28-58 (J. K. Terres) ; Long Island, 1 cf VI-10-59 (F. J. Davis); New York City, 1 cf VII-3-59 (T. Glidaspow). 44 Psyche [March-June Clepsis unifasciana (Duponchel)3 Tortrix unifasciana Duponchel, 1843, Hist. Natur. Lepid. France, Suppl. 4: 135; Klots, 1941, Bull. Brooklyn Entomol. Soc., 36: 12 6. Clepsis unifasciana: Obraztsov, 1955, Tijdschr. v. Entomol., 98: 215 (synonymy) . Diagnosis. — Clepsis unifasciana most closely resembles C. peritana (Clemens) and C. vires cana (Clemens) among Nearctic species but has a rusty or reddish brown tint. From C. peritana it differs also in having a well developed costal fold in the male and a less distinctly marked forewing; from C. vires cana,, in having a longer costal fold, no strongly contrasting outer costal spot, and a dark gray (rather than pale) hindwing (figs. 3 and 4). With their forewing patterns usually obscure, females of all three species look more similar than the males; but the lack of any differentiation of the outer costal spot and the rust-orange coloration of the forewing distinguish C. unifasciana females. The male genitalia of C. unifasciana are at once set apart by the elongate, curving, flattened setae of the medial face of the valva (see Obraztsov 1954; Pierce and Metcalfe 1922). The female genitalia have an elongate sclerotized band in the ductus bursae, somewhat as in Archips argyrospilus (Walker), and are unlike C. peritana and C. virescana. Geographic distribution. — Clepsis unifasciana is widespread in the Palearctic, having been recorded from the British Isles, main- land Europe, northwest Africa, Turkey, Syria, and the Amur and southern Ussuri region of the eastern Soviet Union (Meyrick 1895; Kennel 1910; Obraztsov 1955). In the Nearctic we know of only four localities. Klots’ (1941) record was based on specimens taken at Valley Stream, New York, in western Long Island in 1939. Our latest record comes from this same general area. Klots was next to collect the species, 1 1 years later, immediately north of New York City on the mainland. After another 13 years, C. unifasciana was found in central Connecticut (fig. 6). Biology. — In Europe this species feeds on privet, Ligustrum vulgare (Meyrick 1895; Kennel 1910) ; and at Valley Stream, New York, A. G. Richards, Jr., found adults of C. unifasciana excessively 3Bradley and Martin (1956) exhumed the name consimilana Hiibner, 1822, for this species and relegated unifasciana to synonymy. The Hiibner name had been treated as of doubtful status by Obraztsov (1955) and others, presumably because there is no type; Bradley and Martin gave no explanation for their reversal. 1971] Powell & Burns — Palearctic Moths 45 common on a privet hedge (Klots 1941). Ford (1949), citing Wakely, also listed ivy ( Hedera helix) and apple ( Pyrus malus ) as foodplants in England. According to Kennel (after Disque) and Ford, larvae begin eating in summer, overwinter, and then complete their growth in spring, pupating in June; and adults fly in June and July. American records of adults range from May 30 to July 22, with most occurring between June 19 and July 5. Material examined. — Connecticut: Middlesex Co., Middle- town, 1 cf VII-18-63 (J. M. Burns). NEW YORK: Nassau Co., Sea Cliff, 1 1 cf , 7? VII-5-66 (A. Diakonoff and J. Powell) ; Valley Stream, 5cf VII-1-39 (A. G. Richards, Jr.). Westchester Co., Pelham, 1 cf VII-17-50, 1$ VII-2-53, 36 cf, 18$ V-30 to VII-22-54, icf VI-23-56, icf VI-f-59 (A. B. Klots). Discussion Sporadic data indicate that, subsequent to their initial discovery on Long Island, New York, in the 1930’s, Croesia forskaleana and Clepsis unifasciana reached the adjacent mainland and spread con- siderably. Indeed, Croesia forskaleana not only fanned out in all possible directions on the mainland but also apparently jumped from Long Island to Martha’s Vineyard, an island in Massachusetts about 60 miles eastnortheast of Long Island. This jump is not surprising because it is known that major air masses often move up the Atlantic Coast; that various southern species of Lepidoptera have, at one time and another, briefly attained Martha’s Vineyard and Nantucket; and that many moths have permanently settled these islands without human assistance (Jones and Kimball 1943). Although both immigrant tortricids are thriving in some places, Croesia forskaleana seems to be the better colonizer (cf. figs. 5 and 6). It may have had a head start: our earliest record of it (1932) precedes the earliest record of Clepsis unifasciana (1939) by seven years. On the mainland, Croesia forskaleana was collected first just north of New York City in 1938. In 1950 both species were taken in that area at Pelham — the only mainland locality being sampled even intermittently at that time — and in 1954 both were extremely common there. Both moths are univoltine; and, to judge from their broad Palearctic ranges, both (especially Clepsis unifasciana ) must be ecologically tolerant at temperate latitudes. Croesia forskaleana is a maple (Acer) eater and Clepsis unifasciana is (at least primarily) a privet ( Ligustrum ) eater. Foodplant availability, then, should not notably delay the northeastern invasion of either species. Yet, on an average, the maple-eating Croesia 46 Psyche [March-June forskaleana may be able to cross interurban zones a little more easily than Clepsis unifasciana because several maples are important in the native flora and abundant in rural situations (as well as in cities), whereas all privet is introduced and the majority is planted in yards and kept grounds (albeit a substantial amount has escaped and become established in open woods, in thickets, and along roadsides). The picture of colonization is sketchy, owing in large measure to a paucity of collectors in the Northeast who are interested in local Microlepidoptera. Ironically, however, Powell’s Law — which states that “no systematic entomologist voluntarily works on insects that occur within 1,000 miles of his home laboratory” (Munroe 1969) — has helped as well as hurt : mainland spread has been documented chiefly by workers who left California and sampled northeastern tortricids in our behalf. Since this activity began only in the late 1950’s and was casual, the long lag in appearance of peripheral records beyond the immediate vicinity of New York City, and even the apparent distributional limits themselves, may be artifacts. One or both species may have expanded in New England and states to the south earlier or farther than our records show. Both moths are attracted to lights, sometimes in large numbers, and both are adapted to urban situations. Therefore it would not seem too idealistic to hope that collectors residing in the megalopolis — particularly at its northern and southern ends in the areas of Boston and Washington — ■ might watch for these tortricids and sharpen our view of their expanding perimeters. Acknowledgements We owe much of our information to two individuals who deliber- ately gathered northeastern Microlepidoptera for us. C. W. O’Brien (Texas Tech University, Lubbock), while a student at the Uni- versity of California, Berkeley, made a collection in Connecticut. P. A. Opler (Organization for Tropical Studies, San Jose, Costa Rica) accumulated a large number of light-trap samples from Massachusetts, Rhode Island, New York, and New Jersey as a by-product of work with the U. S. Army in 1962. These collections, together with those we made, are deposited in the California Insect Survey, University of California, Berkeley, and in the Museum of Comparative Zoology, Harvard University, Cambridge. C. P. Kimball (Barnstable, Massachusetts) responded to queries on the occurrence of the two species in the area of Cape Cod and adjacent islands. The following persons permitted use of material 1971] Powell & Burns — Palearctic Moths 47 in institutional and private collections in their care: A. B. Klots and F. H. Rindge (American Museum of Natural History, New York) ; Bryant Mather (Clinton, Mississippi) ; and R. W. Hodges (U. S. National Museum, Washington). Literature Cited Baker, H. G. and G. L. Stebbins, eds. 1965. The genetics of colonizing species. Academic Press, New York. 588 p. Beckwith, R. C. 1962. A new tortricid on maple in Connecticut. Sci. Tree Topics 2(9) : 15. Bradley, J. D. and E. L. Martin 1956. An illustrated list of the British Tortricidae. Part 1. Tortricinae and Sparganothinae. Entomol. Gazette 7: 151-156. Burns, J. M. 1966. Expanding distribution and evolutionary potential of Thymelicus lineola (Lepidoptera : Hesperiidae) , an introduced skipper, with special reference to its appearance in British Columbia. Canadian Entomol. 98: 859-866. Corliss, J. M. 1952. The gypsy moth, p. 694-698. In Insects. U. S. Dept. Agr. Year- book of Agr., 1952. Ford, L. T. 1949. A guide to the smaller British Lepidoptera. South London Entomol. and Natur. Hist. Soc., London. 230 p. Jones, F. M. and C. P. Kimball 1943. The Lepidoptera of Nantucket and Marthas Vineyard Islands, Massachusetts. Publ. Nantucket Maria Mitchell Assoc. 4: 1-217. Kennel, J. 1908-21. Die Palaearktischen Tortriciden. Zoologica 21 (part 54) : 1- 742. Klots, A. B. 1941. Two European Tortricidae (Lepidoptera) not hitherto recorded from North America. Bull. Brooklyn Entomol. Soc. 36: 126-127. Lawrence, J. F. and J. A. Powell 1969. Host relationships in North American fungus-feeding moths (Oecophoridae, Oinophilidae, Tineidae). Bull. Mus. Comp. Zool. 138: 29-51. MacKay, M. R. 1962. Larvae of the North American Tortricinae (Lepidoptera: Tor- tricidae). Canadian Entomol., Suppl. 28: 1-182. Meyrick, E. 1895. A handbook of British Lepidoptera. Macmillan and Co., London. 843 p. Munroe, E. 1969. Insects of Ontario: geographical distribution and postglacial origin. Proc. Entomol. Soc. Ontario 99: 43-50. 48 Psyche [March-June Obraztsov, N. S. 1954. Die Gattungen der Palaearktischen Tortricidae. I. Allgemeine Aufteilung der Familie und die Unterfamilien Tortricinae und Sparganothinae. Tijdschr. v. Entomol. 97: 141-231. 1955. Ibid., 1. Fortsetzung. Tijdschr. v. Entomol. 98: 147-228. 1956. Ibid., 2. Fortsetzung. Tijdschr. v. Entomol. 99: 107-154. Pierce, F. N. and J. W. Metcalfe 1922. The genitalia of the group Tortricidae of the Lepidoptera of the British Islands. Publ. by the authors, Liverpool, England. 101 p. Popham, W. L. and D. G. Hall 1958. Insect eradication programs. Ann. Rev. Entomol. 3: 335-354. Powell, J. A. 1964 a. Biological and taxonomic studies on tortricine moths, with refer- ence to the species in California. Univ. Calif. Publ. Entomol. 32: 1-317. 1964 b. Occurrence in California of Oinophila v-flava, a moth probably introduced from Europe (Lepidoptera: Tineoidea). Pan-Pacific Entomol. 40: 155-157. 1964c. Two scavenger moths of the genus Borkhausenia introduced from Europe to the West Coast of North America (Lepidoptera: Oecophoridae) . Pan-Pacific Entomol. 40: 218-221. Smith, R. F. 1959. The spread of the spotted alfalfa aphid, Therioaphis maculata (Buckton), in California. Hilgardia 28: 647-685. Swatschek, B. 1958. Die Larvalsystematik der Wickler (Tortricidae und Carpo- sinidae). Akademie-Verlag, Berlin, Abhandl. Larvalsystematik Insekten 3 : 1-269. A NEW SCENOPINIDAE (DIPTERA) FROM BERMUDA1 By L. P. Kelsey2 While examining the Scenopinidae in the collection of the Museum of Comparative Zoology, I noted a specimen collected in Bermuda. As there are relatively few records of Scenopinids from islands this specimen was retained for more detailed study and was found to be a new species in the Velutinus Group as evidenced by the narrowed bursal cavity. This species would key to Scenopinus nubilipes Say in the keys to the Nearctic females (Kelsey, 1969) but may be separated from that species on the basis of the bursal cavity, con- formation of the 8th sternum and the markings on the frons. In the accompanying illustrations scale marks equal one half millimeter, the shorter applies to the head and wings, the longer to the terminalia. Figure 1. Scenopinus bermudaensis n. sp. 2 ; a. wing; b. c. lateral and frontal aspects of head; d. enlarged detail of antennae; e. f. lateral and ventral aspects of 8th and 9th segments, missing portions outlined by dashed lines ; g. roof of bursal cavity. Published as Miscellaneous Publication No. 633 with the approval of the Director of the Delaware Agricultural Experiment Station. Publication No. 408 of the Department of Entomology and Applied Ecology. 2Associate Professor, Department of Entomology and Applied Ecology, University of Delaware, Newark, Delaware, 19711. Manuscript received by the editor May 3, 1971. 49 50 Psyche [March-June Scenopinus bermudaensis n. sp. Figure i Holotype: (female) — Bermuda, May 15, 1909, F. M. Jones. Coll. ; Museum of Comparative Zoology, Cambridge, Mass. Type no. 32001. Length body: 4.0 mm., wing 3.2 mm. Female Head red-brown ; eyes brown with a very narrow postocular ridge; frons broad, rugose, swollen on lower portion, divided by a shallow median groove ending in a shallow depression on lower fourth, frons elevated above eye margins ; ocellar tubercle red-brown, distinct, cut off; ocelli orange; mouthparts yellow-brown, well developed ; palpi red-brown ; oral cavity bordered by silvery pubes- cence that extends to bulge on lower frons; antennae with first segment red-brown, short; second segment orange-brown; third segment red-brown, pubescent, about twice as long as broad; see figure for details. Thorax with dorsum red-brown, rugose; humeral and supra-alar calli orange-brown; pleural areas red-brown; wings brown fumose, veins brown; halter stem red-brown, knob red-brown; legs with femora and tibiae red-brown, tarsi orange-brown. Abdomen red-brown; tip of 9th tergum and sternum broken from specimen; see figure for conformation of 8th sternum and bursal cavity. Male unknown . References Kelsey, L. P. 1969. A Revision of the Scenopinidae (Diptera) of the World. Nat. Museum, Bull. 277: 1-336, 208 figures. U.S. DIFFERENTIATION OF THE CARABID ANTENNA CLEANER1 By T. F. Hlavac2 Museum of Comparative Zoology Harvard University The antennae of most carabids are groomed by tightly packed setae on the protibia. The mouthparts are not employed in antenna cleaning in ground beetles, but are so used in many 'Coleoptera (Jander, 1966, table 2). A protibial antenna cleaner may be pri- mitively absent only in Nototylus (Banninger 1927: 771) but is secondarily lost in advanced paussines (Darlington 1950: 65). There is great variation in the degree of development of the cleaning setae and in protibial structure. This paper describes and analyzes struc- tural differentiation of the carabid antenna cleaner, and presents preliminary data on grooming behavior. Work on the fine structure and histology of the cleaning setae and associated glands is in progress and will be reported elsewhere. The protibiae of about 100 genera representing 60 tribes were studied; 50 species were measured. A description of methods em- ployed and a formal list of carabids examined in this study will be presented in a paper on prothoracic morphology of the Coleoptera. Structure. The antenna grooming setae are located on the medial face of the tibia; there are two types of setal aggregations. The major cleaning element is the setal band (Figs. 1, 21 SB). It is composed of very tightly packed setae arranged in a single file that always begins near the anterior spur and may extend nearly hori- zontally across the width of the medial face (Figs. 2, 7, 9) or may extend vertically up the length of the tibia for a considerable dis- tance (Figs. 1, 3, 15 SB). The second type of cleaning cluster con- sists of relatively short, less densely packed vertical rows of setae, usually in single file, originating above the tibial spurs (Figs. 1, 21 ASR, PSR). In a few carabids dense setal rows, which are prob- ably used in grooming, extend above both the anterior and posterior JA preliminary version of this work was submitted as a portion of a thesis to the Biology Department, Harvard University, in partial fulfillment of the requirements for the Ph.D. degree. 2I thank Drs. P. J. Darlington, Jr. and J. F. Lawrence for reading the manuscript. This work was supported, in part, by NSF GB12346, P. J. Darlington, Jr., Principal Investigator; and NSF GB 19922, Reed Rollins, Principal Investigator. Manuscript received by the editor May 14, 1971. 5 52 Psyche [March J une Figs. 1-3. Medial views of distal region of protibia. Fig. 1. Metrius contractus ; Fig. 2. Carabus nemoralis. Fig. 3. Pterostichus lucublandus. Fig. 4. Carabus nemoralis mesotibia. medial view. Fig. 5. Pterostichus lucublandus , medial view of protibia with a section of the antenna in place for grooming. spurs (Fig. 7) ; in others only the anterior row is so developed (Figs. 9, ii, 13, 21, 23, 25). In many advanced forms both rows are either absent or consist of widely spaced setae that may not be employed in grooming (Figs. 17, 19, 31). One or a few long, thick, sinuous bristles at the proximal end of the band serve to guide the antenna to, and clamp it against, the setal band during grooming, (Figs. 1, 22 CLS). In a few genera ( Omophron , Elaphrus) these enlarged setae are straight. Clamp setae are probably modified band setae. Each group of protibial antenna cleaning setae can be homologized with clusters of bristles on a non-compressed meso- or metatibia. Tibial setae in the Carabidae can be divided into a number (4-6) of vertical rows extending nearly the full length of the tibia and into a horizontal ring of setae that run about the circumference of 1971] Hlavac — A ntenna Cleaner 53 the tibial apex, (Fig. 4 R. Rng). The ring and row setae are sub- ject to much variation in size, number and density. In addition, other setae may be present that are not organized into distinct clusters but are distributed randomly over the tibial surface. When the insertions of the spurs of the middle or hind tibia are more than slightly separated, a setal row extends above each, e.g., Carabus (Fig. 4). In such forms, a number of ring setae are found between the spurs. The medial aspect of the tibia consists then of some horizontal ring setae between the vertical setal rows and the spurs. The geometry of these setal aggregations relative to each other and to the spurs on the mesotibia is similar to that of the protiba of some carabids, e.g., Carabus , Opisthius (Figs. 7 and 9). As a preliminary hypothesis, the setal band of the protibia is considered to be serially homologous to that portion of the setal ring between the spurs in the other legs, and the anterior and pos- terior rows are serially homologous to the setal rows above the spurs. This hypothesis assumes, of course, the existence of a similar setal organization on all tibiae in primitive forms before some of the protibial setae became densely packed and specialized for antenna cleaning. The evolution of the carabid protibia can be looked at as divergence from the unspecialized configuration of the mesotibia. One important component of this divergence is the modification of protibial structure for antenna grooming. The amount of proximal lengthening of the setal band is the best single indicator of pro- tibial specialization for antenna cleaning. The length of the setal band (measured along the tibial long axis from the anterior spur insertion to the most proximal clamp seta) over the total length of the tibia (up to the articular head) or SB/Tb X 100 is a simple measure of setal band development. This method underestimates the length of a sinuous or horizontal band, par- ticularly those of grades A and C. In the least specialized forms, (cicindelines, Opisthius ), the setal band is almost completely hori- zontal, located near the distal margin of the medial face, and SB/Tb is less than 10%. In carabines (except Pamborus , Jeannel 1941, Fig. 98), cychrines and in the nebriines the band first extends verti- cally for a very short distance, where it is confluent with the anterior row, and then swings horizontally across the medial face; SB/Tb is between 10 and 20 per cent. In these forms, while the band is shifted proximally for only a short distance, the posterior spur is shifted as well to maintain the geometric relationship between spurs and band. Much greater proximal shifts of the band are associated with a shift in spur position in many carabids. For example, SB/Tb 54 Psyche [March-June 1971] Hlavac — A ntenna Cleaner 55 is 33 per cent in Hiletus (Figs, io, n), 37 per cent in Nomius Figs. 12, 13), 46 per cent in Pterostichus (Figs. 14, 15) and 52 per cent in Plelluomorphroides (Figs. 16, 17). In each case the posterior spur is located close to or at the level of the most proximal clip seta. In a few carabid taxa, a large proximal development of the setal band has occurred without any movement of the posterior spur. For example, SB/Tb is 30 per cent in Trachypachus (Figs. 20, 21 ), 40 per cent in Metrius (Figs. 24, 25), 55 per cent in Tropopsis (Figs. 26, 27), and 58 per cent in Gehringia (Figs. 28, 29). Bell (1964) stated that only one tibial spur is found in Gehringia. In fact, two small apical spurs are present (Figs. 28, 29) as well as a “spinose hair” which Bell thought earlier authors had mistaken for the posterior spur. The setal band in a few forms differs little in position from that of ring setae on the mesotibia (compare Figs. 3, 4, 6, 9). Addi- tional modification of the setal band occurs with (Figs. 16-19) or without (Figs. 20-28) a proximal shift in the posterior spur. In each case, the band is expanded and the tibia is modified in similar ways, (compare Figs. 14, 15, with 26, 27). Jeannel (1941) pro- posed the terms Isochaeta and Anisochaeta for two taxa based on the position of the posterior protibial spur. This arrangement was followed by Bell ( 1967) but not by Ball ( 1963) or Lindroth ( 1969). Basing conclusions on the relationships of higher categories or sim- plistic data from part of a functional system is unsatisfactory on methodological grounds. Jeannel’s arrangement is also faulty on practical grounds. The spur insertions of the mesotibia and of the least specialized protibiae are level or nearly so, i.e., the spurs are isochaetous. Yet, Jeannel considered some of these slightly differ- entiated forms to be anisochaetous, e.g., cicindelines, Opisthius. Figs. 6-31. Posterior (even numbered Figs.), and medial views of cara- bid protibiae. Figs. 6, 7, Opisthius richardsoni’, Figs. 8, 9, Carabus nemo- rails’, Figs. 10, 11, Hiletus ‘versutus; Figs. 12, 13, Nomius pygmaeus ; Figs. 14, 15, Pterostichus lucublandus ; Figs. 16, 17, H elluomorphroides texana ; Figs. 18, 19, Agra sp. ; Figs. 20, 21, Trachypachus gibbsi; Figs. 22, 23, Metrius contractus, Figs. 24, 25, Mystropomus regularise, Figs. 26, 27, Tro- popsis mar ginicollis ; Figs. 28, 29, Gehringia olympica; Figs. 30, 31, Platy- cerozaena panamaensis. Setal aggregations, indicated by dotted lines, limits of antennal channel shown with a dashed line. Scale limits beneath odd numbered figs, indi- cate width of antennal segment 9. Lettered brackets enclose protibiae of the same grade of development; see text for explanation. 56 Psyche [March-June A different arrangement is obtained by considering divergence from an unspecialized tibia on the basis of posterior spur position relative to the tibial apex and to the most proximal point of the setal band. The protibia of Opisthius , for example, is little modified, yet cleaning setae are present; the antenna cleaners of other cara- bids can be derived from this sort of configuration along two parallel evolutionary pathways. In the isochaetous pathway, large proximal shift of the setal band occurs without shift of the posterior spur; while in the anisochaetous pathway, shift of band and spur occur together. In each pathway, divergence from an unspecialized protibia has similar components: 1 ) Proximal development, elongation and differentiation of the setal band into a distal and a proximal cleaning portion. 2) Origin and development of the medial expansion and the cleaning arc. 3) Shift of the antennal channel from vertical to oblique. 4) Anterior-posterior compression of the tibia. 5) Degree of development of anterior and posterior setal rows. Based on the degree of development in the above mentioned factors, the forms examined can be divided into three grades diagnosed as follows : Grade A : Setal band short, almost entirely horizontal, located close to distal rim of medial face; confluent zone, if present, very short; SB/Tb less than 20 per cent. Medial expansion absent. Antennal channel long, vertical. Anterior row always present, posterior row may not be used in cleaning. Tibia not compressed antero-posteriorly. Grade B: Setal band long with distinct vertical section and con- fluent zone, SB/Tb between 26-58 per cent, but is usually less than 40 per cent. Confluent zone short, ranging from 15-35 per cent of the length of the band. Medial expansion present, except in Miletus , but is usually not shifted far anteriorly. Antennal channel shallow and developed far above clip setae, or not. Tibia not compressed antero-posteriorly. Grade C: Setal band, long (SB/Tb 33-69 per cent) divided into a large distal region, or confluent zone, and a proximal clean- ing arc. Distal region varying from 33-69 per cent the length of the setal band. Medial expansion well developed anteriorly. Channel deep, short, does not extend above clip setae. Anterior and posterior rows, if present, usually do not form cleaning aggregations. 1971] Hlavac — A ntenna Cleaner 57 Grade A: Cicindelini, 'Carabini* (in part), Cychrini*, Opisthiini*, Enceladini Grade B: Trachypachidini*, Omophronini, Notiophilini*, Ela- phrini, Loricerini, Promecognathini1**, Siagonini**, Migadopini, Hiletini*, Nomiini, Psydrini, Merizo- dini**, Bembidiini, Pogonini, Carabini (only Pam- borus*) , Ozaenini (only Metrius ** and Mystropo- mus **) Grade C: Members of 40 other tribes. ^denotes groups with open procoxal cavities, and ** forms with advanced grade B configurations. Table 1. Distribution of grade A and B antenna cleaners within the Carabidae. The distribution of antenna cleaner grades within the Carabidae, is given in Table 1. Each grade is characterized more fully below. The setal band in grade A is almost entirely horizontal and situated very close to the distal rim of the tibia. In some forms ( Opisthius, Figs. 6, 7; Cicindela ) there is almost no confluent region between the anterior row and a vertical section of the band, while in others ( Carabus , Figs. 8, 9, Nebria) there is very small con- fluent zone. The gross shape of the tibia is only slightly modified for antenna cleaning. The proximal region is slender; distally it is broadened (Figs. 2, 8). The medial face bears a slight concavity or antennal channel that extends up the length of the tibia far above the setal band. In some grade A forms, e.g., Carabus (Figs. 2, 9), the anterior row is dense distally and is a cleaning element while the posterior row consists of a few widely spaced setae and is not a part of the cleaning system. In others, e.g., Enceladus , Cicindela , Sca- phinotus, and Opisthius (Figs. 6, 7), both the anterior and posterior rows are modified distally into triangular cleaning tufts, many setae wide, that may obscure portions of the setal band. Proximally, the setae of each row are widely spaced and in a single file. Grade B configurations are variable. Hiletus (Figs. 10, 11) and Mystropomus (Figs. 24, 25) represent the least and most modified extremes respectively. It is likely that if more forms were examined, the gap between grades B and C would be quite small. The setal band ranges from 26-58 per cent of the length of the tibia and is divisible into a straight distal, confluent region and a proximal sinuous portion that cleans the antenna. The vertical con- 58 Psyche [March-June fluent zone is short, frequently less than 20 per cent the length of the band. This section is the largest in Mystropomus , Metrius and Promecognathus where it is 30, 34, and 35 per cent respectively. The proximal section begins by curving posteriorly and goes hori- zontally across the tibial face and then curves dorsally and extends vertically up the tibia. Except in Hiletus (Figs. 10, 11), the vertical proximal section of the band is found on part of a flattened, medially expanded section of the tibia. The medial expansion is shaped like a triangle whose apex is at the level of the clip or guide setae and/or the most proximal point of the setal band, (Figs. 1, 22 MEx). The distal side of this triangle is curved in a gentle arc and bears band setae; while the proximal side, which is less flattened, forms the posterior border of the antennal channel. The medial expansion varies greatly in development and orientation. In some forms ( Nomius, Figs. 12, 13) it is almost entirely vertical with only a slight anterior shift. The channel is parallel to the tibial long axis, as in Carabus (Fig. 2). In others ( Mystropomus , Metrius , Figs. 22-25, and Promecognathus ) the expanded region and the band are shifted distinctly anteriorly, but the channel extends far above the clip setae. As the tibia is expanded antero-medially, the clip setae and the proximal part of the band come closer to the level of the confluent zone. The anterior setal row consists of tightly packed short setae in single file and is probably used in antennal grooming (Figs. 1, 11, 13, 23, 25). The posterior row is formed from large, widely spaced setae (Fig. 21) or is absent (Fig. 23). In Grade C, the setal band is divided into a very long, straight, vertical distal region and a curvaceous proximal cleaning section. The anterior row is absent in many of these forms, in which case the non-grooming section of the band is called the distal zone (Fig. 3D). The size of the distal section of the band readily characterizes most grade 'C configurations; it ranges from 33 to 69 per cent of the band length (15 to 34 per cent in grade B forms). This large distal increase in a non-cleaning part of the band results, of course, in a proximal shift of the cleaning elements (Figs. 14, 16, 26). The proximal antenna cleaning part of the band occurs on a vertical, highly curved region of the tibia. This cleaning arc, in posterior view, is distinct (Fig. 26 Ac). The cleaning arc has been formed by a complex set of modifications in tibial molding. The flattened medial expansion is larger than that of grade B configura- tions. The tibia along the distal region of the band is also produced medially. Antero-posterior compression of the entire tibia brings the ends of the cleaning arc closer together so that they lie in nearly 1971] Hlavac — A ntenna Cleaner 59 the same vertical plane. (Compare medial and posterior views of Notnius , Figs. 12, 13, with those of Helluomorphroides , Figs. 16, 17). The deep antennal channel begins at, and has the same curva- ture as, the cleaning arc. It then extends obliquely across the medial face and ends slightly above the clip setae (Fig. 3 Ch). The anterior row is absent in grade C anisochaetous forms, but may be present and be a cleaning element in some ozaenines, e.g., Tropopsis (Fig. 27). In a few genera of the lebiine complex the posterior spur is lost, e.g., Agra (Figs. 18, 19) ; Taicona, Lebia (Habu, 1967, Figs. 17, 18). The tibia, in a few genera, has become very setose distally, e.g., Tropopsis (Fig. 27) ; and in some of these, the setae are so densely packed that the distal zone of the band is not recognizable, e.g., Agra , Platycerozaena (Figs. 19, 31). In many anisochaetous forms, a short linear cluster of relatively fine setae originates close to and above the insertions of the anterior spur and extends posteriorly for a short distance, e.g., Helluomorphroides, Pterostichus (Figs. 3, 15, 17 DC) ; (Habu 1967, Figs. 5-29). These distal cluster setae are modified ring setae and may play a role in grooming the mesotibia. Behavior. Antennae cleaning and other aspects of grooming be- havior of two carabid species were studied in detail and photo- graphed. The grade A antenna cleaner of one species, Scaphinotus ( Brennus ) stratiopunctatus (Chaudoir) (Cychrini), is similar to Figs. 32-34. Antenna cleaning behavior of Scaphinotus (Brennus) stratio- punctatus (Chaudoir). See text for details. 6o Psyche [March -June that of Carahus (Figs. 2, 8, 9) ; while the grade C configuration of the other species, Pristonychus complanatus Dejean (Agonini), is similar to that of Pterostichus (Figs. 3, 14, 15). Placing a drop of clove oil on an antenna stimulates rapid groom- ing behavior, even under the high light intensities used for motion picture photography. Except for rate, antenna cleaning behavior of beetles stimulated with clove oil differs little from that observed in carabids subjected to less violent stimuli (talc, india ink) at low light levels. Scaphinotus begins an act of antenna grooming by lowering the body and by lowering the head and twisting it ventrally towards the side about to be cleaned. The antenna is held nearly perpendicu- lar to the body. The proleg is then raised and lowered on the antenna near segment one. The antenna is now located between the tibial spurs and held against the setal band by the clip setae. The proleg is depressed until it touches the substrate, moving seg- ments one to four or five through the cleaning setae (Fig. 32). The rest of the antenna is drawn through the antenna cleaner by raising the head and the rest of the body (Figs. 33, 34). The proleg is stationary after reaching the substrate. As the antenna is being cleaned, it is bent into three sections (Fig. 33). The segments about to be cleaned lie on or close to the substrate, next to the tarsi, and are nearly perpendicular to the Figs. 35, 36. Antenna cleaning behavior of Pristonychus complanatus De- jean. See text for details. 1971] Hlavac — - A ntenna Cleaner 61 segments just cleaned which are in the channel above the setal band. From the channel, the proximal segments curve broadly to the antennal insertion. Only rarely are both antennae cleaned simul- taneously. In PristonychuSj the proleg is lowered on the extended antenna. The large clip setae engage the antenna and press it against the setal band (Fig. 35). The antenna is drawn through the cleaner by the proleg’s posterior movement, which is generated by coxal rotation, as well as by dorsal movements of the head and thorax (Fig. 36). During cleaning, the antenna is not bent and the apical segments do not touch the substrate. In both species, the protibial antenna cleaner is rubbed against the mesotibia after grooming the antenna several times. Occasionally the protibia is drawn through the mouth parts. The meso- and metatibia bear grooming setae. Fragmentary data indicate that a grooming system involving, in part, anterior movement of particulate matter to the protibia where it is ingested may be present in carabids. Such a system is known in ants (Sudd 1967). Only about half the surface area, but most of the sensory regions, of a flattened antennal segment can come in contact with the band setae during a single act of grooming (Fig. 5 SE). Rotation of the antenna through movement of segment one could permit the other half to be cleaned. Discussion. The effect of grooming on cuticular surfaces in cara- bids and other insects is poorly known. Particulate matter is, of course, quickly removed. The metapleural gland in ants secretes a material with antibiotic properties that is spread over the body sur- face by grooming (Maschwitz, et al, 1970). The association of single cell glands with antenna cleaning setae and other grooming bristles in carabids and other forms suggests that more than me- chanical removal of detritus is involved in grooming. In any case, until these grooming functions become known, it would be useless to speculate on the specific selection pressures that have led to the development and differentiation of the carabid antenna cleaner. The arguments for biological improvement to be developed below are based on differences in mechanics, not on differences of ultimate function. The conclusions drawn from such observations are first approximations. The grade C antenna cleaning mechanism is an improvement over the grade A type. For effective cleaning, the width of the antennal segment being groomed should be parallel to, and lie snugly against, the setal band, as in Fig. 5. If otherwise, only part of the surface 62 Psyche [March-June of a segment will be groomed, or uneven pressure will be applied. In grade A systems, the setal band is horizontal and located close to the tibial apex. These facts impose restrictions on grooming behavior. The setal band and the flattened surfaces of the apical segments can be parallel only if the tibia is placed in a narrow zone near the level of the antennal insertion. Outside this zone, dorsal movement of the head or posterior rotation of the coxa will draw the antenna obliquely through the band. This is especially so since the antennal segments (looked at as an ellipse in cross section) are capable of little motion parallel to the semi-major axis. The pro- tibia of Scaphinotus, and perhaps other grade A forms as well, is stationary after reaching the substrate, maintaining the geometric relationship between the antennal surface and the band throughout a sequence. Since the band is located close to the tibial apex, and since the spurs contact the substrate during grooming, the antennal segments about to be cleaned are dragged across the substrate (Fig. 33). And, the antenna, as a whole, undergoes considerable bending, parallel to the semi-minor axis. In grade C configurations, the cleaning arc is vertical and located above the tibial apex. Movement of the tibia will not pull the antenna out of alignment with the band since the antenna is gripped by the clip setae and is capable of considerable movement parallel to the semi-minor axis. While the antenna is drawn through the cleaning setae, by movements of both the head and proleg, the apical segments do not contact the substrate nor is the antenna bent (Figs. 35, 36). In permitting use of an additional grooming movement, coxal rotation, by developing vertical cleaning elements; and in prevent- ing the antenna from contacting the substrate, by moving the cleaning seatae proximally, the grade C system is an improvement over grade A. The setal band in grade B forms has a vertical grooming component that is not developed far above the tibial apex (Fig. 13). In these forms, coxal rotation can be a grooming move- ment but the antenna may come in contact with the substrate. Grade B systems may be intermediate functionally as well as struc- turally. And so, the three grades of antenna cleaners, defined above on structural criteria, represent grades of improvement in the antenna cleaning mechanism. There are two types of evidence which suggests that the three grades represent sequences of historical development: 1971] Hlavac — Antenna Cleaner 6 3 1 ) the nature of putative reductions of the antenna cleaner mechanism, and 2) correlation of presence of either grade A or B cleaners with that of another presumably primitive character — the open procoxal cavity. Reduction of the antenna cleaner is found in very few groups. In several ozaenine genera, e.g., Platycerozaena (Figs. 30, 31), Physea , Ozaena (Banninger 1927, Figs. 1, 2, 7) the curvature of the cleaning arc is reduced and the antennal sensory regions are very much smaller than those of related genera, e.g., Tropopsis (Figs. 26, 27). A very long, straight setal band is present in primitive paussids, e.g., Protopaussus, Carabidomemnus , Eohomopterus and some species of Arthropterus. In these forms, the antennae are not greatly modi- fied and bear a moderate number of setae. The setal band is absent in other paussines, where the antennal segments may be greatly dilated and fused together (Darlington 1950: 60-65, Figs. 45-103, I 20-127). The antennae of Adelotopus (Pseudomorphinae) are not as long as the head and bear very few setae and sensillae. The antenna cleaner is greatly reduced, and may no longer be used in grooming. It consists of a few widely spaced setae arranged in a vertical row between the spurs. A single large straight seta near the posterior spur is probably a vestigial clip seta. In related genera, e.g., Sphal- lomorpha the antenna is long, bears dense setation, and the cleaner is a normal grade C type. In each case, both the antenna and the cleaner have undergone reduction. Furthermore, the grade C cleaners are simply reduced ; there is no evidence of reversal towards a grade B configuration. The setose regions of forms with either grade A or B cleaners are quite variable in size, but except for the trachypachidines, they are much larger than in the forms cited above as examples of reduction. Except for Gehringia , carabids with open procoxal cavities have either a grade A or B antenna cleaner, while forms with closed coxal cavities have either a grade B or C cleaner, except for the cicindelines and Enceladus (Table 1). Closed coxal cavities and related structures represent a mechanical improvement in the pro- mesothoracic joint. Open cavities are probably primitive in carabids, (Hlavac, unpublished observations). Improvement of the antenna cleaner is then correlated with improvement of the pro-mesothoracic j oint. Reduction of the antenna cleaner mechanism is a rare phenomenon. 64 Psyche [March-June The least specialized cleaners are found in forms with the least specialized and possibly primitive pro-mesothoracic joint. The grades of antenna cleaners probably represent sequences of historical develop- ment, and the terms primitive, intermediate, and advanced can be applied to A, B, and C grade configurations, respectively. It has been argued above that two developmental pathways have been employed in the parallel differentiation of the carabid antenna cleaner, and that the grades may represent historical sequences of mechanical improvement. The major cleaning element, the setal band, is a modified portion of the horizontal ring of setae about the tibial apex. Mechanical improvement of a primitive, i.e., grade A, configuration consists of a shift of the setal band from horizontal to vertical so that coxal rotation can be used in antenna grooming. The pathways are geometric alternatives; the setal band is shifted proximally and vertically with or without the posterior spur. Im- provement then implies choice of pathway. Polyphyletic adoption of a given pathway is readily visualized. Similarity of pathway is consistent with, but is not in itself, evidence for phylogenetic rela- tionship. Each case must be dealt with separately and evidence collected from other functional systems before conclusions are reached. When such evidence becomes available, the distributions of the several types of antenna cleaner will be germaine to a discus- sion of carabid evolution. Summary 1. An antenna cleaner, formed from setal aggregations, is present on the protibia of nearly all carabid beetles. 2. The major cleaning element, the setal band, consists of tightly packed bristles arranged in single file. The setal band is serially homologous with a section of the apical ring of setae between the tibial spurs on the meso- and metalegs. 3. In the least specialized, and probably primitive antenna cleaners, the setal band is apical, horizontal and located between the nearly level spur insertions. In others, the band is lengthened, developed proximally and may be divided into cleaning and non- cleaning sections. 4. Based on the degree of divergence of the setal band and related structures, the forms examined can be divided into three grades. 5. The most specialized configuration is found in the majority of carabid tribes; the least modified antenna cleaner is present in only a few tribes. 1971] Hlavac — A ntenna Cleaner 65 6. Specialized antenna cleaners may have evolved in parallel along two pathways: the setal band is shifted with or without the posterior spur. 7. In permitting use of coxal rotation as a grooming movement and in preventing the antenna from being dragged across the sub- strate, antenna grooming behavior of a form with a highly specialized cleaner represents a mechanical improvement over a form with a slightly modified configuration. 8. Reduction of the antenna cleaner is uncommon and always occurs with reduction in antennal setation. 9. The least specialized antenna cleaners are found in forms with the least specialized pro-mesathoracic joint. 10. The grades of antenna cleaners may represent historical sequences of mechanical improvement. Abbreviations Used in the Figures ASp — anterior spur ASR — interior setal row Ch — antennal channel CLS — clip seta(ae) CON — confluent region of SB and ASR D — distal region of SB (— confluent region when ASR is present) DC — distal cluster PSp — posterior spur PSR — posterior setal row R — row(s) of setae on mesotibia Rng — apical ring of setae on mesotibia SB — setal band SE — setose regions of the antenna References Cited Ball, G. E. 1963. Carabidae. In R. H. Arnett, Jr., The Beetles of the United States, pp. 55-181. Banninger, M. 1927. Die Ozaenini. Deutsche Ent. Zeit. (1927), pp. 177-216. Bell, R. T. 1964. Does Gehringia belong to the Isochaeta? Coleop. Bull. 18: 59-61. 1967. Coxal cavities and the classification of the Adephaga. Ann. Ent. Soc. Amer. 60: 101-107. Darlington, P. J., Jr. 1950. Paussid beetles. Trans. Amer. Ent. Soc. 76: 47-142. Habu, A. 1967. Carabidae. Truncatipennes group. Fauna Japonica, Tokyo, 1-338. Jander, U. 1966. Untersuchungen zur Stammegeschichte von putzbewegungen von Tracheaten. Z. Tierpsychol. 23: 799-844. Jeannel, R. 1944. Coleopteres Carabiques. Faune de France 39: 1-571. 66 Psyche [March-June Lindroth, C. H. 1969. The ground-beetles of Canada and Alaska, Part 1. Opus. Ent. Suppl. 35: I-XLVIII. Maschwitz, V., K. Koob, and H. Schildknecht. 1970. Ein beitrag zur funktion der metathoracalariise der Ameisen. J. Insect Physiol. 16: 387-404. Sudd, J. H. 1967. An introduction to the behavior of ants. St. Martins Press, N.Y. 1-200. NOTES ON THE PHASMATODEA OF THE WEST INDIES: TWO NEW GENERA* By Carl Farr Moxey The Biological Laboratories, Harvard University, Cambridge, Massachusetts. The West Indies have a diverse, but poorly known, fauna, of stick-insects, about 80 species being recorded in the literature. In preparing a review of the West Indian Phasmatodea over the past couple of years, I have accumulated probably the largest collection of Antillean stick-insects ever assembled. This has been the result of borrowing material from a number of institutions and of recent collections made in the West Indies; included in this material are representatives of the two genera described below. Thanks are due to the following who helped supply the material described in this paper: Dr. David Rentz and Dr. W. Wayne Moss, Academy of Natural Sciences of Philadelphia (ansp) ; Dr. Ashley B. Gurney, U.S. National Museum (usmnh) ; Dr. Robert J. Lavigne, Uni- versity of Wyoming (rjl) ; Dr. Howard E. Evans, Museum of Comparative Zoology (mcz) ; and Mr. Will Dirk and Dr. George E. Drewry, Puerto Rico Nuclear Center and Luquillo Experimental Forest, Puerto Rico. Dr. Niilo Virkki of the Agricultural Experi- ment Station, Rio Piedras, Puerto Rico, has been helpful in eluci- dating the cytogenetics of one of the new species. T. Preston Webster of the Biological Laboratories, Harvard University, has been in the West Indies three times, and, when not collecting ^/zo/zV-lizards, has brought back a number of interesting phasmatids. Finally, I would like to acknowledge support for my recent collecting trip in Puerto Rico and St. Thomas by an Evolutionary Biology Grant from Har- vard University (NSF Grant, GB 19922, R. 'C. Rollins, Principal Investigator) . Fam. Phasmatidae, subfam. Phibalosomatinae Genus Taraxippus new genus Female: Body form elongate, subcylindrical, extremely spinose. Head elongate, the vertex swollen and spinose. Antennae longer than the anterior legs; scape depressed; pedicel subconical. Com- pound eyes small, but protruding; ocelli absent. Pronotum subrectangular ; defensive gland opening present. Pro- sternum transverse, lyriform. Mesothorax elongate, swollen dorsally and expanded laterally just behind the apex; dorsal pre-median * Manuscript received by the editor May 27, 1971. 67 68 Psyche [March-June Figure 1. Mirophasma ? cirsium (Redtenbacher) , Alta de los Cruces, Colombia, (mcz). Dorsal view. Figure 2. Taraxippus paliurus n. sp., St. Louis du Nord, Haiti, (ansp). Dorsal view. Figure 3. Lamponius restrictus (Redtenbacher), Jayuya, Puerto Rico. (ansp). Dorsal view. Figure 4. Mirophasma ? cirsium. Lateral view. Figure 5. Taraxippus paliurus. Lateral view. Figure 6. Lamponius restrictus. Lateral view. 1971] M oxey — Phas?natodea 69 swelling of the notum concave in the middle and expanded laterally into strong spines. Mesosternum granulose and laterally spinose. Metathorax transverse, rectangular. Median segment about 0.8 times the length of metanotum. Abdominal segments transverse, segments II-VII laterally expanded into spinose lobes, VIII-X narrower than the preceding. Supraanal plate small, triangular. Cerci short, slightly curved. Sternite VII with a postero-median praeopercular organ ; subgenital plate elongate, extending beyond the apex of the abdomen, acuminate apically. Legs elongate and slender. Anterior femora straight basally. Four posterior tibiae anareolate. First tarsomere slightly longer than the next two together. Tegmina and wings absent. Male: Unknown. Type species: Taraxippus pciliurus Moxey, new species. Distribution : This genus is known only from the type locality in the northwestern part of Haiti. Derivation of name: Taraxippus, the horse-scarer, was the name given to the ghost of Glaucus. The state of the suprageneric classification of the phasmatids is in such chaos that it is extremely difficult to place this remarkable genus in the present scheme. At first glance, it would appear to belong to the pygirhynchine genus Mirophasma Redtenbacher (1906), but it can readily be excluded from this group on the basis of its anareolate tibiae and the relatively large median segment. It would thus seem to belong to the Phibalosomatinae, being somewhat related to Lamponius Stal (1875), from which it may be distin- guished by its basally straight anterior femora and the elongate first tarsomere. It is unfortunate that the male of Taraxippus is not known, for it would help to resolve its relationships; I have some reservations about describing it at all from a single specimen, but it is so interesting, that I feel more is to be gained by placing a name in print than bv not. Tbe females of Taraxippus f>aliurus n.sp. (Figures 2 and 5), Tamponius restrictus (Redtenbacher, 1908)1 (Figures 3 and 6), and 1Originally described in the genus Pericentrus Redtenbacher (1908), this species was placed in Antillophilus Carl (1913) by Rehn & Hebard (1938). My recent collecting in Puerto Rico convinces me that Antillophilus (type species: A. brevitarsus Carl), is a junior synonym of Lamponius Stal (type species: P\girhynchus guerini Saussure, 1868). Thus, the species of Lamponius are: L. guerini (Saussure), L. portoricensis Rehn (1903), L. kfugi Redtenbacher (1908), L. bocki Redtenbacher (1908), L. restrictus (Redtenbacher, 1908), new combination, L. brevitarsus (Carl), n. comb., and L. dominicae Rehn & Hebard (1938). 70 Psyche [March-June Mirophasma ? cirsium Redtenbacher (1906) 2 (Figures 1 and 4) form a stunning convergent complex. In all three, the body and legs have become extremely spiny, the vertex of the head and the mesonotum swollen, and abdominal segment VII of the female laterally expanded. The color in life is either green or a mottled green and brown. The habitat of Mirophasma is unknown, but from the altitude of capture (2200 m) and its overall appearance, I would assume that it, like the Taraxippus and Lamponius species, inhabits wet mossy forest, where the spinosity and green color would provide excellent camouflage from a predator. Taraxippus paliurus new species (Figures 2 and 5) Type locality: High mountains near St. Louis du Nord, Haiti. Color light green in life, light reddish brown preserved. Acanthotaxy3 : Head with supra-antennal, supra-orbital, lateral and medial coronal spines; a pair of median spines is situated just anterior to the well-developed occipital median spines; scape with a single spine. Pronotum with anterior, posterior, and postero-lateral pronotal spines present; mesonotum with the anterior mesal, pre- median, post-median, posterior, inter-posterior, and lateral mesonotal spines; anterior and medio-lateral metanotal spines present, medially there is a pair of strong compound spines; mesopleura with lateral, supra-coxal and mesopleural spines; metasternum with a pair of antero-lateral spines; metapleura with lateral, supra-coxal, and meta- pleural spines; medial spines of median segment strong, anterior and posterior spines reduced. Abdominal tergites II-VII with anterior, medial, lateral, and full posterior series of spines; VIII with anterior and postero-lateral spines, full posterior series robust; IX with the anterior and second paired posterior spines strongly reduced, first paired posterior and postero-lateral spines present; X with the typical complement of spines, although the second paired posteriors are reduced; abdominal sternites II-VI with two paired lateral and two paired medial spines; VII with three paired lateral spines. All femora and tibae armed with thorn-like spines. Holotype: A female, pinned. High Mts. near St. Louis du Nord. 2My specimen of this species, from the locality of Alta de los Cruces, Colombia, 2/10, 2200 m (MCZ), agrees well witth Redtenbacher’s descrip- tion, except for its much larger size (55 mm as opposed to 25 mm for his specimen). Either my specimen represents a closely related new species, or his specimen was immature. 3As used by Rehn & Rehn (1939) on the Obriminae of the Philippines. 1971] Moxey — Phasmatodea 7i On wet mossy tree. April, 1929. Haiti. E. C. Leonard, Coll. Light green in life. (ansp). Derivation of name: From the thorny shrub, Paliurus australis. Measurements of type: Length 57 mm Length of mesonotum 12.5 mm Length of metanotum 4.2 mm Length of median segment 3-3 mm Length of anterior femur 13-5 mm Length of median femur 10.2 mm Length of posterior femur 13.5 mm Genus Agamemnon new genus Body form elongate and slender; surface rugose or granulate. Head elongate, rectangular, with a pair of tubercles or spines between the eyes. Antennae longer than the anterior legs; scape depressed; pedicel subconical. Compound eyes small, but prominent; ocelli absent. Pronotum elongate, rectangular; defensive gland opening present. Prosternum transverse, lyriform. Mesothorax elongate, cylindrical in the male, slightly narrowed anteriorly in the female; notum of female medially carinate. Metathorax elongate, rectangular, cylin- drical in the male; notum medially carinate in the female. Median segment about O.5-0.7 times the length of the metanotum. Abdomen cylindrical in the male, segments II-VIII elongate, IX quadrate, X transverse, rounded posteriorly; abdomen broader in the female, segments II-VII subquadrate, VIII-X narrower than the preceding. Supraanal plate of female prominent, elongate. Cerci short, curved in male, straight in female. Vomer of male triangular, with a short sclerotized tip ; subgenital plate fornicate, apex rounded ; genitalia with a dextral sclerotized “hook.” Sternite VII of female with a highly specialized postero-median praeopercular organ ; sub- genital plate elongate, exceeding the end of the abdomen, apex rounded. Legs elongate and slender. Anterior femora strongly curved basally. Four posterior tibiae anareolate. First tarsomere longer than the second, but not longer than the next two together. Tegmina and wings absent. Type species: Agamemnon iphimedeia Moxey, new species. This genus is most closely related to Ocnophila Brunner (1907), from which it may easily be distinguished as follows: 72 Psyche [March-June 12 1971] Moxey — Phasmatodea 73 Ocnophila 1. First tarsomere of hind tarsi longer than the next two together (Figure 21) 2. Median segment less than O.40 times the length of metanotum 3. Supraanal plate of female small, not elongate (Figure 26) 4. Praeopercular organ of fe- male unspecialized 5. Subgenital plate of female barely exceeding the end of the abdomen (Figure 26) 6. Abdominal segment X of male with a median projec- tion posteriorly Distribution: The genus is known only from the West Indies, with the type species, A. iphimedeia from eastern Puerto Rico and a second species, described by Saussure (1868) as Pygirhynchus thomae ^ from St. Thomas, and herein recorded from western Puerto Rico. The females of these two species may be distinguished : 1. Supraanal plate subquadrate, broadly rounded and with a median notch apically, shorter than abdominal segment X ; mesonotum more than three times the length of the metanotum; anterior femora shorter than the posterior .... A. iphimedeia new species, i'. Supraanal plate lanceolate, longer than abdominal segment X; mesonotum less than three times the length of the metanotum ; anterior femora longer than the posterior A. thomae (Saussure). 1. 2. 3* 4- 5- 6. Agamemnon First tarsomere of hind tarsi not longer than the next two together (Figures 19 and 20) Median segment more than O.45 times the length of metanotum Supraanal plate of female large, elongate (Figures 24 and 25) Praeopercular organ of fe- male specialized (Figures 29 and 30) Subgenital plate of female exceeding the end of the abdomen by one-third its length (Figures 24 and 25) Abdominal segment X of male broadly rounded pos- teriorly Figure 7. Agamemnon iphimedeia n. sp., Luquillo Experimental Forest, Puerto Rico. (rjl). Male, dorsal view. Figure 8. A . iphimedeia, Luquillo Experimental Forest, Puerto Rico. (rjl). Female, dorsal view. Figure 9. A. thomae (Saussure), Aguadilla, Puerto Rico, (usmnh). Female, dorsal view. Figure 10. A. iphimedeia. Male, lateral view. Figure 11. A. iphimedeia. Female, lateral view. Figure 12. A. thomae. Female, lateral view. 74 Psyche [March-June Figure 13. A. iphimedeia. Male, head and prothorax. Figure 1+. Ocnophila poeyi (Bolivar), Upper Ovando River, eastern Oriente, Cuba. (mcz). Male, head and prothorax. Figure 15. A. iphimedeia. Female, head and prothorax. Figure 16. A. iphimedeia. Female, head and prothorax with well-devel- oped spines, (ansp). Figure 17. A. thomae. Female, head and prothorax. Figure 18. Ocnophila poeyi. Yunque de Baracoa, Oriente, Cuba. (mcz). Female, head and prothorax. 1971] Moxey — - Plias?natodea 75 Derivation of name: Agamemnon was a Greek hero of the Trojan War. Agamemnon iphimedeia new species (Figures 7, 8, 10, and 11) Lamponius sp. Ill/ Gunther -f- Lamponius sp. (No. 232) T Lamponius sp. V/Roberts. Virkki, 1970:G-57. Type locality: Luquillo Experimental Forest, eastern Puerto Rico. Color dark reddish brown to black. Female: Elongate. Head with a pair of tubercles between the eyes, vertex tuberculate (Figures 15 and 16) ; first seven antennal flagellomeres not as decidedly elongate as in the male. Pronotum with a pair of tubercles on the anterior, a pair of stubby spines on the posterior margin; the posterior spines may occasionally become enlarged (Figure 16). Abdominal segment VII depressed, VIII slightly narrowed posteriorly, IX transverse and narrowing, X sub- quadrate, slightly narrowed ; supraanal plate subquadrate, broadly rounded and notched apically; abdominal sternite VII with the lateral carinae terminating in a blunt, posteriorly directed spine, behind which sometimes is another smaller one (Figure 29). Sub- genital plate with a median carina in the apical two-thirds; ovipositor valves crossed (Figure 31). Femora and tibiae with the margins subdentate. Male: Elongate, cylindrical. Head with a pair of strong spines between the eyes, vertex tuberculate (Figure 13); first seven an- tennal flagellomers each very elongate, the remaining shorter, lighter colored. Pronotum with a pair of large tubercles on the anterior margin and a pair of large, anteriorly curved spines on the posterior margin (Figure 13). Mesonotum granulose, the granules being numerous anteriorly. Subgenital plate strongly fornicate, with a median longitudinal carina and a transverse V-shaped ridge (Figures 22 and 27). Genitalia only slightly chitinized, with an irregularly lobed basal mass displaced somewhat dextrad, and with a strongly chitinized dextral hook (Figure 28). Anterior coxae with a lateral spine; femora and tibiae with the margins subdentate. Egg: Large, ovoid, surface coriaceous. Micropylar plate shield- shaped, about twice as long as broad. Operculum slightly convex. Length 3.5 mm. (Figure 32). Penultimate nymphal instar: Female — similar to the adult, but the antennae are shorter, the abdomen tapers gradually apically, and 76 Psyche [March-June 1971] Moxey — Phasmatodea 77 the subgenital plate reaches only to the middle of segment X. Length 55 mm. (Figures 34 and 35). Cytogenetics: Virkki (1970) reported the karyotype of this species as 211=34 + XX; the male (Virkki’s number 180) 4 haploid number being I7n + X. In the male, there was one pair of acro- centric, or almost acrocentric, autosomes forming a long rod bivalent and two pairs of metacentrics forming rings, the chiasmata being localized near the centromere. The other autosomes are small- to medium-sized. The X-chromosome is large and submetacentric. Habitat: I found this species never more than two-thirds of a meter off the ground on P/^gr-shrubs in the Luquillo Forest. Lamponius portoricensis Rehn, the common species of stick-insect in the Forest, also occurs frequently on Piper but is usually found at heights of about one to two meters. Derivation of name: Iphimedeia, a daughter of Triops, is a noun in apposition to the generic name. Rearing: In captivity, this species can be raised on Rhododendron spp., Persea americana, and Parthenocissus tricuspidata leaves. One female was kept alive from July to December, 1969; apparently she was a virgin at the time of capture, for she never laid any eggs during the time I observed her. From the recent collecting trip, I had three living females, one of which molted from a nymph to imago. The following notes on defensive behavior derive from observations made on these four. Behavior5: i) Primary defence — The insect is nocturnally active. 4Although I have not seen this specimen (it was sent by Virkki to Klaus Gunther in Berlin), the brief description given by Gunther {in litt .) con- vinces me that assignment to this species is extremely probable. 5In this discussion, I have followed the format and terminology used by Robinson (1969). Figure 19. A. iphimedeia. Female, left hind tarsus. Figure 20. A. iphimedeia. Male, left hind tarsus. Figure 21. Ocnophila poeyi. Female, left hind tarsus. Figure 22. A. iphimedeia. Male, lateral view of apex of abdomen. Figure 23. Ocnophila poeyi. Male, lateral view of apex of abdomen. Figure 24. A. iphimedeia. Female, lateral view of apex of abdomen. Figure 25. A. thomae. Female, lateral view of apex of abdomen. Figure 26. Ocnophila poeyi. Female, lateral view of apex of abdomen. Figure 27. A. iphimedeia. Male, ventral view of apex of abdomen. (cfm). C, cercus; SGP, subgenital plate; V, vomer. Figure 28. A. iphimedeia. Male, ventral view of apex of abdomen with the subgenital plate removed to expose the genitalia, (cfm). C, cercus; DGH, dextral genital hook; V, vomer. 78 Psyche [March -June . i 1971] Moxey — Phas?natodea 79 During the day, it may assume a resting attitude as shown in Figure 36, although it will also rest on the substrate (Figures 37 and 38). ii) Secondary defence — The insect does not display and has almost no escape behavior. It is in general unresponsive to tactile stimula- tion, although it may move the stimulated part of the body away from the source of the disturbance (Figure 39). After repeated pinching, it will walk away slowly, rocking moderately as it does so, and then assume a resting position again. If grasped, the insect becomes immobile, with the fore limbs protracted, and the inter- mediate and posterior limbs extended in the lateral plane (Figure 40) . The species will not regurgitate, and, although defensive gland openings are present on the pronotum, the gland is much reduced and probably non-functional (Figure 41). In a 70 mm specimen of A. iphhnedeicij the gland is only 2 mm long, whereas in an Anisomorpha buprestoides (Stoll) of the same length, the gland is 10 mm long (Figure 42). Holotype: A female, preserved in alcohol. El Verde Research Station, El Verde, Puerto Rico. 22.iii.70. T. P. Webster, (mcz). Allotype: A male, preserved in alcohol. Same data as type. Copulated with female type in plastic collecting bag. (mcz). Paratypic material: El Yunque, P. R. Alt. 1600 ft. Feb. 22, 1927. Coll: S. T. Danforth. r$G (ansp). El Toro Trail, 1st. This specimen is contained in the type collection of the Academy of Natural Sciences of Philadelphia under J. A. G. Rehn’s manuscript name “Lamponius danforthi,” with type number 5709. Figure 29. A. iphimedeia. Female, sternite VII and base of subgenital plate. PO, praeopercular organ; S, lateral spine, just below which can be seen the small additional spine. Figure 30. A. thomae. Female, sternite VII and base of subgenital plate. PO, praeopercular organ; S, lateral spine. Figure 31. A. iphimedeia. Female, ventral view of apex of abdomen with the subgenital plate removed to show the crossed ovipositor valves. IV, inferior valve of ovipositor; SAP, supraanal plate; SV, superior valve of ovipositor. Figure 32. A. iphimedeia. Eggs in dissected female, (cfm). M, micro- pylar plate; O, operculum. Figure 33. A. thomae. Eggs in female abdomen. M, micropylar plate; O, operculum. Figure 34. A. iphimedeia. Penultimate nymphal instar of female, (cfm). Note the regenerating left hind leg. Figure 35. A. iphimedeia. Nymphal female, ventral view of apex of abdomen, (cfm). SGP, subgenital plate. Figure 36. A. iphimedeia. Female resting on twig. (cfm). Figure 37. A. iphimedeia . Female resting on ground, (cfm). 8o Psyche [March-June mile. Sierra de Luquillo, Puerto Rico. 23.iii.70. T. P. Webster. 2 c? <3 • (cfm). 1 st !/? mile on El Toro Trail, El Yunque, Puerto Rico, c. 2500'. T. P. Webster, S. Rand, E. E. Williams, W. Hall. 26-27.vi.1969. 1? (cfm). Night. Tropical wet forest. El Verde, P. R. xii. 12.69. R. Lavigne i?. (rjl). Mossy forest. El Yunque, P. R. 3200'. iii.28.70. R. Lavigne, F. Lavigne, Preston Webster. 1 cT 1$. (RJL). El Yunque, c. 3000 ft., P. R. May, 1938. Darling- ton. icf. (mcz). El Verde. 3.9.64. N. Virkki. 4$$. 7 (ansp). 1st ^4 mile of El Toro Trail, Luquillo Experimental Forest, Puerto Rico. 7.iv.i97i. George Drewry and C. F. Moxey. 2 immature 99- (cfm). 1st Yz mile of El Toro Trail, Luquillo Experimental Forest, Puerto Rico. 12.iv.1971. Will Dirk and C. F. Moxey. 1$. (cfm). El Verde Research Station, Luquillo Experimental Forest, Puerto Rico. May, 1971. data. i$. (usmnh). Will Dirk. 19. (cfm). No Measurements of types: Female Male Length 73 mm 65 mm Length of mesonotum 19.5 mm 19.8 mm Length of metanotum 4.6 mm 5.5 mm Length of median segment 3.0 mm 2.9 mm Length of anterior femur 16.5 mm 15.6 mm Length of median femur 14.4 mm 14.6 mm Length of posterior femur 17.7 mm 17.4 mm Length of supraanal plate 1.0 mm — Agamemnon thomae (Saussure, 1868) new combination (Figures 9 and 12) Pygirhynchus thomae Saussure, 1868: 64; Saussure, 1872: 170; Kirby, 1904: 408. Type locality: St. Thomas, West Indies. 'Color light brown. Female: Elongate, slender. Head with a pair of short, erect spines between the eyes; vertex with very small tubercles (Figure 17). Pronotum with small tubercles. Mesonotum laterally tuber- culate, less than three times the length of the metanotum. Abdomen longitudinally rugose; segment VII not depressed, but narrowed posteriorly, VIII-X with the lateral margins subparallel, X slightly narrowed apically; supraanal plate lanceolate, medially carinate, reaching to the end of the subgenital plate, thus having a beak-like appearance (Figure 25) ; lateral carinae of sternite VII each ending ’These are specimens numbered 244, 250, 251, and 252 by Virkki, 1970: G-57. 1971] Moxey — Phasmatodea 81 Figure 38. A. iphimedeia. Female resting on substrate, (cfm). Figure 39. A. iphimedeia. Same specimen as in Figure 38, after the fore legs and antennae have been touched several times, (cfm). Figure 40. A. iphimcdeia. Female after having been grasped and dropped, showing the extended position of the legs. (cfm). Figure 41. A. iphimedeia. Female, defensive gland of dissected speci- men. (cfm). DG, defensive gland. Scale line equals 1 mm. Figure 42. Anisomorpha huprestoides (Stoll). Female, defensive gland of dissected specimen, (cfm). DG, defensive gland. Scale line equals 1 mm. 82 Psyche [March-June in a small, blunt, posteriorly directed spine; each side of praeopercular organ with an oblique, blunt spine (Figure 30). Subgenital plate longitudinally rugose and medially carinate in the distal three- quarters. Margins of femora and tibiae minutely granulate, lower lateral and lower median carinae of hind four femora each with two subapical granules; posterior femora shorter than the anterior. Male: Unknown. Egg: Large, subcylindrical, surface granulate. Micropylar plate planaria-shaped, about three times longer than broad. Operculum flat. Length 3.5 mm. (Figure 33). Locality: Aguadilla, Puerto Rico. From brush in field near Ramey Air Force Base. April 28, 1948. Oakley, et al. 48 16407. (usmnh). Habitat: The region around Aguadilla does not have any par- ticularly high land and is considerably drier than the Luquillo rain forest. Unfortunately, on my recent collecting trip, I was unable to find this species either on St. Thomas or near Aguadilla. My specimen agrees extremely closely with Saussure’s description, except for the spines between the eyes, the somewhat shorter legs, and the absence of spines on the femora. Measurements : Length 68 mm Length of mesonotum 15.5 mm Length of metanotum 6.0 mm Length of median segment 2.9 mm Length of anterior femur 13.0 mm Length of median femur 10.5 mm Length of posterior femur 12.7 mm Length of supraanal plate 4.0 mm Literature Cited Brunner v. Wattenwyl, K. 1907. Die Insektenfamilie der Phasmiden. II. Lieferung. Phasmidae Anareolatae (Clitumnini, Lonchodini, Bacunculini). Leipzig; Wilhelm Engelmann. Pp. 181-338, pis. VII-XV. Carl, J. 1913. Phasmides nouveaux ou peu connus du Museum de Geneve. Rev. Suisse de Zool. 21(1): 1-56, pi. 1. Kirby, W. F. 1904. A Synonymic Catalogue of Orthoptera. Vol. I. Orthoptera Euplexoptera, Cursoria, et Gressoria. (Forficulidae, Hemimeri- dae, Blattidae, Mantidae, Phasmidae.) London, x+501 pp. 1971] Moxey — Phasmatodea 83 Redtenbacher, J. 1906. Die Insektenfamilie der Phasmid'en. I. Lieferung. Phasmidae Areolatae. Leipzig; Wilhelm Engelmann. Pp. 1-180, pis. I-VI, 2 figs. 1908. Ibid. III. Lieferung. Phasmidae Anareolatae (Phibalosomini, Acrophyllini, Necrosciini) . Pp. 341-589, pis. XVI-XXVII. Rehn, J. A. G. 1903. Notes on West Indian Orthoptera, with a list of the species known from the island of Porto Rico. Trans. American Ent. Soc. 29(2) : 129-136. Rehn, J. A. G. &M. Hebard 1938. New genera and species of West Indian Mantidae and Phasmidae (Orthoptera). Trans. American Ent. Soc. 64: 33-55, pis. III-IV. Rehn, J. A. G. & J. W. H. Rehn 1939. The Orthoptera of the Philippine Islands, Part I. — Phasmatidae ; Obriminae. Proc. Acad. Nat. Sci. Philadelphia 90: 389-487, pis. 31-38, 6 text-figs. Robinson, M. H. 1969. The defensive behaviour of some orthopteroid insects from Panama. Trans. Royal Ent. Soc. London 121 (7) : 281-303, 1 plate, 15 figs. Saussure, H. de. 1868. Phasmidarum novarum species nonnullae. Rev. & Mag. Zool. 2« ser., 20: 63-70. 1872. Famille des Phasmides. Etudes sur les myriapodes et les insectes. In M. Edwards, Recherches Zoologiques pour servir a l’histoire de la fauna de PAmerique Centrale et du Mexique. Paris. 6me partie, Livr. 2, pp. 133-201, pis. 3-4. Virkki, N. 1970. Karyotype rearrangements and malformation of gonads in walk- ingsticks (Phasmatoptera) of the El Verde Radiation Center. In H. T. Odum, A Tropical Rain Forest. A study of irradiation and ecology at El Verde, Puerto Rico. U.S. Atomic Energy Commission. Pp. G-51-G-62, 10 figs. THE MALE GENITALIA OF BLATTARIA. VI. BLABERIDAE: OXYHALOINAE* By Louis M. Roth Pioneering Research Laboratory U. S. Army Natick Laboratories Natick, Massachusetts 01760 Princis (1965) included the following genera in the Oxyhaloidae: Oxyhaloa Brunner, Griffiniella Karny, Nauphoeta Burmeister, Henschoutedenia Princis, Jagrehnia Princis, Coleoblatta Hanitsch, Pronauphoeta Shelford, Leucophaea Brunner, Pelloblatta Rehn, Ileminauphoeta Saussure, Gromphadorhina Brunner, Ateloblatta Saussure, and Aeluropoda Butler. In this paper I shall illustrate the male genitalia of 8 of the above genera. I have not seen males of Coleoblatta , Ileminauphoeta , and Ateloblatta. Princis (1961) included Pronauphoeta and Pelloblatta with a (?) in the Oxyhaloi- dae but did not question their placement in this family in his Cata- logus (Princis, 1965). The male genitalia and subgenital plate of Pronauphoeta are so different from other members of the Oxyha- loinae [I follow McKittrick (1964) in using subfamily rather than family rank], that I do not include it in this subfamily. I have placed Pelloblatta in the Panchlorinae (Roth, 1971). The genus Ploceophilus Rehn was placed by Rehn (1965) in the Oxyhaloinae; it includes one species, P. kohlsi Rehn which lives in the communal nests of the Social Weaver Bird in southwest Africa. Acording to Rehn (1965) the species is related to OxyhaloaJ and he stated that Ploceophilus could be separated from Griffiniella by its [ Ploceophilus ] lappet-like tegmina and absence of wings in both sexes. However, Princis (personal communcation) believes that Rehn was wrong in his interpretation of Griffiniella and that Ploceophilus kohlsi is actually Griffiniella heterogamia Karny. Materials and Methods The male genitalia of museum specimens were treated with 10% KOH, dehydrated, cleared in xylol and mounted in Permount. The source of each of the specimens whose genitalia are illustrated is given using the following abbreviations: (AMNH) = American Museum of Natural History, New York; (ANSP) = Academy of Natural Sciences, Philadelphia; (BMNH) = British Museum * Manuscript received by the editor June 4, 1970. 84 1971] Roth — Blattaria 85 Fig. 1. Male genitalia of Nauphoeta cinerea (Olivier) (dorsal view). Ll =: first sclerite of left phallomere; L2vm = median sclerite of left phallomere (L2 ventromedial) ; L2d = dorsal sclerite of L2 ; R2 — hooked sclerite of right phallomere ; ret — retractable portion of R2 which lies in a membranous sheath. (Natural History), London; (CSIRO) = Division of Entomology, CSIRO, Canberra, Australia; (CUZM) = Copenhagen Univer- sity, Zoological Museum, Denmark; (L) = Zoological Institute, Lund, Sweden; (N) = U. S. Army Natick Labs., Natick, Mass.; (USNM) = United States National Museum, Washington, D. C. The number preceding the abbreviations refers to the number assigned the specimen and its corresponding genitalia (on a slide) which are deposited in their respective museums. Results and Discussions Princis (1961, p. 444) used the male subgenital plate as his final key character in distinguishing the Oxyhaloidae. This plate has a laterally directed recurved pointed projection posterior to each stylus (Figs. 2-5; arrows in 3). McKittrick (1964, p. 45) suggested that this shape may be the closest to the ancestral type and that all other shapes of subgenital plates in the Blaberidae could be derived from it by differential reduction. The subgenital plate is an excellent character for distinguishing Oxyhaloinae because the internal geni- talia (Fig. 1 ) of the 8 genera used in this study are all basically similar. The L2d is separated from L2vm, and is a sclerotized plate 86 Psyche [March-June which is an integral part of the preputial membrane. The prepuce has no distinctive shape, other than this sclerotization. The Li is markedly reduced and, in most species, its sclerotization is restricted mainly to the region of the cleft. The tip of the genital hook (R2) is more lightly sclerotized than the main body of the hook, and in some species is a separate pointed or rounded lobe which may break off. Gurney (1965) has drawings of the genital hooks of Nauphoeta cuiereci and three species of TIenschout edema, but erred in indicating that they are part of the left phallomere. The genera listed earlier are arranged linearly according to Princis (1965). The genitalia are essentially so basically similar in the genera studied here that there is little reason to alter this arrangement. However, there are certain differences in structure of L>2d and. R2 which allow the genera to be placed into three tribes. 1. Oxyhaloini ( Oxyhalod) (Figs. 29-46). Characteristically the upper right side of the L2d is extended into a long relatively narrow arm (Figs. 29, 3 2, 35, 38, 41, 44). The genital hook (R2) differs from all other genera of Oxyhaloinae. The rounded outer surface has minute setae and the apical lobe appears to arise from the dorsal surface as a thinly sclerotized membrane and extends from behind the tip of the darkly pigmented hook (Figs. 30, 33, 36, 39, 42, 45). 2. Nauphoetini ( Griffiniella , Nauphoeta , Henschoutedenia, Leu- cophaea, and Jagrehnia) (Figs. 6-25, 47-139). The curved genital Figs. 2-5. Male subgenital plates of Oxyhaloinae (ventral views). 2. Nauphoeta cinerea. 3. Leucophaea maderae (Fab.). 4. (188 USNM) Henschoutedenia sordida (Shelford) (Allotype of Nauphoeta pro c era Rehn). Mt. Coffee, Liberia. 5. Gromphadorhina brunneri Butler. Madagascar, (scale — 0.5 mm). 1971] Roth — Blattaria 87 hook is usually relatively slender and its tip in some species appears to be a distinct joint attached to the main body of the hook, or it is a more lightly pigmented point which blends into and is an integral part of R2 (Figs. 120, 123, 126, 129). Gurney (1965, p. 11) stated that in N. cinerea, H. procera , H. flexivitta, and II. tectidoma the genital hook differed . . in closeness of the apex to the oppo- site base, and in the position and shape of the flange near the base.” The flange (Fig. 72, f) is found in Leucophaea and J agrehnia, but may be poorly developed (Figs. 81, 105) or absent (Figs. 129, 132, 135, 138) in some species of these genera. In Jagrehnia idonea and J . madecassa , R2 appears to be closer to the Gromphadorhini (see below) than to other species of Jagrehnia. Apparently the tips have been broken off (Figs. 132, 135, 138) and these resemble the damaged genital hooks (Figs. 155- 157) of Grom- phadorhina. The genital hooks of Jagrehnia can be arranged to show a trend from an elongated slender form (Fig. 111) to a stouter (Fig. 129) more robust shape (Fig. 138) approaching that found in the Gromphadorhini (Fig. 157). Jagrehnia idonea , J. madecassa , and Gromphadorhina spp. are all Malagasy species. 3. Gromphadorinini ( Gromphadorhina Aeluropoda ) (Figs. 26- 28, 140-157). In this tribe the retractable portion of R2 is unusually short and therefore cannot be extruded to the same extent found in other genera of Oxyhaloinae (and most other species of Blaberi- dae). The genital hook is robust, black, and the tip is lightly pigmented and resesmbles a nonarticulated segment. In Grompha- dorhina there is a distinct indentation on the inside margin between the tip and main body of the hook (Figs. 144 [arrow], 147, 150, 153). The retractable portion of R2 (Fig. 1, ret) of males of Blattaria in which females mount and palpate his dorsum prior to copulation, is relatively long. Thus, the genital hook is extruded for a con- siderable length and is used in the initial seizure, or to pull down the female’s subgenital plate while she is above him. The short retractable portion of R2 in Gromphadorina probably is correlated with the difference in precopulatory behavior of this genus. In G. portentosa the female does not mount and palpate the male’s dorsum during courtship; the male simply backs into the female to make connection (Barth, 1968). Nothing is known about the mating behavior of Aeluropoda , but it may be similar to Gro?nphadorhina. 88 Psyche [March-June Figs. 6-11. Adult male Oxyhaloinae. 6. N auphoeta cinerea. 7. (28 CUZM). Henschoutedenia elegans (Shelford). Cameroon Republic (det. Princis). 8. (1450 L). Henschoutedenia mombuttu (Rehn). Usambara, Nguelo, Tanganyika (det. Princis). 9. (134 ANSP). Henschoutedenia flexivitta (Walker). Ruanda (det. Rehn). 10. (1451 L). Henschoutedenia occidentalis (Fab.), (det. Princis). 11. (1452 L). Henschoutedenia tectidoma Gurney. Probably from Cameroon Republic, (det. Princis). (scale = 5 mm). 1971] Roth — Blattaria Figs. 12-17. Adult male Oxyhaloinae. 12. (133 ANSP). Hcnschoute- denia epilamproides (Shelford). Yaounda, Cameroon, West Africa (det. Rehn). 13. (1453 L). Henschoutedema bicolor (Shelford). Cameroon Republic (det. Princis). 14. (131 ANSP). Lcucophaea capelloi (Bolivar). Kafakumba, Congo (det. Rehn). 15. Leucophaea maderae. 16. (128 ANSP). Leucophaea grandis (Saussure). Entebbe, Uganda. 17. (1460 L). Leuco- phaea pustulata (Hanitsch). Bunduki-Uluguru Mts., Tanganyika Terr, (det. Princis). (scale — 5 mm). 90 Psyche [March-June Figs. 18-24. Adult male Oxyhaloinae. 18. (152 ANSP). Leucophaca puerilis Rehn. Paratype. Duala, Cameroon Republic. 19. (1454 L). Ja- grehnia minuta (Shelford). East Africa, (det. Princis). 20. (162 ANSP). Jagrehnia gestroiana (Saussure). (det. by Rehn as N auphoeta sudanensis Werner, a synonym). Northeast Africa. 21. (1457 L). Jagrehnia made- cassa (Saussure). Annanarivo, Sikora (det. Princis). 22. (1456 L). Jagrehnia testacea (Brunner) (det. Princis). 23. (123 ANSP). Jagrehnia invisa circumdata (Rehn). Niangara, Congo (det. Rehn). 24. (1455 L). Jagrehnia invisa invisa (Rehn). Yaounde, Cameroon (det. Princis). (scale =: 5 mm). 1971] Roth — B lot t aria 91 Figs. 25-28. Adult male Oxyhaloinae. 25. (173 ANSP). Jagrehnia idonea (Rehn) (scale =: 5 mm). Paratype. Madagascar. 26. (22 BMNH). A eluropoda insignis Butler. Madagascar. 27. Gromphadorhina hrunneri Butler. Madagascar. 28. Gromphadorhina chopardi Lefeuvre. Madagas- car. (Figs. 26-28, scale = 10 mm). [March-June Figs. 29-37. Male- cockroach genitalia of Oxyhaloa spp. 29-31. (12 CUZM). 0. buprestoides (Saussure). Uganda (det. Princis). 32-34. (13 CUZM). O. deusta (Thunberg). East Congo (det. Princis). 35-37. (42 BMNH). O. ferreti Reiche and Fairemaire. Samburu District (det. Cho pard). (scale — 0.2 mm). Psyche 1971] Roth — Blattaria 93 Figs. 38-46. Male cockroach genitalia of Oxyhaloa spp. 38-40. (1448 L). O. perspicua Shelford. Bingerville, Ivory Coast (det. Princis). 41-43. (14 CUZM). 44-46. (1449 L). 0 . minima. Kribi, South Cameroon (det. Princis). (scale — 0.1 mm). 94 Psyche [March-June Figs. 47-55. Male cockroach genitalia of Oxyhaloinae. 47-49. (N). Griffiniella larvalis Princis. 50-53. (N). Nauphoeta cinerca (det. Roth). 54-55. (3 CSIRO). N. cinerea. Australia (det. Roth), (scale, Figs. 47-49 0.1 mm, Figs. 50-55 = 0.2 mm). 1971] Roth — B l at t aria 95 Figs. 56-64. Male cockroach genitalia of Henschoutcdenia spp. 56-58. (133 ANSP). H. epilamproides (from specimen shown in Fig. 12). 59-61. (30 CUZM). H. flexwitta. Gubi, East Congo (det. Princis). 62-64. (27 MCZ). H. flexivitta. Bitye Ja River, Cameroon, West Africa (det. Rehn). (scale — 0.2 mm). 96 Psyche [March-June Figs. 65-73. Male cockroach genitalia of Henschoutedenia spp. 65-67. (1452 L). H . tectidoma (from specimen shown in Fig. 11). 68-69. (188 USNM). H. sordida (Shelford). Allotype of N auphoeta procera Rehn, a synonym. Mt. Coffee, Liberia. 70-73. H. elegans. 70. (28 CUZM) (from specimen shown in Fig. 7). 71-73. (135 ANSP). Lolodorf, Cameroon, (f — flange), (scale = 0.2 mm). 1971] Roth — Blattaria 97 Figs. 74-82. Male cockroach genitalia of Henschoutedenia spp. 74-76. (1450 L). H. mombuttu (from specimen shown in Fig. 8). 77-79. H. occi- dentalis. 77. (1451 L). (from specimen shown in Fig. 10). 78-79. (31 CUZM). (det. Princis). 80-82. (1453 L). H. bicolor, (from specimen shown in Fig. 13). (scale — 0.2 mm). 98 Psyche [March-June Figs. 83-91. Male cockroach genitalia of Leucophaea spp. 83-85. (131 ANSP). L. capelloi. (from specimen shown in Fig. 14). 86-88. (23 AMNH). L. capelloi. Niangara, Congo (det. Rehn). 89-91. (152 ANSP). L. puerilis. Paratype. (from specimen shown in Fig. 18). (scale = 0.2 mm). 1971] Roth — Blattaria 99 Figs. 92-100. Male cockroach genitalia of Leucophaea spp. 92-94. (22 AMNH). L. grandis. Kinshasa, Belgian Congo (det. Rehn). 95-97. (128 ANSP). L. grandis. (from specimen shown in Fig. 16). 98-100. (29 CUZM) . L. puerilis. Cameroon Republic (det. Princis). (scale -gg 0.2 mm). 100 Psyche [March-June Figs. 101-109. Male cockroach genitalia of Leucophaea spp. 101-103. (1460 L). L. pusiulata. (from specimen shown in Fig. 17). (det. Princis). 104-109. (N). L. maderae (det. Roth), (scale = 0.2 mm). 1971] Roth — Blattaria IOI Figs. 110-118. Male cockroach genitalia of Jagrehnia spp. 110-112. (48 BMNH). J. paolina (Giglio-Tos) . Kenya (det. Kevan as Nauphoeta punctipennis Chopard, a synonym). 113-115. (1454 L). J. minuta. (from specimen shown in Fig. 19) (det. Princis). 116-118. (32 CUZM). J. minuta. (det. Princis). (scale = 0.2 mm). 102 Psyche [March-June Figs. 119-130. Male cockroach genitalia of Jagrehnia spp. 119-121. (162 ANSP). ./. gestroiana. (from specimen shown in Fig. 20). 122-124. (123 ANSP). J. invisa circumdata. (from specimen shown in Fig. 23). 125-127. (1455 L). J. invisa invisa, (from specimen shown in Fig. 24). 128-130. (1456 L). J. testacea. (from specimen shown in Fig. 22). (scale — 0.2 mm). 1971] Roth — Blattaria 103 Figs. 131-139. Male cockroach genitalia of J agrehnia spp. 131-133. (173 ANSP). ./. idonea. (from specimen shown in Fig. 25). 134-136. (159 ANSP). J. madecassa. Madagascar (det. Rehn). 137-139. (1457 L). J. madecassa. (from specimen shown in Fig. 21). (scale = 0.2 mm). Figs. 140-148. Male cockroach genitalia of Oxyhaloinae. 140-142. (22 BMNH). Aeluropoda insignis. (from specimen shown in Fig. 26). 143 - 145. (N). Gromphadorhina chopardi. Madagascar. 146-148. (N). Gromphadorhina javanica Saussure. Madagascar, (scale = 0.2 mm). 104 Psyche [March-June Figs. 149-157. Male cockroach genitalia of Gromphadorhina spp. 149- 151. (N). G. portentosa (Schaum). Madagascar. 152-156. (N). G. brun- neri. Madagascar. 157. (N). G. portentosa. (The tips of the genital hooks in Figs. 155-157 are broken off.) (scale — 0.2 mm). io6 Psyche [March-June Summary The male genitalia of 8 genera of Oxyhaloinae show remarkable basic uniformity of characters, especially in the structure of L2d and the marked reduction in Li. Differences in characters of the genital hooks (R2) were used to separate the genera into three tribes as follows : 1. Oxyhaloini. — Oxyhaloa. 2. Nauphoetini. Nauphoeta , Griffiniellcij II enschout edema. Leu - cophaea, and J agrehnia. 3. Gromphadorhinini. — Gromphadorhina and Aeluropoda. Acknowledgments I thank the following for the loan of museum material: Dr. N. Jago, Academy of Natural Sciences, Philadelphia, Dr. Jerome G. Rozen, Jr., American Museum of Natural History, New York, Dr. David R. Ragge, British Museum (Natural History), London, Dr. S. L. Tuxen, Zoological Museum, Copenhagen, Dr. Karl Princis, Zoological Institution, Lund University, Sweden, Dr. Ashley Gurney, United States National Museum, Washington, D. C., and Dr. K. H. L. Key, CSIRO, Canberra, Australia. I also thank Mr. Sam Cohen for taking the photographs. References Barth, R. Jr. 1968. The mating behavior of Gromphadorhina portentosa (Schaum) (Blattaria, Blaberoidea, Blaberidae, Oxyhaloinae) : an anomalous pattern for a cockroach. Psyche 75: 124-131. Gurney, A. B. 1965. Two new cockroaches of the genera Pelmatosilpha and Hen- schoutedenia, with a key to the West Indian species of Pelma- tosilpha (Dictyoptera : Blattaria). Proc. Roy. Entomol. Soc. Lond. (B) 34: 5-11. McKittrick, F. A. 1964. Evolutionary studies of cockroaches. Cornell Univ. Agric. Exp. Sta. Memoir 3 89, 197 pp. Princis, K. 1961. Zur systematik der Blattarien. Eos 36: 427-449. 1965. Orthopterorum Catalogus. Pars. 7: Blattariae: Subordo Bla- beroidea: Fam.: Oxyhaloidae, Panesthiidae, Cryptocercidae, Chorisoneuridae, Oulopterygidae, Diplopteridae, Anaplectidae, Archblattidae, Nothoblattidae. ’s-Gravenhage pp. 284-400. Rehn, J. A. G. 1965. A new genus of symbiotic cockroach from southwest Africa (Orthoptera; Blattaria; Oxyhaloinae). Notulae Naturae No. 374: 1-8. Roth, L. M. 1970. The male genitalia of Blattaria. VIII. Panchlora, Anchoblatta, Biolleya . Pcllohlatta, and Achroblatta. Blaberidae: Panchlorinae. Psyche (in press). STUDIES ON THE BIOLOGY OF THE CHRYSOPIDAE II. THE FEEDING BEHAVIOR OF THE ADULT OF GHRYSOPA CARNEA (NEUROPTERA)1’ 2 By Joseph K. Sheldon3 and Ellis G. MacLeod Department of Entomology, University of Illinois, Urbana, Illinois Introduction Although it has long been known that the larvae of the Chryso- pidae are obligate predators of small, soft-bodied arthropods, the feeding habits of their adults have never been studied in detail. The adults of such species as C. chi Fitch, C. incompleta Banks, C. oculata Say, C. nigricornis Burmeister, and C. quadripunctata Fitch are believed to feed primarily, like their larvae, on living prey (Smith, 1922; Burke and Martin, 1956; MacLeod, unpubl.). On the other hand, adults of a number of other species, including C. earned Stephens, will not accept such food, although several other substances are readily consumed. Smith (1922) maintained adults of C. earned (=C. plorihunda Fitch) on a weak sugar solution plus crushed aphids, but he did not ascertain the separate contributions of these dietary items toward adult survival or toward yolk deposition in the female. Finney (1948), whose studies were the first to be directed toward develop- ing methods for the mass rearing of C. carnea for biological control, found that a diet consisting solely of honey resulted in a very low level of oviposition, but when this food was supplemented with a coccid honeydew a much higher level was achieved. Neumark ( 1952) reported obtaining similar “high” levels of oviposition using aphid honeydew as the sole food, while Sundby (1966, 1967) was success- ful in maintaining adults and securing reasonable numbers of eggs from females which were fed upon a diet of honey and pollen. As Finney’s mass rearing procedures had proven uneconomical because of the difficulty of obtaining adequate supplies of honeydew, Hagen (1950) investigated several alternatives. After establishing that the difference in the fecundity of adults fed either honey or honeydew alone was not due to quantitative or qualitative differences Tart I of this series appeared in 1967. J. Insect Physiol. 13: 1343-1349. This work was supported by a grant from the National Science Founda- tion (GB 8644). 3Present address: Dept, of Biology, Eastern Baptist College, St. Davids, Pa. 19087. Manuscript received by the editor, May 12, 1971. 107 io8 Psyche [March-June in the carbohydrate components, he supplemented the honey diet with several synthetic foods known to contain high proportions of proteins (or their products of hydrolysis) and B-complex vitamins. All were found to raise egg production well above the level obtained on honey alone. This early work led to further efforts toward developing highly nutritious, semi-defined, laboratory diets which would permit the efficient mass rearing of large numbers of adults (Finney, 1950; Burke and Martin, 1956; Hagen and Tassen, 1966, 1970; Ridge- way et al.y 1970). None of these investigations attempted to determine the natural feeding habits of the adults of C. carnea. We have therefore cen- tered our attention on a qualitative attempt to discover the food utilized by adults under natural conditions. This has involved an analysis of the gut contents of field-collected individuals and observa- tions on the feeding behavior of adults in the field. In addition, as large numbers of wild adults were found feeding on corn pollen during the summer, we have investigated the efficiency of this ma- terial as food. Materials and Methods Since it has been sugested that honeydew is an important natural food of C. carnea (Hagen, 1950; Neumark, 1952), we began our study with a microscope examination of leaf-surface honeydew in order to obtain a standard of comparison for our study of gut con- tents. This was carried out using two methods. In the first, leaves covered with honeydew were washed in a beaker containing distilled water, and the washings were then poured into a centrifuge tube and spun down. The pellet was placed in a drop of warm glycerine jelly on a microscope slide, a cover slip was added, and the prepara- tion was gently squashed to flatten it. The second method utilized unfed, lab-reared adults which were permitted to feed on leaves covered with fresh honeydew, and an examination was then made of the slide-mounted gut contents. In these preparations the crop, midgut, and hindgut were removed in an isotonic 0.75% NaCl solution (MacLeod, unpublished) and these were mounted directly into glycerine jelly as described above. Studies of the gut contents were also made from similar whole mounts of the crop, midgut, and hindgut of 133 field-collected individuals which had been col- lected at various times throughout the year. These samples were taken, depending on the season, from woodlands or agricultural fields 1971] Sheldon & MacLeod — Chrysopidae 109 at several localities in 'Champaign County, Illinois during 1970 (Table 1). An experiment examining the nutritive value of corn pollen utilized the offspring of 20 females collected at Urbana, Illinois on 15 June 1970 and maintained as a mass culture in an environmental chamber at 25±i°C and at a photoperiod of LD — 16/8. The females had constant access to water and a food consisting of a 1:1: 1 volumetric mixture of Food Wheast®, sucrose, and water (Hagen and Tassen, 1970; Ridgway et al.f 1970). A sufficient number of eggs for the experiment was collected from a single night’s oviposition and these were placed singly in cotton-stoppered, two-dram shell vials. The rearings were carried out at the tempera- ture and photoperiod experienced by the adults in the mass culture, as described previously (MacLeod, 1967), except that vials contain- ing the immature stages were kept in a desiccator over a saturated solution of KBr which maintained a relative humidity of 80%. Upon emerging, the young adults were immediately divided by sex, and 40 of each sex were randomly distributed among the four diet groups shown in Table 2. Each group, still maintained under the same temperature and photoperiodic regimen as before, was provided daily with fresh food. The hand-collected pollen was fed dry, while the sucrose was presented in the form of a highly concentrated solu- tion absorbed in a small cotton pledget. A cotton pledget saturated with water was also available in all four groups at all times. The efficiency of these diets was measured by an examination of the condition of the reproductive system five days after eclosion. In males it was noted whether sperm had shifted from the testis to the seminal vesicle. This is an important initial step in the ontogeny of full reproductive activity by the male (MacLeod, 1967,* Sheldon and MacLeod, in prep.). In the females the basal diameter of the largest ovariole in each ovary was measured,4 the number of mature eggs (diam. — 0.41 mm) was noted, and the number of yolky oocytes was counted. These scores, indicating the degree of development of the female reproductive system, should be excellent indicators of the adequacy of the diet since the process of yolk deposi- tion is undoubtedly responsible for the largest nutritional require- 4The number of ovarioles per ovary in C. carnea is nearly always 12. We have occasionally encountered specimens with 11 ovarioles in one ovary and 12 in the other, but this is the greatest departure from the normal 12/12 state which we have observed. When yolk synthesis is underway, all of the ovarioles of a female are similarly active and usually they all have a similar basal diameter. no Psyche [March-June 1971] Sheldon & MacLeod — Chrysopidae 1 1 1 ment placed on females. All measurements of ovarioles and oocytes were made in isotonic NaCl Ringer’s solution. The results were analyzed using Student’s t test. Field observations of feeding behavior were carried out at night in both forest edge and field communities using a Burgess “Safari Light” with a “cool white” bulb for illumination. This method has the advantage over a regular flashlight of uniformly illuminating a broad area rather than casting a narrow beam. A few individuals seemed to be disturbed by this procedure as they took wing and flew toward the light; however most were apparently undisturbed and seemed to continue their normal activities. Results Analysis of Gut Contents. Honeydew, a secretion emitted from the anus of various Homoptera, consists largely of unabsorbed plant sap to which certain excretory products may be added by some species. From the leaf washings we found that many foreign objects are rapidly trapped in honeydew, the most obvious of these being minute, nonorganic fragments such as small particles of rock, and such other objects as pollen grains, spores of various types, and portions of insect cuticle (particularly exuviae of aphids and lepi- dopteran scales) . The quantity of pollen present varies in propor- tion to the amount being produced in the near vicinity. Also directly associated with the honeydew are sooty-molds (Dematiaceae) grow- Figures 1-4 Photomicrographs of selected areas of the midgut contents of species of ChrysoPa illustrating the principal types of digestive debris found in C. carnea and in the predaceous species C. nigricornis and C. oculata. All figures are prints from polaroid negatives made through a Zeiss Photo- microscope using phase optics. The scale lines in figs. 1, 3, and 4 repre- sent 0.1 mm. The scale of fig. 3 is the same as that of fig. 4. Fig. 1. Acer saccharum pollen from C. carnea. The gut contents of this individual consisted almost exclusively of this single species of pollen. Fig. 2. Residue of honeydew feeding from C. carnea. Visible are fruiting bodies of sooty-molds of the genera Helminthosporium and Alternaria , pollen grains, and a lepidopteran scale. Also visible are setae derived from integumental grooming. Fig. 3. Cuticular remains, mostly antennal fragments, from C. nigriconrnis. Also present are a few scattered pollen grains and fruiting bodies from the sooty-molds Alternaria and Fumago. Fig. 4. Cuticular remains from C. oculata, including the tarsal tips and pre-tarsal claws from several legs and antennal fragments. A few pollen grains are also visible. In both figs. 3 and 4 the irregular dark objects are the fragments of darkly pigmented cuticle. Abbreviations: P — pollen grain; Se — seta; Sc — -lepidopteran scale; SM — fruiting body of a sooty-mold. 1 12 Psyche [March-June ing directly on the surface of the honeydew. Fruiting bodies of Alternaria spp. are very common, with Piricauda spp., Helmintho- sporium spp., and Fumago spp. somewhat less abundant. In addition to the presence of honeydew in the gut of most adults examined, many individuals were also found to contain a considerable amount of pollen (fig. i). We considered an individual to be feed- ing selectively on pollen whenever we found compact masses of one species of pollen predominating in the gut (Table i, columns 3 and 4), while individuals were considered to be feeding largely on honeydew when only occasional, scattered pollen grains, usually of several different species, were present along with the other honey- dew materials mentioned above (Table 1, column 2). Of the 133 specimens examined, 48 had a sufficient amount of one type of pollen in the gut to indicate preferential feeding. Several species of pollen were detected in these individuals, including (at various times of the year) Catalpa bignoniodes, Acer saccharum , Ulmus sp., Carya sp., Celtis occidental™, Zea mays, and a monoporate type of grass pollen. In most of these cases the gut contained evidence of honey- dew in addition to the pollen. Only five of these 48 individuals were found with nearly pure pollen. Of the remaining 85 specimens (Table 1, columns 1 and 2) most had a few scattered pollen grains representing several plant species. The first indication of preferential pollen feeding in our samples occurred in the middle of April and coincided with the early spring flowering of a number of forest trees. Some preferential pollen feeding continued throughout the spring and summer and was par- ticularly noticeable in samples from agricultural fields taken when the extensive stands of field corn tasselled. This utilization of corn pollen was coinfirmed both by examination of the gut contents of the specimens collected on the corn and by direct field observation of feeding adults. The frequent presence of a characteristic type of insect seta in the gut contents at first suggested that either C. carnea was picking up a very common type of seta from the debris on the honeydew or that occasional arthropods were being taken for food (fig. 2). The origin of these setae was ultimately found to be from the grooming of their own integument, as we recovered large numbers of such setae from the gut contents of a lab-reared individual that had been fed only sucrose. This origin was substantiated by a microscopic examination of the integument, which showed that most of the body setae are identical to those recovered from the gut. 1971] Sheldon MacLeod — Chrysopidae 1 1 3 co S c g a <-■> — -q - 4-. ’r| __ c mh > T3 4J - T3 +H p— t OJ o go ”o £ « <-> 2 £»S cn ;> o ~a 'o £ -a Cl, iu C “■Os T3 C C ■3 2 Ji If pollen present, only a few scattered grains of several species. Psyche [March-June 114 Pollen Feeding Experiment. In all 40 of the males the sperm shift had taken place by day five, and no other obvious differences between the dietary groups were present. The results for the females, summarized in Table 2, show an obvious dietary effect. We found that both pure corn pollen and pure sucrose resulted in signifi- cantly less oocyte development compared to the two combined or to Food Wheast® alone. This was true in comparisons of the mean ovariole diameter, the mean number of yolky oocytes, and the mean number of mature eggs per female (P < 0.05 in all cases, except in the three cases where sx is zero and where a t test cannot be made. In these cases the differences are obvious.) Even though by itself corn pollen seems to be a poor diet, it is significantly better than sucrose alone with respect to the ovariole diameter and the number of yolky oocytes (P < 0.05 in both cases). No statistical com- parison was made between these two groups for the number of mature eggs as only a single individual (pollen fed) had matured any. The diets of pollen plus sucrose vs. Food Wheast® were not found to differ in any of the three parameters examined. Field Observations. The observations made in the field were consistent with the data derived from the study of the gut contents. Direct observation of the feeding of adults on pollen in the spring was not possible because of the low numbers of C. carnea present in the habitats sampled and the high frequency of nights too cool for extended activity of the chrysopids. During the middle of the summer a large number of adults were found feeding at night on corn pollen. They were either directly on the tassels or were on the leaves feeding on shed pollen which had accumulated, primarily in the mid-rib depression. On warm, calm nights during the peak of pollen production there were at times as many as two or three individuals per corn plant, and as many as 200-300 individuals could be collected in an hour. Feeding of adults on honeydew was also observed on several occasions during the summer. These observa- tions, made at night, revealed that the adults simply walk along leaf surfaces, stopping periodically to scrape at them. Consistently, a close examination of each area where a chrysopid had stopped revealed a concentration of honeydew. Discussion The seasonal nutritive cycle of C. carnea. The probable use of honeydew as a natural food by C. carnea was pointed out by both Hagen (1950) (who used the synonym C. calif ornica Coq. for this species) and Neumark (1952). That honeydew has a high food 1971] Sheldon & MacLeod — Chrysopidae US value has been demonstrated by several authors who have noted the presence of the mono- and disaccharides fructose, glucose, and sucrose and such trisaccharides as raffinose and gluco-sucrose. Several more- complex sugars are also present along with many of the essential amino acids (Gray and Fraenkel, 1954; Ewart and Metcalf, 1956; Auclair, 1958; Burns and Davidson, 1966). All of these analyses were based on pure honeydew collected shortly after deposition, so that there was no chance for the contaminants which we have men- tioned above to have developed on it. The quantitative nutrient content of honeydews found under natural conditions should vary considerably from that noted above depending on the amount of sooty-mold and trapped pollen which is present since Todd and Bretkerick (1942) have shown that both of these materials contain considerably more amino acids than are present in pure honeydew. Many of the data relating to the chemical composition of different pollens (Todd and Bretherick, 1942; Free, 1970) have indicated the presence of rather high proportions of carbohydrates. Thus in an analysis of pollens from 26 different plant species, Todd and Bretherick (1942) found mean values of 25.71% reducing sugars, 2.71% non-reducing sugars, and 2.55% starch along with mean values of 21.60% crude protein, 4.96% lipids (“ether extract”), 2.70% ash, and 11.19% water. These analyses, however, were based on pollens which had been collected by bees, the workers of which add various amounts of honey and nectar to the pollen during the process of collection and storage (Ribbands, 1953 and references therein), so that the true carbohydrate content of bee-collected pollen is actually lower. Such lower values are shown in a similar analysis of six species of hand-collected pollen, in which Todd and Bretherick (1942) found a mean percentage of only 2.59% reducing sugars, while the absolute amounts of the other major constituents were of approximately the same order of magnitude as those in the samples of pollen collected by bees. Comparisons made beween the different species of these hand-collected pollens show that there is some interspecific variation in the proportions of the major nutrients, particularly starch, and it is obvious that only detailed studies of specific pollens can determine their exact nutritional characteristics. The low proportion of carbohydrate reported for most pollens is consistent with the results of the only other study which has examined the effectiveness of pollen as a complete diet for females of C. carnea (Sundby, 1967). This work indicated a carbohydrate inadequacy for such a diet, since, although there was a low level of oviposition after feeding on (timothy) pollen, a considerably higher level was 1 1 6 Psyche [March-June Table 2. — The effect of diet on the reproductive maturation of lab-reared females of Chrysopa carnea. Diet* Mean Ovariole Diameter Mean Number of Yolky Oocytes per Female Mean Number of Mature Eggs per Female Pollen Only 0.21 ± 0.03mm" 4.6 ± 1.08 0.2 ± 0.20 Sucrose Only 0.14 ± 0.01mm 0.7 ± 0.47 0.0 ± 0.00 Sucrose + Pollen 0.41 ± 0.00 mm 28.0 ± 2.42 12.2 ± 1.67 Wheast® 0.41 ± 0.00 mm 26.5 ± 2.14 12.4 ± 1.5 * N =: 10 females per dietary group b The ovariole diameter in a teneral or diapausing female is approximately 0.07 mm; in a field-collected, reproductively active female it is about 0.41 mm. The standard error of the mean is given for each group. achieved when pollen (of an unstated species) was supplemented with honey. We found a similar marked increase in the reproduc- tive potential of females in the present study when our experimental diet of corn pollen was supplemented with sucrose (Table 2). In this case, however, the carbohydrate inadequacy of corn pollen was unexpected since hand-collected samples of this pollen have been shown to have the exceptionally high value of 36.59% carbohydrate (Todd and Bretherick, 1942). This large proportion of carbo- hydrate is due primarily to a high content of starch, 22.4% (as opposed to 2-3% found in other hand-collected pollens), and the proportions of reducing and non-reducing sugars, 7.31% and 6.88% respectively, are much lower and closer to the means of other hand- collected pollens. The results of a separate study on the utilization of dietary starch by adult chrysopids, which we initiated after the surprising results from the corn pollen experiments were obtained and which will be published elsewhere, show that in C. carnea a large proportion of the starch incorporated into experimental diets remains in an unaltered form in the feces. These findings suggest that the large quantity of starch present in corn pollen is nutritionally unavailable to C. carnea and that the lower concentrations of sugars which are present provide an insufficient carbohydrate source for maximal egg production. The importance of pollen in the natural diet of C. carnea can be 1971] Sheldon & MacLeod — Chrysopidae 1 17 best appreciated when it is considered in conjunction with the sea- sonal cycle of this species. The adults undergo a reproductive dia- pause during the cold months of the year (MacLeod, 1967; Tauber et al^ 1970), and in the early fall migrate from the field commu- nities into the forest edge areas (Zeleny, 1965; Sheldon and MacLeod, in prep.) where they eventually enter the leaf litter and spend the winter. Dissections show that throughout the winter the gut is largely empty, most individuals containing only a few scat- tered setae, pollen grains, and sooty-mold fragments. There is no evidence of extensive feeding. It seems likely that this material represents the undigested residue of feeding late in the autumn, particularly as it is difficult to account otherwise for the presence of pollen in the gut contents at this time. During unseasonably warm periods of the winter there is some movement of the adults within the litter and a few may temporarily leave this habitat and fly about, so that some of this particulate matter which we have seen in our winter dissections may derive from mid-winter feeding on persistent honeydew containing pollen grains and mold fragments. Dissections made during December and January of 1969-1970 do, however, demonstrate the long-term persistence of the residue of late autumn feeding, since this interval had not been interrupted by warm periods. The gut contents of active mid-winter individuals sometimes include a large amount of clear liquid, which suggests that they imbibe free water at this time. With the arrival of warmer weather in the early spring, the adults move out of the litter and begin to search for food. The specimens in the early spring samples in our study (Table 1 — April 1 - 1 5 ) show typical honeydew remains in their gut contents, although most specimens are far from full. There is no honeydew production at this time of the year, and the source of this material in the gut remained an enigma until we noticed that early spring adults could be beaten, with considerable success, from the dry, persistent leaves of such tree species as Quercus alba and Q. palustris. Since many of these leaves had areas of what appeared to be old honeydew on their surfaces, we placed a number of such leaves in a cage with newly emerged, lab-reared adults. An eaxmination of the gut contents of these adults 24 hours later gave results quali- tatively identical to what we had encountered in the guts of our field-collected samples. We have not determined the nutritional adequacy of old honeydew, and it is possible that such other food sources as fermenting sap flows, where one might expect to find similar contaminants to those found on honeydew, are the actual n8 Psyche [March-June sources of the gut contents of our early spring adults. The congre- gating behavior of the adults and the feeding behavior of our lab- reared series suggest otherwise, however, and it is likely that an analysis of the nutritional content of the honeydew from last-year’s leaves would show that, although limited in amount, this is potentially an important initial food source for early spring insects. By the middle of April the pollen production of a number of forest trees begins and this provides focal points of large sources of food which are apparently heavily utilized by the adults of C. earned. The importance of this early pollen becomes evident when the dietary requirements of the females are considered. Males are probably able to complete their reproductive maturation on a smaller energy budget than females, since spermatogenesis is completed in the pupal stage (MacLeod, unpubl.) and, upon adult eclosion, the males have only to shift the mature sperm from the testes to the seminal vesicle (MacLeod, 1967), secrete spermatophores, and mate. Females, on the other hand, have large, continuing nutritional requirements associated with the synthesis of yolk. The spring pollen should provide an energy source adequate for the males to initiate repro- ductive activity, while it conceivably also provides a sufficient source of carbohydrate and amino acids for the production of a limited number of eggs by the females. When it becomes possible for the adults to supplement their pollen diet with a more nearly optimal carbohydrate source such as fresh honeydew or, possibly, nectar, our experiments indicate that this combination should then constitute an excelent diet for maximal reproductive activity. Obviously, as noted above, the nutritional adequacy of this early pollen probably varies from one plant species to another. After their initial, preferential pollen feeding, the adults appear to disperse within the woodlands and into other habitats (Sheldon and MacLeod, in prep.) where they probably feed opportunistically on both honeydew and pollen. Near the middle of the summer, in our area of study, there is again a sharp increase in pollen feeding at the time of anther anthesis of the field corn. This is followed in the late summer by a decreased pollen utilization, which probably corresponds to a reduction in the availability of pollen, until by late fall only honeydew remains are present in the gut analyses. In connection with the suggestion made above that C. carnea is probably unable to utilize starch as a carbohydrate source, it is perhaps noteworthy that, other than corn pollen, none of the major food sources which we have found this species to utilize contain much starch. The failure of C. carnea to make use of starch may 1971] Sheldon & MacLeod — Chrysopidae 119 indicate the evolutionary loss of this ability, since the present ecology of this species provides adequate simple sugars from honeydew and seldom brings adults into contact with foods containing large con- centrations of starch. Notes on the feeding behavior of other Nearctic Chrysopidae. C. carnea is one of a large number of chrysopids which seem to feed opportunistically either as “leaf scrapers” or on pollen. Unlike such predators as C. nigricornis and C. oculata, which usually have numerous, obviously chewed fragments of cuticle in addition to pollen and portions of sooty-molds in their gut (figs, 3, 4), these non-predaceous species lack arthropod fragments other than lepi- dopteran scales ingested during honeydew feeding and the setae scraped from their own cuticle during grooming. By an examination of gut contents we infer the absence of pre- dation and an extensive reliance on the leaf-scraping habit in a rather large number of Nearctic species of several genera, including the close taxonomic relatives of C. carnea within the genus Chrysopa. These close relatives, which comprise the Carnea Group (Subgenus Chrysopa , sens, str ., MacLeod, unpubl.), include C. downesi Smith, C. externa Hagen, C. harrisi Fitch, and C. rufilabris Burmeister. The feeding behavior of C. co?nanche Banks and C. inohave Banks, which are western members of this group, have not yet been examined by us. Although C. rufilabris has been reported by Smith (1922) to be predaceous and to feed readily on aphids, we have been unable to confirm this. Our analyses of the gut contents of field-collected specimens of this species fails to indicate predatory food habits and we have not been able to entice them to feed on aphids in the labora- tory. Burke and Martin (1956) were also unable to observe feeding of C. rufilabris on aphids. Limited field observations on feeding adults and studies of gut contents of several species of Chrysopiella and of Eremochrysa fra- terna Banks suggest that these species feed exclusively on pollen. Acknowledgements We are indebted to D. P. Rodgers for his assistance in identifying the sooty-molds encountered in our analyses of the gut contents; and to R. B. Selander, J. H. Willis, P. W. Price, and A. W. Haney for their constructive criticism of the manuscript. We are also grateful for the use of the facilities of the E. N. Huyck Preserve, Rensselaerville, New York, and the Jackson Hole Biological Re- search Station, Moran, Wyoming where field studies of the feeding behavior of several of the species were carried out. 120 Psyche [March-June References Cited Auclair, J. L. 1958. Honeydew excretion in the pea aphid Acyrthosiphon pisum (Harr.) (Homoptera: Aphididae). J. Insect Physiol. 2: 330-337. Burke, H. R. and D. F. Martin 1956. The biology of three chrysopid predators of the cotton aphid, j. Econ. Entomol. 49: 698-700. Burns, D. P. and R. H. Davidson 1966. The amino acids and sugars in honeydew of the tuliptree scale, Tourney ella liriodendri, and in the sap of its host, yellow poplar. Ann. Entomol. Soc. Amer. 59: 1071-1073. Ewart, W. H. and R. L. Metcalf 1956. Preliminary studies of sugars and amino acids in the honeydews of five species of coccids feeding on citrus in California. Ann. Entomol. Soc. Amer. 49: 441-447. Finney, G. L. 1948. Culturing Chrysopa calif ornica and obtaining eggs for field dis- tribution. J. Econ. Entomol. 41: 719-721. 1950. Mass-culturing Chrysopa calif ornica to obtain eggs for field dis- tribution. J. Econ. Entomol. 43: 97-100. Free, J. B. 1970. Insect pollinators of crops. Academic Press. London and New York. 544 pp. Gray, II. E. and G. Fraenkel 1954. The carbohydrate components of honeydew. Physiol. Zool. 27: 56-65. Hagen, K. S. 1950. Fecundity of Chrysopa calif ornica as affected by synthetic foods. J. Econ. Entomol. 43 : 101-104. Hagen, K. S. and R. L. Tassen 1966. The influence of protein hydrolysates of yeasts and chemically defined diets upon the fecundity of Chrysopa carnea Stephens (Neuroptera) . Vestnik Cs. spol, zool. (Acta soc. zool. Bohemo- slov. ) . 30: 219-227. 1970. The influence of food, Wheast® and related Saccharomyces fragilis yeast products on the fecundity of Chrysopa carnea (Neuroptera: Chrysopidae) . Can. Entomol. 102: 806-811. MacLeod, E. G. 1967. Experimental induction and elmination of adult diapause and autumnal coloration in Chrysopa carnea (Neuroptera). J. Insect Physiol. 13: 1343-1349. Neumark, S. 1952. Chrysopa carnea St. and its enemies in Israel. Ilanoth, Forest Res. Sta., I. 127 pp. Ribbands, C. R. 1953. The behavior and social life of honeybees. Dover Publications, Inc., New York. 352 pp. Ridgeway, R. L., R. K. Morrison, and M. Badgley 1970. Mass rearing a green lacewing. J. Econ. Entomol. 63: 834-836. 1971] Sheldon MacLeod — Chrysopidae 121 Smith, R. C. 1922. The biology of the Chrysopidae. Cornell Univ. Agr. Exp. Sta., Memoir 58: 1291-1372. SundbY, R. A. 1966. A comparative study of the efficiency of three predatory insects Coccinella septempujictaia L. (Coleoptera, Coccinellidae) , Chrys- opa carnea St. (Neuroptera, Chrysopidae), and Syrphus ribesii L. (Diptera, Syrphidae) at two different temperatures. Ento- mophaga 11: 395-404. 1967. Influence of food on the fecundity of Chrysopa carnea Stephens (Neuroptera, Chrysopidae). Entomophaga 12: 475-479. Tauber, M. J., C. A. Tauber, and C. J. Denys 1970, Diapause in Chrysopa carnea (Neuroptera: Chrysopidae) II. Maintenance by photoperiod. Can. Entomol. 102: 474-478. Todd, F. E. and O. Bretherick 1942. The composition of pollens. J. Econ. Entomol. 35: 312-317. Zeleny, J. 1965. Lace-wings (Neuroptera) in cultural steppe and the population dynamics in the specie.? Chrysopa carnea Steph. and Chrysopa phyllochroma Wesm. Acta Entomol. Bohemoslov. 62: 177-194. CAMBRIDGE ENTOMOLOGICAL CLUB A regular meeting of the Club is held on the second Tuesday of each month October through May at 7:30 p.m. in Room B-455, Biological Laboratories, Divinity Ave., Cambridge. Entomologists visiting the vicinity are cordially invited to attend. The illustration on the front cover of this issue of Psyche is a reproduction of the drawing by James W. Chapman of the Smaller Elm Bark Beetle, Scolyfus multistriatus. This was included in his account of the discovery of this insect in the elm trees of the Harvard Yard, in October, 1909 — the first report of this European insect in the United States (Psyche, 1910, Vol. 17, pi. 4). BACK VOLUMES OF PSYCHE The Johnson Reprint Corporation, 111 Fifth Avenue, New York, N. Y. 10003, has been designated the exclusive agent for Psyche, volumes 1 through 62. Requests for information and orders for such volumes should be sent directly to Johnson Reprint Corporation. Copies of issues in volumes 63-77 are obtainable from the editorial offices of Psyche. Volumes 63-77 are $6.00 each. F. M. Carpenter Editorial Office, Psyche, 1 6 Divinity Avenue, Cambridge, Mass. 02138. FOR SALE Classification of Insects, by C. T. Brues, A. L. Melander and F. M. Carpenter. Published in March. 1954, as volume 108 of the Bulletin of the Museum of Comparative Zoology, with 917 pages and 1219 figures. It consists of keys to the living and extinct families of insects, and to the living families of other terrestrial arthropods; and includes 270 pages of bibliographic references and an index of 76 pages. Price $9.00 (cloth bound and postpaid). Send orders to Museum of Comparative Zoology, Harvard College, Cambridge, Mass. 02138. / PSYCHE A JOURNAL OF ENTOMOLOGY Vol. 78 September, 1971 No. 3 CONTENTS The Predatory Behavior of the Golden-Web Spider Nephila clavipes (Araneae: Araneidae). Michael H. Robinson and Heath Mirick 123 Trechoblemus in North America, with a Key to North American Genera of Trechinae (Coleoptera: Carabidae). Thomas C. Barr, Jr 140 Mechanisms Controlling Copulatory Behavior in Wolf Spiders (Ara- neae: Lycosidae). Jerome S. Rovner 150 The Larva of Heliocausus larroides (Hymenoptera, Sphecidae). Howard E. Evans 166 Flights of the Ant Formica dakotensis Emery. Mary Talbot 169 The Male Genitalia of Blattaria. VII. Galiblatta, Dryadoblatta, Poroblatta, Colapteroblatta, Nauclidas, Notolampra, Litopeltis, and Cariacasia (Blaberidae: Epilamprinae) . Louis M. Roth 180 The South American Castianeirinae. I. The Genus Psellocoptus (Araneae: Clubionidae) . Jonathan Reiskind 193 The Genus Oonops (Araneae, Oonopidae) in Panama and the West Indies. Part 2. Arthur M. C bickering '203 CAMBRIDGE ENTOMOLOGICAL CLUB Officers for 1971-1972 President C. S. Henry, Harvard University Vice-President P. VYebster, Harvard University Secretary H. F. J. Nijhout, Harvard University Treasurer F. M. Carpenter, Harvard University Executive Committee A. F. Newton, Jr., Harvard University M. Corn, Harvard University EDITORIAL BOARD OF PSYCHE F. M. Carpenter (Editor), Fisher Professor of Natural History, Harvard University P. J. Darlington, Jr., Professor of Zoology, Emeritus, Harvard U niversity W. L. Brown, Jr., Professor of Entomology, Cornell University; Associate in Entomology, Museum of Comparative Zoology E. 0. Wilson, Professor of Zoology, Harvard University H. W. Levi, Professor of Biology and Curator of Arachnology, Museum of Comparative Zoology H. E. Evans, Alexander Agassiz Professor of Zoology, Harvard University PSYCHE is published quarterly by the Cambridge Entomological Club, the issues appearing in March, June, September and December. Subscription price, per year, payable in advance: $4.50 to Club members, $6.00 to all other subscribers. Single copies, $2.00. Checks and remittances should be addressed to Treasurer, Cambridge Ento- mological Club, 16 Divinity Avenue, Cambridge, Mass. 02138. Orders for missing numbers, notices of change of address, etc., should be sent to the Editorial Office of Psyche, 16 Divinity Ave., Cambridge, Mass. 02138. For previous volumes, see notice on inside back cover. IMPORTANT NOTICE TO CONTRIBUTORS Manuscripts intended for publication should be addressed to Professor F. M. Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass. 02138. Authors contributing articles over 4 printed pages in length may be required to bear a part of the extra expense, for additional pages. This expense will be that of typesetting only, which is about $13.50 per page. The actual cost of preparing cuts for all illustrations must be borne by contributors: the cost for full page plates from line drawings is ordinarily $12.00 each, and the full page half-tones, $20.00 each; smaller sizes in proportion. AUTHOR’S SEPARATES Reprints of articles may be secured by authors, if they are ordered at the time proofs are received for corrections. A statement of their cost will be furnished by the Editor on application. The March-June, 1971, Psyche (Vol. 78, nos. 1-2) was mailed December 30, 1971. The Lexington Press, Inc., Lexington, Massachusetts PSYCHE Vol. 78 September, 1971 No. 3 THE PREDATORY BEHAVIOR OF THE GOLDEN-WEB SPIDER NEPHILA CLAVIPES (ARANEAE: ARANEIDAE) By Michael H. Robinson and Heath Mirick Smithsonian Tropical Research Institute, P.O. Box 2072 > Balboa, Canal Zone (Panama) Introduction Details of the role of wrapping behavior in the predatory activities of Nephila clavipes (Linnaeus) were given by Robinson, Mirick & Turner (1969). Their account also gave broad details of the total behavior of this species. At that time,, the publication of an exhaustive account of the predatory behavior of N. clavipes was in- tended and anticipated. Since then, however, the senior author has carried out studies of the behavior of other species of Nephila in Africa and Asia, and with co-workers is currently engaged in a study of the behavior and ecology of Nephila maculata (Fabricius) in New Guinea. It now seems appropriate for us to leave details of part of our work on N. clavipes for inclusion in a broad compara- tive paper and publish here those aspects which relate most directly to the main points cited in outline by Robinson, Mirick & Turner (1969). This paper therefore presents a summary model of the predatory behavior of Nephila clavipes , based on the investigations of the present authors in the summer of 1968 and further experiments carried out by the senior author in 1969. We give emphasis to the investigations and experiments that led to establishing some of the major aspects of the model and leave detailed descriptions of be- havior units and behavior sequences for inclusion in the projected comparative paper. We have also left consideration of the temporal aspects of the behavior sequences for inclusion in the later paper. Materials and Methods Our basic observations and some of our experiments were car- ried out with captive adult female spiders. The spiders were not confined to cages but were released in a large screened insectary at 123 124 Psyche [September the Barro Colorado Island research station of the Smithsonian Trop- ical Research Institute. We maintained a minimum of fifteen spiders in captivity during the period of study in 1968. The experiments carried out in 1969 required a larger number of spiders and we used free-living spiders in the Barro Colorado forest as well as a number of captive spiders. Even with access to twenty or more non-captive spiders it was necessary to use some spiders more than once in an experimental series ; details of how these were manipulated to obviate the possible effects of experience is given below in the appropriate section. Captive spiders were collected from the same areas as the non-captives and all were of unknown age and previous experience. We made repeated presentations of a number of different prey items in order to establish the basic patterns of the spider’s behavior. This involved fifty presentations of each of seven prey types. These were chosen for their relevance to the natural diet of the spider (see later) and because they presented large differences in size, weight and type of activity after striking the web. We used the data obtained from these observations to prepare ethograms of the type used by Robinson & Olazarri (1971) and then used these etho- grams as a basis for the integrated model. Fifty presentations of each prey item meant that a proportion of the spiders received the same type of prey more than once. In general we presented but one prey per spider each day. In an attempt to avoid any possible effects of experience we avoided successive presentations of the same type of prey. In all cases at least two days (usually more) inter- vened between one presentation and the next presentation of the same type. Usually another type of prey, or several other types of prey, would be presented between presentations of the same type. In addition we made presentations of all the prey types to a large number of free-living spiders as a check on the behavior of our captive spiders. In the course of our initial observational work we presented the following prey items: moths (living & dead), grasshoppers (living), crickets (living & dead), Tenebrio beetles (living), dragonflies (liv- ing), Tfigona sp. (living), pentatomids (living), and blowflies (living). Later when we attempted to elucidate the stimuli which the spiders were capable of detecting at various stages in the process of predation we used a number of experimental techniques involving modified insects in the form of ‘dummies’. These techniques are de- tailed in the appropriate section below. All the insects that we used were weighed and measured before being presented to the spiders. Where insects were presented dead 1971] Robinson Cff Alirick - — Nephila clavipes 125 they were killed by freezing to avoid contamination by chemical killing agents. Behavioral observations were supplemented by cine- photography and the duration of behavior units was recorded on a Rustrak multi-channel chart recorder. Natural History In Panama, and wherever we have encountered Nephila clavipes in South America (Colombia & Venezuela), the spider is most fre- quently found at the edges of forest clearings, alongside forest trails and across forest streams and watercourses. It thus apparently ex- ploits areas within the forest that are in all probability flightpaths of insects. The structure, size, and siting of N, clavipes webs is consistent with the view that the spider specialises on prey that are in flight above the herb layer rather than moving about in it. Al- though the web may be sited with part of the prey capture area within the herb layer it is most frequently above this or stretched across gaps in the vegetation. The structure of the web has been described by Gertsch (1948) and Peters (1954, 1955). For its area, which is large, the web has a very fine mesh which is far less penetrable by small insects than the much smaller but coarse-meshed web of Argiope argentata (Fabricius), as was shown by Robinson, Mirick Sc Turner (1969). The web of the adult spider is not a complete orb but is U-shaped with the hub very close to the upper bridge thread. Kaston Sc Kaston (1953, p. 176) give an excellent figure of the N. clavipes web. Webs of several adult spiders are often built in close proximity and fairly large aggregations of the spider may occur in apparently favourable areas (Shear 1970, has commented briefly on this phenomenon). The adult web is frequently equipped with a barrier web con- sisting of a complex of strong lines arranged in a non-symmetrical manner above and/or below the main plane of the orb. There is considerable variation in the structure of barrier webs. Robinson Sc Robinson (1970) have suggested that they may function as early warning devices enabling the spider to detect the approach of pos- sible predators. Certainly the spider often responds to manipulation of the barrier web by escape or other forms of defensive behavior. Web renewal is not a daily occurrence, and in captivity the spider may only renew part of the web at a time. (This is also certainly the case with free-living N. maculata but we have not made exten- sive observations on web renewal by free-living N. clavipes ). Our observations on the natural prey of N. clavipes , although limited in scope, confirm the deductions based on web structure and siting i.e. that flying prey of small to medium size may be the 126 Psyche [September speciality of this species. We found flies, bees, wasps and small lepi- dopterans to be predominant in the prey found in webs and amongst the corpses that are occasionally suspended from the barrier web after they have been consumed by the spider. 1971] Robinson C5f Mirick — Nephila clavipes 127 Predatory Behavior 1. The basic pattern Figure 1 is a summary diagram of the complex of possible se- quences of predatory behavior that we have observed to be given by adult female N. clavipes to a variety of prey presented in several ways. It has the same summary function that Figure 2 of Robinson (1969) had in relation to the behavior of Argiope argent ata and is not meant to be a flow diagram or relate to any cybernetic con- ventions. It does, however, illustrate the temporal sequence and relationships between behavior units, their degree of association, and the effects of some stimuli on the course of the sequence. Like most models it is certainly much more simple than reality. In the predatory behavior of N. clavipes alternative behavior units are available at several functional stages and the ‘decision’ to employ one or other of these is shown, on the diagram, to be the result of a discrimination. We recognise that in some cases our assumption of the basis (or bases) for the discrimination may be an oversimpli- fication; this matter is discussed at the appropriate place in the text. The diagram employs simple conventions. The behavior units are shown as circles connected by lines. The arrows on the lines show the direction of change from one behavior unit to the next. In the upper half of the diagram two conventions are used to denote ‘choice’ points. Some of the circles are divided into two halves by vertical lines so that two behaviors may follow the behavior shown by the circle. In addition, here and throughout the diagram, small square boxes on the lines connecting behavior circles represent places where behavior may be switched from one course to another. Dotted lines from oblong boxes suggest imputs to the system that are de- pendent on stimulus properties of the prey. Behavior occurring up to first contact with the prey is shown in a much simplified form. The activity spider on hub (which could be amplified as ‘spider at hub in predatory position waiting for prey’) is represented as an ongoing activity by a vertical line beneath the behavior circle. If a prey item, or in fact any item above a certain weight, strikes the web the spider is alerted (the square box has an imput from prey strikes web and an output to spider alert , this represents a diversion from the ongoing activity spider at hub). The overt behavioral change following impact may be a momentary in- crease in the flexion of the spider’s legs prior to an almost immediate movement towards the prey or a more sustained adoption of this alert position. 128 Psyche [September After the spider has been alerted two things may occur. If the prey is in a state of sustained vibration the spider almost always makes an immediate approach to it. If on the other hand, the prey is immobile or merely making spasmodic movements the spider usually plucks the web. If the prey has remained in the web until the spider plucks, this behavior is likely to be followed by an approach to the prey (during which the plucking movement may be repeated). When the prey escapes after the initial alerting impact, the spider usually returns to its pre-alerted state after plucking. As examples of the simplification involved in our model we can cite the following examples that occur in the fairly complex section that we have traced so far. Thus a further alternative behavior that can occur on the impact of prey is not shown. Instead of the spider being alerted it may show an immediate escape response. This usually takes the form of the spider running upwards from the hub, out of the web and onto the support lines (or even onto nearby vegetation). Similarly, if it did not result in an illegibly complex diagram, we could show at least three distinct approach-to-prey behaviors. There is a rapid, unhestitant, approach that is made to rapidly vibrating insects and a hesitant, much interrupted by pluck- ing, approach to non-vibrating insects. In addition very large or very heavy insects are approached in a slow ‘deliberate’ manner, in which legs I & II are flexed far back over the prosoma in a very characteristic ‘cautious’ gait. After arrival at the prey the spider may immediately attack with- out a perceptible pause, or may touch (with the tarsi) and palpate (with the pedipalps) the prey, before attacking. We have not shown these ‘investigative’ stages in our diagram. There are three basic attack behaviors, all of which involve the use of the chelicerae. In no case have we ever seen Nephila clavipes (or any other species of Nephila , for that matter) use the strategy of attack wrapping. This matter is extensively discussed by Robin- son, Mirick & Turner (1969). The three basic forms of biting attack are as follows: 1. An attack similar to the seize and pull out behavior of Argiope argentata (see Robinson & Olazarri, 1971). This is given to very small or light prey. 2. A long bite in which the bite is not immediately followed by pulling out movements but is sustained in situ. This long bite may be accompanied by a special posture in which legs I & II are raised off the web and the opisthosoma is raised at its apex so that the body presents a concave dorsal aspect. 1971] Robinson & Mirick — Ncphila clavipes 129 3. The third attack strategy we call bite & back off. It consists of a rapid forward lunge, a short-duration bite and a rapid with- drawal to a distance at which the prosoma is well away from the prey. We have shown this behavior as a repeatable unit since the lunge, bite, retreat sequence may be repeated a large number of times (we have records of over 12 repetitions) before a sustained long bite ensues. In many cases, when this form of attack is em- ployed, the spider can be seen to open the chelicerae until they are almost held horizontally, before making the rapid forward lunge that terminates in the bite. The type of attack strategy that the spider employs seems to be largely determined by parameters of weight and/or size of prey; our experimental analysis of these factors is described later. We have not been able to determine what factor or factors mediate the decision to cease repetitions of bite & back-off and commence su- stained biting. This process is not dependent on reduction of activity by the prey (that might be consequent on a series of bites) since it occurs in the spider’s behavior towards large dead prey. Behavior following the initial attack phase is somewhat more complex than is the case with araneids that are efficient at enswathing their prey in silk. Attack is almost always terminated by pull-out movements. The body of the spider is lowered, on flexed legs, during the biting attack and pulling out consists of strong extensions of the leg pairs. These result in the spider pushing down on the web and pulling up on the prey. Very small prey, adhering to a small area of viscid spiral, are quickly freed, as are lepidopterans which do not adhere strongly because of their loose wing scales (see Eisner et al 1964). Other prey may be subjected to repeated pull out movements before being freed. Prey that are not readily freed by pulling movements are wrapped in the web and then cut out by the snider. Robinson, Mirick & Turner (1969) called this wrapping at the capture site. Type 1. We have been able to show that the spider can be induced to wrap prey (in this way) if its pulling out attempts are blocked experimentally (see later). It seems possible that the pull out movements enable the spider to gauge the degree of adhesion of the prey to the web and that the ‘decision’ to continue pulling, or to wrap in situ and then cut out, is influenced by this information. A further complication arises from the fact that the spider may wrap, at the capture site, prey that have already been freed from the web by pulling (Post-immobilization wrapping at the capture site. Type 2, of Robinson, Mirick & Turner 1969). 130 Psyche [September (The functional interpretation of these behaviors is that pulling prey from the web results in less web damage than wrapping in situ and subsequent cutting out. This can easily be substantiated by com- paring the web damage resulting from the two techniques of prey- removal. Wrapping after removal from the web has a trussing effect which reduces the bulk of the prey and may facilitate its transportation to the feeding site.) After the prey is freed from the web, and trussed in some cases, it is transported to the hub of the web. Here the spider employs one of two techniques. It may either carry the prey held in the chelicerae, perhaps supported by one or both of the first legs, or it can carry the prey package on a silk thread hanging from the spin- nerets and supported by one or both of the fourth legs. Prey carried to the hub in the jaws are wrapped on arrival at the hub and then suspended from the hub silk as the spider turns to assume its preda- tory posture. Prey that are carried suspended on silk are not wrapped on arrival at the hub but are suspended. Nephila clavipes does not store prey at the capture site but carries all prey to the hub where it is hung until previously caught prey are consumed. Very small prey, carried to the hub in the jaws, may not be wrapped on arrival (but prey as small as stingless bees — io-30mg in weight — are regularly wrapped on arrival at the hub). The return to the hub from the capture site is carried out in a forwards direction after the spider has turned to face the hub at the capture site. ( Nephila maculata frequently backs slowly up the web, without turning, when carrying prey in its jaws.) When the spider has assumed its normal head-down predatory position it may undertake more or less extensive grooming activities before taking up the prey in its jaws and anterior legs. These are very similar to those that Robinson & Olazarri (1971) described for Argiope argentata. Prior to the commencement of feeding the spider often carries out extensive manipulations of the prey during the course of which small bites are given to region after region of the prey body. We have a few records of prey being wrapped after removal from the web and then being transported in the jaws. Most prey that are trussed in this way are then carried suspended on silk behind the spider. Heavy prey are also carried in this way, but as in the case of Argiope argentata , the weight threshold for the changeover from carry in jaws to carry on silk varies from individual to individual, and from time to time within individuals (see Robinson 1969, p. 1 70-1). 1971] Robinson £s? Mirick — Neph'la clavipcs I3i 2. Experimental investigations To investigate the possibility that certain aspects of the predatory pattern are responses to relatively simple stimuli we carried out a number of investigations using ‘modified’ insects as dummies. We manipulated the size and weight of some insects and also modified such parameters as the strength of their adhesion to the web. We were able to compare the responses of the spiders, at various stages in the predatory process, to such dummies, using adequate controls or using their response to unmodified insects as a baseline. (a) The bite & back-off attack behavior Our observations on the responses of N. clavipes to a range of prey items (see page 123, above) showed that the bite & back-off attack behavior was only given to large and heavy prey. Since all of these that were presented to the spiders were alive there was a possibility that the response could be to size, weight or specific activity of large heavy prey, or a combination of any of these factors. Preliminary tests showed that the response was given to dead (im- mobile) prey so that although it remained possible that activity could enhance the response it was not the important stimulus. We then decided to manipulate the parameters of size and weight. Using small acridiids (25-3omm, 400-550mg) as prey, we added lead shot to some to double the weight (approximately), increased the length of others by inserting a wooden tooth pick in line with the long axis, and with the third group we increased both the length and the weight. The dead dummies were presented at right angles to the radii of the web. This ensured that the weight of prey was dis- tributed over as wide an area of web as was covered by the length of the insect (or the insect + toothpick). In fact it meant that the maximum dimension of the prey was at right angles to the spider as it approached across the web. The insects were vibrated elec- trically at 250 cps until the spider left the hub on its predatory excursion. At that stage the vibrator was switched off so that the prey was motionless when the spider came in contact with it. The results of this experiment are shown in Table 1. In addition to the form of biting attack we noted whether the spider raised legs I & II off the web during the attack (see page 127, above), and whether the prey was wrapped in situ, or free wrapped, after the attack. There was a significant increase in leg raising during the attack in the case of the weighted insects. There was a slight numerical, but not a statistically significant, increase in the number of attacks on the long dummies that involved leg raising. There were three attacks out of ten, on heavy insects, that involved the bite and back off behavior. 132 Psyche [September Table 1. Acridiids weighted and unweighted, lengthened and normal. Behaviors. Prey A Prey B Prey C 1. Bite and back off. yes 4 1 1 no 6 9 9 2. Bite, legs raised. yes 9 4 2 no 1 6 8 3. Wrap in web. yes 7 7 2 no 3 3 8 Ten spiders chosen at random from wild population receive A, ten captive spiders B ten wild spiders C. A is acridiid with weight added and lengthened (total weight ca. 1000 mg length 70 mm). B is acridiid lengthened to 70 mm weight 400-550 mg. C is unmodified length 25-30 mm weight 400-550 mg. Statistical analysis — Fisher’s exact probability: differences between A & B, A & C, B & C for Bite and back off are not significant; difference between A & B significance level 0.05, difference between A & C significance level 0.005, no signficant difference between B & C, all for Bite, legs raised; for wrap in web difference between A & B and B & C level of significance 0.05. All levels one-tailed Although this number is not statistically significant we regarded the occurrence of this behavior in attacks on the weighted insects as being highly suggestive. We then carried out a further series of experiments, using similar sized acridiids, in which we quadrupled their weight. We found that very heavy dummies of small size often dropped out of the webs before the spider reached them. We therefore in- creased the length of both experimental (very heavy) dummies and controls (normal weight) by adding the toothpick to the insect. This worked in a perfectly satisfactory way to distribute the weight over a greater number of web members. As before, we vibrated the dummies until the spider commenced its predatory excursion. The results are given in Table 2. There is a statistically significant effect of weight on the occurrence of the bite & back-off attack behavior. Note that we did not induce this attack behavior in all the presentations; we consider that with large, heavy active insects there may be some heterogeneous summation. We attempted to test for the possible additive effect of activity, at the moment of contact 1971] Robinson & Mirick — Nephila clavipes 1 33 Table 2. Acridiids weighted and unweighted all lengthened Prey A to 70 mm. Prey Behaviors. yes 8 2 1. Bite and back off no 4 10 yes 11 4 2. Bite, legs raised no 1 8 yes 12 11 3. Wrap in web no 0 1 yes 4 8 4. Quit the web no 8 4 Prey A weights 1700-1800 mg. Prey B weights 400-550 mg. Statistical analysis — Fisher’s exact probability: Difference between A & B for bite and back off, level of probability 0.025; difference between A & B for bite, legs raised, level of probability 0.01 ; difference between A & B for wrap in web not significant; difference between A & B for quit the web not significant. All levels one-tailed with the prey, by tapping weighted and control dummies from behind the web. (We did this when the spider was in tarsal contact with the prey and before it had attacked). This led to a numerical, but non statistically significant, increase in the number of bite & back off attacks. It is very difficult to standardize simulated prey move- ments. A by-product of these experiments was the suggestion that the wrapping of prey in situ might be a response to the failure of the spider to pull out the prey. The number of wrap in situ responses was significantly greater in the case of the artificially lengthened prey used in the first experiment. These were, to the observer, obviously a much greater problem to the spider at the pull out stage. We therefore decided to carry out some experiments to see if increasing the adhesion of the prey to the web, or its ‘apparent* adhesion, would affect the spider’s behavior at this stage in the predatory process. ( b ) Pulling out and wrapping behavior As a first simple experiment we used domestic crickets as prey. We simply presented 20 dead unmodified crickets and scored the 134 Psyche [September Table 3. Wrap in situ behavior (i). Experiment with ‘winged’ and normal crickets. Wrap in web Pull out alone No result Winged crickets 18 1 1* Normal crickets 1 19 — * Spider disappeared before completion of pair. Behavior to live dragonflies matched in weight with crickets. Wrap in web Pull out alone 20 0 Time of persistent pull out attempts (before inception of wrap in situ.) averages. Winged crickets 194.5 secs Dragonflies 68.7 secs number of them that were wrapepd in situ and the number that pulled out and subsequently wrapped at the hub. We then matched the sizes and weights of these crickets to a second set to which we attached thin paper ‘wings’ at right angles to the long axis of the body. These dummies were presented in such a way that the ‘wings’ greatly increased the surface adhering to the web. We then scored the number of wrap in situ and wrap at hub responses. The results are shown in Table 3. The increased adhesion resulted in a highly significant increase in the number of prey that were subjected to being Wrapped and then cut out rather than being pulled out. The spiders made very persistent attempts to pull out the winged crickets, two succeeded and the remainder averaged 194.5 seconds of abortive pulling-out attempts. This is very interesting since the spiders started wrapping dragonflies (of lower weight) after only 69.7 seconds of pulling out attempts. (Testing these two sets of data-pull out times for dragonflies and winged crickets, with the Mann-Whitney U test, shows that the difference is significant; p is less than 0.001). We also carried out a further experiment on this aspect of the predatory process. In this case we passed a thread through the thorax of the crickets and presented twenty crickets with the thread hanging below the insect and twenty in which we passed the thread through the web and then held it from behind. In the second case we were able to prevent the spider from pulling the cricket from the web by exerting a force in the opposite direction. The results are shown in Table 4. Again the spiders that were unable to pull the prey from the web wrapped it at the capture site and then cut 1971] Robinson & Mirick — Nephila clavipcs 135 Table 4. Wrap in situ behavior (ii). Experiment with stringed crickets. Wrap in web Pull out alone Reject Experimentals 19 0 1 Controls 2 18 0 Experimentals were held by string from behind the web. it free. In this experiment the spiders made extended and persistent efforts to pull the prey before initiating the wrapping behavior. The process of free wrap proved to be much more difficult to elicit under experimental conditions. We had observed that this behavior occurred most frequently in the treatment of butterflies and moths. These could be freed from the web by pulling but were, nonetheless, bulky and cumbersome insects. Other cumbersome in- sects such as dragonflies adhered strongly to the web, were wrapped and then cut free of the web, and were thus largely trussed and packaged at this stage. They were not therefore suitable for experi- ments on free wrap behavior. We reasoned that free wrapping was a response to insects that could be removed from the web by pulling but were too bulky to be transported without becoming entangled in the web. The spider makes movements during the pulling out process that could enable it to gauge the bulk of the prey as it is removed from the web. These movements involve legs I & II. The tarsi are passed along the adhering margins of the prey and ease them away from the viscid elements of the web. Our observations on the removal of butterflies and moths from the web suggested that the cumbersomeness of detached prey was, to a large extent, a function of apparently chance factors. Thus it was partially dependent on the point on the prey body at which the spider exerted the pull out movements (i.e. the point at which the prey was held in the jaws), and partially a result of the orientation of the wing and body sur- faces in relation to the sticky elements of the web. (Some idea of the complexity of these factors can be gained by visualizing the process of picking up a randomly cast down book, by the spine, from a sticky surface. A winged insect, like the book, may open in a variety of ways, depending on where it is seized and where it is stuck down.) Starting from this point we tried to present a series of moths to the spiders with the entire dorsal surface of their wings adhering to the web but at differing orientations to the radii (and therefore the hub and the spider). We hoped that these would be seized at different points on the body and that our sole experimental variable 136 Psyche [September would be the ‘apparent’ bulk of the prey beneath the spider during the prey-removal process. Our aims proved to be extraordinarily difficult to achieve and the results, although suggestive, are not convincing. In an attempt to manipulate the bulkiness of a series of presentations of butterflies ( Anartia sp.) we trimmed the wings of half of them down to small (5mm long) stubs. We then matched the weights of pairs of intact and mutilated specimens and presented the pairs, successively, in random order to the spiders. Only the intact insects elicited any free wrap behavior, and only 24% of these were so treated. This result is not statistically significant. If the number of free wrap occurrences is compared not with the total number of presentations of intact butterflies, but with that number less the number wrapped in situ or subjectively scored as presenting below average bulk on removal from the web, the result is signifi- cant. However this depends on our subjective assessment of the bulk and is unsatisfactory. These results do not enable us either to accept or reject, with confidence, the hypothesis that the free wrap response is related to the bulkiness of the prey after its removal from the web. We have yet to design an adequate test for this. Discussion A number of features of the predatory behavior of Nephila clavipes are of interest from the comparative standpoint. The most im- portant of these, in our view, are the total reliance of the species on biting, as an attack strategy, and the fact that the spider does not store prey in the web at the capture site. Both these features represent marked differences from the behavior of araneids belonging to the genera Argiope, Araneus and Eriophora. Reliance on attack (immobilization) wrapping, as the principal means of attack, prob- ably extends to a much greater number of araneid genera. These aspects of the predatory behavior of N. clavipes have been discussed in some detail by Robinson, Mirick & Turner (1969). These authors suggested that “advanced” spiders would obtain at least two advantages from the addition of immobilization wrapping to their behavioral repertoire. They would be enabled to attack large and/or dangerous prey without closing to the potentially dangerous contact distance involved in biting, and also could achieve a considerable economy in time spent at the capture site in subduing the prey. The bite & back off attack behavior, that we have de- scribed for N. clavipes , immediately suggests to the observer that it is a danger-avoiding device. Our experiments on the stimuli that 1971] Robinson Mirick — Nephila clavipes 137 evoke this behavior gave results which are entirely consistent with this view. In fact increasing the weight of the prey eventually results in the suppression of attack behavior and its supercession by escape. We obtained evidence that N. clavipes would lose some large prey by escapes, during the prolonged process of a bite & back off attack. Prey of similar size presented to the much smaller Argiope argentata were not lost during wrapping attacks in any of our presentations. (The senior author has obtained similar results in comparing the performance of Nephila maculata and Argiope aemula ( Walckenaer) , dealing with large acridiids and melo- lonthid beetles.) We have also recorded injuries inflicted on N. clavipes during biting attacks on prey that had biting mouthparts and have one record of injury following the defensive kicking of an acridiid. The economy in time spent at the capture site that is a potential consequence of attack wrapping was examined in detail by Robinson, Mirick & Turner (1969), who suggested that a major factor in this economy was the ability of the spider to leave an attack-wrapped prey in situ after delivering a short bite. These authors argued that once the prey was wrapped the spider could safely leave it and not transport it to the hub until the bite had taken effect. With biting attacks, on the other hand, the bite could not be terminated until the prey had been safely subdued by its effects. The economy in time that results from this process is greatly exaggerated if one compares time spent in bite & back off attacks with time spent in wrapping attacks on similar prey by other spiders. (This comparison will be made in the projected comparative paper; it was not made by Robinson, Mirick & Turner (ibid) because the data for the attacks of Argiope species on very large prey was not then available). It is interesting that although N. clavipes wraps prey at the capture site it does not store them there. Once this type of wrapping be- havior has evolved it would seem but a short step to utilize it to enable the spider to interrupt the predatory process after the attack phase and defer the removal of the prey from the web, and its transportation, until later. This step, according to Robinson, Mirick & Turner (ibid) would be advantageous in circumstances where large numbers of prey arrived in rapid succession, or where prey left at the hub during an attack might be in danger of being stolen by kleptoparasites in the absence of the spider. At the time these authors suggested that N. clavipes may transport all prey to the hub because the depredations of kleptoparasites might be more diffi- cult to detect if prey were stored at a number of capture sites in a i3§ Psyche [September very large web. The arguments dependent on the effects of klepto- parasites are not easy to resolve. We have seen prey stolen from the hub, during the absence of the spider, on many occasions. We have also seen N. clavipes respond to, attack, and eat, kleptoparasites that were moving about near to the periphery of the web. Other hypotheses to explain the absence of capture-site food storage by N. clavipes include the possibility that this spider has not evolved a sufficiently efficient wrapping technique to safely allow the storage of prey in situ , or that the presence of prey packages in an already fine meshed web might render it more conspicuous, and hence avoidable, to flying prey. It seems worth stressing the fact that N. clavipes can be induced to wrap prey in situ if these are made difficult to remove from the web by pulling. This simple function of post immobilization wrapping at the capture site may have been obscured by the fact that such wrapping can serve other functions in the predatory strategy of other araneids. The function of simplifying the safe removal of prey from the web seems to us, on a priori grounds, to be basic and probably primary. Similarly the existence of free- wrapping behavior suggests that the trussing or packaging function of wrapping is an important one in its own right, and not merely the useful by-product of a process serving another function. Both these opinions derive from our study of the behavior of N. clavipes, and, as far as we know, were not anticipated by earlier studies of more “advanced” araneids. If we now consider the model shown in Figure I our earlier comment that this represents a very considerable simplification can now be expanded. We have detailed some of the behaviors that are not included in the model on pp. 132- 134. In our account of the ex- perimental side of our studies it is obvious that the investigation of the effect of external stimuli on the course of predatory sequences is not complete. Although we have shown that some behavior units can be brought into play in response to simple stimuli we have not shown that these are the only effective stimuli in all cases. We have made no progress at all in investigating the effect of internal factors on the behavior of the spider. In all these respects the model is inadequate, although it is already quite complex. We have also ignored any discriminations that may be involved in the termination of acts of behavior after they are brought into plav. The spider must, for instance, both start and stop the process of bite & back off, and start and stop the process of pulling out the prey. Peters ( 1931, 1933a, 1933b), in elegant studies of the behavior 1971] Robinson & AAirick — Nephila clavipes 39 of Araneus diadematus, was able to show that the change in stimula- tion brought about as a result of one behavior could be the trigger for the next behavior in the sequence. This approach has not yet been made in the case of N. clavipes. It should be productive. Bibliography Eisner, T., R. Alsop and G. Ettershank 1964. Adhesiveness of spider silk. Science, N.Y. 146: 1058-1061. Gertsch, W. J. 1948. The spider book. Comstock Press. Kaston, B. J. and E. Kaston 1953. How to know spiders. W. C. Brown & Co. Peters, H. M. 1931. Die Fanghandlung der Kreuzspinne ( Epeira diademata L.). Experimented Analysen des Verhaltens. Z. Vergl. Physiol. 15: 693-748. 1933a. Weitere Untersuchungen uber die Fanghandlung der Kreuzspinne {Epeira diademata Cl.). Z. Vergl. Physiol. 19: 47-67. 1933b. Kleine Beitrage zur Biologie der Kreuzspinne Epeira diademata Cl. Z. Morph. Okol. Tiere. 26: 447-468. 1954. Estudios adicionales sobre la estructura de la red concentrica de las aranas. Commun. Inst. Trop. El Salvador. 1: 1-18. 1955. Contribucions sobre la etologia y ecologia comparada de las aranas tejedoras tropicales. Commun. Inst. Trop. El Salvador. 1-2: 37-46. Robinson, M. H. 1969. Predatory behavior of Argiope argentata (Fabricius). Am. Zool. 9: 161-173. Robinson, M. H., H. Mirick and O. Turner 1969. The predatory behavior of some araneid spiders and the origin of immobilization wrapping. Psyche. 76: 487-501. Robinson, M. H. and J. Olazarri 1971. Units of behavior and complex sequences in the predatory be- havior of Argiope argentata (Fabricius) (Araneae: Araneidae). Smithsonian Contributions to Zoology. 1-36. Robinson, M. H. and B. Robinson 1970. Prey caught by a sample population of the spider Argiope argen- tata in Panama: a year’s census data. Zool. Journal. Linnean Soc. Lond. 49: 345-357. Shear, W. A. 1970. The evolution of social phenomena in spiders. Bull. Brit. Arachnol. Soc. 1 : 65-76. TRECI-IOBLEMUS IN NORTH AMERICA, WITH A KEY TO NORTH AMERICAN GENERA OF TRECHINAE (COLEOPTERA: CARABIDAE) By Thomas C. Barr, Jr. The University of Kentucky, Lexington Trechoblemus Ganglbauer is a genus of trechine beetles (Tre- chinae: Trechini: Trechina) previously known only from Europe and Asia. It formed the type genus of Jeannel’s “Serie phyletique de Trechoblemus T and is generally regarded as closely related to cavernicolous trechines in Japan, the Carpathians and Transylvanian Alps of eastern Europe, and eastern United States (Barr, 1969; Jeannel, 1928, 1962; Ueno and Yoshida, 1966). The large cave beetle genus Pseudanophthalmus Jeannel, with approximately 175 species in caves of ten eastern States, the monobasic genus Nea- phaenops Jeannel, from Kentucky caves, and the dibasic genus N ebonites Valentine, from Tennessee and Kentucky, are part of the Trechoblemus complex. The apparent restriction of Trechoblemus to Eurasia led previous investigators to conclude that, with respect to the richly diverse trechine fauna in caves of eastern United States, “there are no im- mediate, ancestral genera now present in North America” (Barr, 1969, p. 83). Although there is at least one edaphobitic (obligate in soil) species of American Pseudanophthalmus known (P. sylvaticus Barr, 1967), in the mountains of West Virginia, it has already lost eyes, wings, and pigment, and merely indicates that many of the “regressive” evolutionary changes in ancestral Pseudanophthal- mus may have taken place in the soil or deep humus before the beetles became restricted to caves. Most of the species of Pseuda- nophthalmus from eastern Europe (Barr, 1964) are also eyeless edaphobites. In March, 1971, I received a series of 5 trechines for determina- tion from Dr. Richard L. Westcott, State Department of Agri- culture, Salem, Oregon. These specimens were all taken in black light traps in the Willamette valley between Salem and Portland, northwest Oregon, and proved to belong to an undescribed species of Trechoblemus , the first species of the genus to be discovered in North America. There are at least two other zoogeographic links between the forests of the Pacific Northwest and those of the southern Appala- chians, when one looks at the carabid faunas as a whole. The 140 1971] Barr — Trechoblemus 141 Figure 1. Trechoblemus westcotti , new species. Hillsboro, Washington County, Oregon. Holotype male, length 4.6 mm. small, burrowing cychrines of the genus Maronetus are endemic to the southern Applachians, but are more closely similar to cychrine genera of the Pacific Northwest than they are to other eastern cychrines (Barr, in press). The subgenus Amerizus of the large genus Bembidion includes only two species, B. ( A .) oblongulum Herbst in the Northwest and B. (A.) wingatei Bland in the southern Appalachians (Lindroth, 1961). Both of these groups, like Tre- choblejnus-Pseudanophthalmus , are more or less humicolous or sub- terranean. Within the large, widely-distributed genus Pterostichus , the species of the California-Northwest subgenus Ilypherpes share a close similarity in appearance and habits with those of the Appa- lachian subgenus Haplocoelus (cf. Barr, 1969, footnote p. 80). Until both subgenera have been carefully studied, however, it is not possible to state whether these similarities are the result of phylogenetic relationship or convergence. The relatively few close ties between the Pacific Northwest and Appalachian forest carabid faunas do suggest, however, that the two faunas have been isolated from each other for a long time. The beginning of the period of isolation presumably began in the Miocene with the establishment of the Great Plains. If the Trechoblemus- like ancestors of Pseudanophthalmus arrived in North America via a Bering land bridge, their arrival was probably pre-Miocene. Cer- tainly the considerable diversity (approximately 20 species groups 142 Psyche [September Figure 2. Trechoblemus 'westcotti, new species. Hillsboro, Washington County, Oregon. Aedeagus of holotype male, length 0.92 mm., left lateral view. at present) and fairly wide distribution that Pseudanophthabnus ancestors had achieved prior to colonization of caves during the Pleistocene must have required a rather long period of time, a sup- position which at least accords with postulating an ancestral stock from the Northwest. Trechoblemus westcotti, new species Figures i, 2 Length 4.3-4.8 mm., mean 4.5 mm. Pale castaneotestaceous, head a little darker, usually with a faint, nebulous, darker fascia on apical half of elytra; integument pubescent, shining, microsculpture finely isodiametric on head and very finely transverse, iridescent, on pro- notum and elytra. Functional wings present. Head rounded; labrum distinctly but not very deeply bisinuate; surface with scat- tered, rather long pubescence, especially on sides and beneath; eyes feebly convex, about 0.3 mm. high X 0.2 mm. long; mentum fused to submentum, mentum tooth conspicuous and deeply emarginate; 1 1 -1 2 prebasilar setae. Pronotum transverse, about 1.3 times wider than long, disc convex, with long, rather dense pubescence; greatest width in apical fourth, widths at apex and base subequal to each other and also to length; sides arcuate in apical half, then con- vergent, distinctly sinuate in basal fifth; anterior angles prominent; hind angles large, sharp, prominent, a little less than right; ante- basal foveae large and deep; base entire; anterior marginal seta placed just behind greatest width, posterior seta just before hind angles. Elytra elongate-oval, 1.8 times longer than wide, broadly 1971] Barr — Trechoblemus 143 subconvex, distinct humeri present, prehumeral borders slightly oblique to mid-line; disc densely pubescent, microsculpture very line, nearly obsolete but perceptibly transverse and iridescent; inner three longitudinal striae more or less complete, fourth and fifth striae joining in apical third, sixth stria shorter than seventh, all striae somewhat irregularly punctulate and progressively shallower toward margin; intervals subconvex; apical recurrent groove rather short and subparallel to suture, joining third stria in a short crosier in advance of anterior apical puncture; humeral group of umbilicate punctures closely spaced against marginal gutter ; two discal punctures on fourth interval, anteriormost behind level of fourth umbilicate; apical triangle normal for genus. Antenna a little more than half total body length, with all segments more or less pubescent. Aedeagus of holotype 0.92 mm. long, gently arcuate, membranous above, with a ventral preapical boss, apex briefly and narrowly produced, finely reflexed at the very tip; copulatory piece single, concave toward right wall of internal sac, apically attenuate and rounded, about one third as long as aedeagus; parameres with five apical setae. Holotype male, Hillsboro, Washington County, Oregon, July 27, 1965, taken in a black light trap by Kenneth Goeden; deposited in collection of the State Department of Agriculture, Salem, Oregon. Four female paratypes, taken in black light traps in northwestern Oregon, as follows: Marion County: Salem, July 15, 1969; 10 miles northeast of Salem, August 9, 1969: Pudding River, 3 miles east of Woodburn, August 2, 1968. Washington County: 5 miles northeast of Newberg, August 1, 1966. Measurements of holotype: total length 4.6 mm., head 0.86 mm. long X 0.88 mm. wide, pronotum 0.84 mm. long X 1.08 mm. wide, pronotum 0.80 mm. wide at apex and 0.82 mm. wide at base, elytra 2.64 mm. long X 1.46 mm. wide, antenna 2.54 mm. long, aedeagus 0.92 mm. long. Discussion: Superficially T. westcotti closely resembles T. micros Herbst and T. postilenatus Bates, European and Japanese species, respectively. The apical recurrent groove, however, is less rounded than in those species, and the aedeagus with its less arcuate form, ventral preapical boss, and briefly produced, truncate, slightly reflexed apex is distinctive. The general form of the aedeagus recalls that of Trechoblemus micro phthalmus Ueno, from Takarajima, one of the Tokara Islands (Ueno, 1955, pp. 404-405, fig. 1), but T. micro- phthalmus is a smaller, brachypterous species with smaller eyes and smaller aedeagus. Among North American species of Trechinae 144 Psyche [September the only species with which it might readily be confused is Lasio- trechus discus (F.), which also has eyes, wings, pubescent integument, and similar coloration. L. discus , however, has large, convex eyes, a strongly cordiform pronotum, an apical recurrent groove which is oblique to the suture, and most of the longitudinal striae vanish before reaching the apex. Key to Nearctic Genera of Trechinae i. Eyes pubescent; form elongate-subparallel, depressed; head with deep, sulcate frontal grooves extended backward onto sides of head; West Indies Perileptus Schaum Eyes glabrous or absent 2 2(1). Form elongate-subparallel, depressed; head with deep, sul- cate frontal grooves extended backward onto sides of head; elytra with second longitudinal stria beginning at level of anterior discal puncture, third stria obsolete behind anterior discal; aedeagus with basal bulb open- ing between two lobes; color black or piceous; in North America known only from Panama to southern Mexico Cnides Motschulsky Elytral striation not as described; frontal grooves not sul- cate; basal bulb of aedeagus closed except for normal basal orifice 3 3(2). Mandibles with a premolar tooth between retinaculum and and mola; color black, form Trechus-li ke, elytral discal punctures not on fifth stria; in North America, known only from Panama Trechisibus Motschulsky Mandibles without a disinct premolar tooth 4 4(3). Elytra with anterior discal puncture on or near fifth stria OR if anterior discal puncture is absent, specimen micro- phthalmous, depigmented, from cave in Mexico 5 Elytra with anterior discal puncture on or near third (rarely fourth) stria OR if anterior discal puncture absent, then specimen is not from cave in Mexico .... 6 5(4). Color rufotestaceous ; eyes reduced to minute, pale areolae: form elongate, frontal grooves short, neck very narrow; inhabitants of caves in Mexico Mexaphaenops Bolivar Color black or piceous to pale piceous; eyes normal, or if reduced to pale areolae (Queretaro, Mexico), then frontal grooves normal, neck not very narrow; moun- 1971] Barr — Trechoblemus 145 tains or caves, Mexico to Costa Rica and possibly Panama Paratrechus Jeannel 6(4) . Eyes normal 7 Eyes absent or rudimentary; inhabitants of caves or of deep soil in mountainous regions 9 7(6). Color black or piceous; glabrous, polished integument; range: most of Canada, United States including Alaska (absent from southern Great Plains and Altantic and Gulf coastal plains), central highlands of Mexico Trechus Clairville Color more or less testaceous, integument densely pubescent 8 8(7). Darker, with prominent, blue-black fascia across elytra; pronotum cordiform; eyes very convex; longitudinal striae disappearing before apex; apical recurrent groove oblique to suture; eastern Canada and New England Lasiotrechus Ganglbauer Paler, elytral fascia nebulous or absent; pronotum trans- verse-subquadrate; eyes feebly convex; at least inner three longitudinal striae complete; apical recurrent groove subparallel to suture, conspicuously joining third stria; northwest Oregon . ... Trechoblemus Ganglbauer 9(6). Mentum separated from submentum by distinct suture; submentum with row of four long (prebasilar) setae; basolateral margins of pronotum with two or three prominent teeth; known only from caves in vicinity of St. Louis and Ste. Genevieve, Missouri X enotrechus Barr and Krekeler Mentum fused to submentum; submentum with transverse row of six to twelve setae; basolateral margins of pro- notum without teeth; range: east of Mississippi River 10 10(9). Head with one pair of supraorbital setae; form elongate, subconvex, elytron without anterior apical puncture (+ +o); last segment of maxillary palp shorter than penultimate segment; length 6-7 mm.; western Penny- royal plateau of Kentucky Neaphaenops Jeannel Head with two pairs of supraorbital setae (rarely more than two) ; last segment of maxillary palp subequal in length to penultimate segment 1 1 11 (10). Anterior tibia subglabrous on external face; anterior apical puncture of elytron very small, not setigerous ( + +0) ; 146 Psyche [September length 6-8 mm., elongate, more or less glabrous, convex; southeastern Kentucky (Estill, Jackson, Rockcastle, Pulaski, Clinton, Wayne, and McCreary counties) and adjacent Tennessee (Fentress County) Darlingtonea Valentine Anterior tibia rather densely pubescent on external face; anterior apical puncture setigerous, or if absent (Adair County, Kentucky), then posterior discal puncture also absent (+++, +O + , 00 +, or +00) 12 12(11). Antenna as long as body, attaining elytral apexes when laid back; length 5. 5-7.0 mm.; head and mandibles very large, all appendages elongate and slender; anterior discal punctures of elytra at level of second umbilicate AND prehumeral borders sharply oblique to mid-line; Cum- berland plateau margin from Jackson County, Kentucky, southwestward to Van Buren County, Tennessee Nelsonites Valentine 13(12). Right mandible with retinaculum 5-tuberculate, anterior two and posterior three teeth separated by a deep emargination ; apex of aedeagus more or less umbonate in ventral view, transfer apparatus a single median ventral sclerite; length 4.5-6.O mm., form robust, elytra usually more or less pruinose; southeastern Kentucky (Jackson, Rockcastle, Pulaski, northern Wayne and McCreary counties) Ameroduvalius Valentine Right mandible with retinaculum usually 2- to 4-tuber- culate, without a conspicuous gap separating teeth; apex of aedeagus not as described ; transfer apparatus of one or two sclerites, placed edgewise in internal sac Pseud anophthal m us Jeannel Pseudanophthalmus appears last because it is the largest and most variable genus of nearctic trechines, and the key has been con- structed to split off the other genera one at a time. The last couplet is the most inconvenient for rapid sorting because it requires re- moving a mandible and/or an aedeagus. As a practical matter this can usually be avoided because most species of Pseudanophthalmus do not occur within the range of Ameroduvalius , but most of those that do are either under 4 mm. in length or have obliquely sloping prehumeral borders. For Pseudanophthalmus species in the same size range as Ameroduvalius (4-5 mm., Wayne and McCreary counties, Kentucky), males can be readily distinguished by the 1971] Barr — Trechoble?nus H7 absence of the diagnostic, wing-like expansions of the aedeagus so characteristic of A. jeanneli (Valentine, 1952). The + ’s and — ’s in couplets 10 and 1 1 are a shorthand for presence ( + ) or absence (- — ) of the anterior discal, posterior discal, and anterior apical punctures, respectively, of the elytra. A Prospectus of the North American Trechinae The following classification is within the framework proposed by Ueno (in Ueno and Yoshida, 1966, footnote, p. 77). The starred (*) subtribes are not represented in North America. Other than adding newly described genera, I have diverged from the “series” classification of Jeannel (1928) only in putting Lasiotrechus into the Trechoblemus series. The “Darlingtonea series” is a new category for two genera described by Valentine (1952) and seen by Jeannel only a year before his death. He felt that Ameroduvalius was not closely related to any trechine genus known to him and had the impression that Darlingtonea was very close to Pseudanophthal- mus (Jeannel, in litt.). Quite probably Jeannel relied on the aedeagal figures drawn by Valentine (op. cit.), and was thus misled about the nature of the transfer apparatus in Darlingtonea ; instead of consisting of two spatulate sclerites, the Darlingtonea copulatory pieces are dorsal/ventral and isotopic, like the two halves of a box. Tribe Trechodini Subtribe Cnidina Cnides Motschulsky — 3 spp. ; southern Mexico to Panama Subtribe Trechodina* Subtrible Plocamotrechina* Tribe Perileptini Perileptus Schaum — 4 spp. ; West Indies Tribe Trechini Subtribe Aepina* Subtribe Aemaloderina Trechisibus Motschulsky — 1 sp. ; Panama (many spp. in South America) Subtribe Trechina 1) Trechoblemus series Trechoblemus Ganglbauer — 1 N.A. sp. ; Oregon Lasiotrechus Ganglbauer — 1 N.A. sp.; eastern Canada and New England 1 48 Psyche [September Pseudanophthalmus Jeannel — approximately 175 spp.; Ala- bama, Georgia, Illinois, Indiana, Kentucky, Ohio, Penn- sylvania, Tennessee, Virginia, West Virginia N ebonites Valentine — 2 spp.; Kentucky and Tennessee Neaphaenops Jeannel — -1 sp. ; Kentucky 2) Aphaenops series Xenotrechus Barr and Krekeler — 2 spp. ; Missouri 3) Darlingtonea series Darlingtonea Valentine — -i sp.; Kentucky and Tennessee Ameroduvalius Valentine — 3 spp.; Kentucky 4) Paratrechus series Paratrechus Jeannel — approximately 20 spp.; Mexico to Costa Rica Mexaphoenops Bolivar — 5 spp.; northeastern Mexico 5 ) T rechus series Trechus Clairville — -approximately 40 spp.; mountainous and northern regions, also central highlands of Mexico Literature Cited Barr, Thomas C., Jr. 1964. The status and affinities of Duvaliopsis Jeannel (Coleopterar Carabidae). Psyche, vol. 71, pp. 57-64. 1967. A new Pseudanophthalmus from an epigean environment in West Virginia (Coleoptera: Carabidae). Psyche, 74: 166-172. 1969. Evolution of the Carabidae (Coleoptera) in the southern Ap- palachians. In: Holt, P. C., ed. The distributional history of the biota of the southern Appalachians. Virginia Polytechnic Inst., Res. Mon. 1, pp. 67-92. 1971. In press. Studies on the cychrine beetles of eastern North America (Coleoptera: Carabidae). Bull. American Mus. Nat. Hist. Jeannel, Rene 1928. Monographic des Trechinae. Morphologic comparee et distribu- tion geographique d’un groupe de Coleopteres (3e livraison). L’Abeille, 35: 1-808. 1962. Les Trechini de l’Extreme-Orient. Rev. francaise d’Entomol., 29: 171-207. Lindroth, Carl H. 1961. The ground-beetles of Canada and Alaska, part 2. Opusc. En- tomol., suppl. 20: 1-200. Ueno, Shun-ichi 1955. Marine insects of the Tokara Islands. VII. New species and new subspecies of the subfamily Trechinae (Coleoptera, Harpa- lidae). Publ. Seto Mar. Biol. Lab., 4: 403-413. and Akira Yoshida 1971] Barr — Trechoblemus 149 1966. A presumptive prototype of the Trechoblemus complex (Cole- optera, Trechinae). Bull, Nat. Sci. Mus., Tokyo, 9: 75-83. Valentine, J. Manson 1952. New genera of anophthalmid beetles from Cumberland caves (Carabidae, Trechini). Geol. Surv. Alabama, Mus. Pap. 34, 41 pp. MECHANISMS CONTROLLING COPULATORY BEHAVIOR IN WOLF SPIDERS (ARANEAE: LYCOSIDAE)1 By Jerome S. Rovner Department of Zoology Ohio University Athens, Ohio 45701 Male spiders use their palps for picking up, storing, and, finally, transferring seminal fluid to the female’s copulatory apparatus. In previous papers (Rovner, in press) I described inter-generic differ- ences in palpal insertion patterns and examined temporal variation in the duration of insertion during mating in the lycosid spiders Lycosa rabida and Schizocosa saltatrix. Using the former species and taking a different perspective in the present study, I sought to determine some of the mechanisms involved in regulating the sequence of events accompanying each palpal insertion, as well as the role of the palps in the orientation of the male throughout mating. Experimental modification of one or the other partner was followed by the male’s performance of behaviors which did not occur during normal copula- tions: courtship, disorientation, tying down the female, and “pseudo- insertions”. Such data led to hypotheses concerned with the control of various elements of copulatory behavior. During mating, lycosid spiders maintain a position (Position II of Gerhardt, 1924) in which the male’s sternum is above the female’s carapace and the partners face in opposite directions (Fig. 1). Each insertion involves the male’s leaning down on one side of the female and scraping one palp (the one closest to that side of the female) against her epigynum. The male’s right palp serves the female’s right copulatory pore; his left palp, her left pore. One or more scrapes of the palp result in engagement of the em- bolus in the copulatory pore. The latter is accompanied by hema- todochal expansion, which forces the embolus into the duct leading to the seminal receptacle. Ejaculation of seminal fluid through the embolus is presumed to occur at maximum expansion of the hema- todocha (Gering, 1953). Subsequent collapse of the hematodocha is followed by disengagement of the embolus and lifting of the palp away from the epigynum. (For details of palpal function during copulation, see Gering, 1953. The hydrostatic system involved in hematodochal expansion, as well as in locomotion, has been studied recently by Wilson, 1970.) This study was supported in part by Ohio University Research Grant 244. Manuscript received by the editor , September 27 , 1971. 150 1971] Kovner — Wolf Spiders I5i Fig. 1. Male and female Lycosa rabid a in copula. The male is above, facing the camera, and has just initiated a pseudo-insertion with his left palp. After about half of the palpal insertions, male L. rabida moisten the palp just used by drawing it between the chelicerae. In such cases, they then usually moisten the opposite palp. Although a bout of palpal moistening often involves a rapid alternation of the palps, it always is initiated in the palp which has just been used in an at- tempted or completed insertion (Rovner, in press). In about half of the insertion sequences the male, rather than moisten the palps, either remains inactive for several seconds or immediately crosses over to the female’s opposite side. Whether or not palpal moistening occurs, male L. rabida then shift over to the female’s opposite side; i.e., palpal alternation is the insertion pattern in this species (Montgomery, 1903). While moving from one side to the other, the male usually taps his palps against the anterior dorsal surface of the female’s abdomen. At this time the female performs the only behavior shown by her during most of the copulation — abdominal swiveling. Her abdomen rotates about its longitudinal axis during each crossing by the male, thereby bring- ing the epigynum within reach of the male’s palp. The above palpal insertion sequence, which is repeated an average of about sixty times by each palp during copulation in L. rabida , is 152 Psyche [September LEAN SWIVEL ABDOMEN co LU K c n o CL CL o o H =F V V. SCRAPE PALP I ENGAGE EMBOLUS I HEMATODOCHAL EXPANSION I HEMATODOCHAL COLLAPSE I REMOVE EMBOLUS 1 MOISTEN SAME PALP 4 MOISTEN BOTH PALPS J Fig. 2. Sequence of events associated with each palpal insertion during copulation in Lycosa rahida. summarized in Fig. 2. Some of the mechanisms associated with these events were suggested by the findings of the present study. Specific questions that I was asking were these : ( i ) Given that the palps serve for transfer of sperm, how essential is sensory information from the palps to the maintenance of the male’s copulatory state? (2) Given that the palps may be aiding in a. sensory capacity to locate a target, the female’s epigynal openings, is information from the palps important for the male’s orientation on the female? (3) Does the pattern of right-left alternation persist after unilateral mod- ification of the male, or will the spider learn to favor the functioning palp? (4) What stimulus determines which palp is moistened first after each insertion? (5) What action by the male elicits abdominal swiveling in the female? Methods Over one hundred individuals of L. rahida were collected as im- mature instars during June, 1968, in a field near Athens, Ohio, 1971] Rovner — Wolf Spiders 153 U. S. A. Each was housed separately in a glass jar prior to pairing in the observation arena. Mealworms ( Tenebrio sp.) served as food ; and cotton-stoppered, water-filled vials provided moisture. Observations on copulatory behavior were made at temperatures of 24-26°C. A manually activated Esterline Angus event recorder, at a chart speed of 15.3 cm/min, was used to record some of the data. Protocol was whispered into the microphone of a tape recorder. Recording instruments were placed on a separate table to reduce possible effects of machine noise. This study involved fifty pairings of virgin, adult spiders. (Al- though housed individually until that time that the mating partners were placed into the arena, the spiders will be referred to in terms of this eventual pairing.) Ten pairs of spiders were not modified experimentally; and their behavior represented that of normal in- dividuals. The remaining forty pairs were divided into the following eight groups, each consisting of five pairs: (1) Males losing both palps prior to the final molt; (2) Males losing both palps after the final molt; (3) Males with both palps fixed dorsally; (4) Males losing one palp prior to the final molt; (5) Males losing one palp after the final molt; (6) Males with one palp fixed dorsally; (7) Females with both copulatory pores sealed; (8) Females with one copulatory pore sealed. Palp removal was accomplished by autotomy. During carbon dioxide-induced anesthesia, the male’s palp was attached to the sub- stratum. After the male’s recovery I prodded him with an artist brush and forced him to pull away from the point of attachment, which resulted in palpal autotomy at the trochantero-femoral joint. This was repeated for males undergoing loss of both palps. When autotomy involved penultimate males, a “stump” or a complete but vestigial palp was present after the final molt. In the latter case the tarsus lacked a genital bulb. Fixation of a palp above the cephalothorax involved positioning the palp into a drop of melted paraffin placed on the adjacent region of the carapace of the anesthetized spider. Paraffin was also used to cover the copulatory pores of anesthetized females. All operations were performed under a dissecting microscope. In each experimental group of five pairs which involved unilateral modification, three individuals were treated on one side (e.g., right palp) and two on the other side (e.g., left palp). Results A variable period of time after mounting the female, many of the males of the experimental pairs performed behaviors which were not 154 Psyche [September Fig. 3. Male Lycosa rabida disoriented 180° from the normal position while above the female in copula. observed during copulation in the normal pairs in this or previous studies of lycosid mating behavior. Males unable to use one or both palps (due to modification of the males) performed courtship display at various times while mounted on the female. Those males possessing one usable palp often followed each successful insertion with a brief period of courtship that was initiated after the shift to the non-functional side and the subsequent adoption of a medial position. Thus the male alternated copulation with courtship. In some cases, courtship was initiated soon ofter mount; e.g., one male began to display after having suc- cessfully inserted the functional palp only three times. Males possessing one usable palp always shifted either to a medial position or completely over to the non-functional side after each successful insertion. However, when visiting the non-functional side, these males typically did not lean ventrad as steeply as did normal males. After resting momentarily, the experimental males either shifted back immediately to the functional side or initiated a bout of courtship display. The average duration of insertion of the func- tional palp in these males ranged from 7.1 sec in one male to 15.8 in another. 1971] Kovner — W olf Spiders 155 Fig. 4. Palpless male Lycosa rabida disoriented from the normal position and using his black legs I to perform elements of courtship display while above the female in copula. Disorientation was shown by most of the males which lacked the use of both palps (due to modification of the males). In this be- havior the male pivoted on the female’s carapace in a clockwise or counter-clockwise direction and temporarily adopted a position other than the normal one. The most typical and most prolonged abnormal position was that in which the male was 1800 out of proper align- ment, i.e., facing in the same direction as the female (Fig. 3). The next most common abnormal position was one of 90° mis-alignment, i.e., the male’s longitudinal axis perpendicular to that of the female. As “copulation” progressed, the male adopted positions even further removed from the normal one by locating himself above the basal segments of the female’s legs on one side (Fig. 4). While in the various abnormal positions, the male spent much of his time in courtship display. Whether in the normal position or disoriented 180°, males lack- ing the use of both palps (due to modification of the males) showed bouts of rapid, oscillatory movements, in which the male either shifted from side to side or slid forward and back while above the female’s carapace. Such bouts of activity were variable in duration, on the average lasting 6.8 sec (N = 140). During these excited 156 Psyche [September movements, the male scraped his chelicerae against the female's cara- pace (or, rarely, abdomen) and alternately spread and closed the chelicerae as they slid over the female’s surface. Some of these bouts terminated with the male’s use of his chelicerae to grasp the rim of the female’s carapace or a basal segment of one of her appendages and to lift that end of her body dorsad. Synchronous erection of his leg spines accompanied each tug by the male. The female did not respond to the male’s tugs, even though: (i) males disoriented i8o° tugged at structures near the female’s face; and (2) puncture of a joint membrane and loss of a drop of hemolymph occurred in a few females due to the male’s vigorous cheliceral grasping. After one or more tugs, the male became inactive but maintained his hold on the female with his chelicerae. The pattern of behavior in males lacking the use of both palps was that of an alternation of inactive periods (with the male located medially or leaning slightly to one side of the female) with periods involving one or more of the above- described behaviors. Abdominal vibrations continued during the male’s inactive periods. Half of the males with both palps unavailable and one-fifth of the males with one palp unavailable occasionally performed “tying down”. In this behavior the male pivoted on the female’s carapace in a clock- wise or counter-clockwise direction while laying down a barely visible silk line over the legs and abdomen of the female. The silk on the legs usually contacted the patellar and tibial segments; less fre- quently, the femora. At a few points in the crude circle, the silk was attached to the substratum. During tying down the male typ- ically moved through a complete 360° and resumed the normal posi- tion. Less often the male momentarily stopped at 1800 and then returned to the normal position by either continuing to 360° or re- versing direction and covering the same ground again. Even though the male stopped briefly at 180° (or, rarely, 90° or 270°), he did not initiate any other behavior at these points, but soon resumed tying down in a return to the normal position. Thus, tying down was distinct from the behavior described above as disorientation, in that the latter: (1) did not involve release of silk; (2) involved pivoting through arcs of 180° or less; and (3) usually was accom- panied by other behaviors while the male was in an abnormal posi- tion. 'Lying down was shown in pairs in which the female was inactive throughout the copulation (except for abdominal swiveling) ; i.e., tying down typically was neither preceded nor followed by activ- ity in the female. When resuming locomotion at the end of the 1971] Rovner — Wolf Spiders 157 mating, females which had undergone one to several bouts of tying down were only slightly hampered by the scant threads on their legs and readily freed themselves. Most of the normal males copulating with females that had one or both copulatory pores sealed performed an atypical behavior which I have termed a “pseudo-insertion”. This occurred when one of the palpal scrapes (insertion attempts) on an unavailable side resulted in complete expansion of the hematodocha, even though the palp was no longer in contact with the female’s body. Synchronously with the hematodochal expansion, the palp was lifted dorsad, sometimes to a relatively high position (with the palpal femur about 450 above the horizontal plane). Leg spine erection accompanied hematodochal expansion, as in normal insertions. The palp was lowered ventrad (and the leg spines dropped) during the subsequent hematodochal collapse. Pseudo-insertion duration averaged 15.5 sec (N = 20). Pseudo-insertions began to occur during a copulation after the male had made a number of visits to an unavailable side of the female. Dur- ing each visit males made from one to ten (usually from two to five) attempts to insert. A pseudo-insertion usually occurred after live or fewer attempts; in some cases later in copulation, the first attempt during a visit to one side resulted in a pseudo-insertion. In many cases if the hematodocha began expansion after partial engagement of the embolus in the paraffin seal, expansion would continue to com- pletion after the palp slipped away from the epigynum. In other instances the palpal tarsus would swing down to the base of the female’s leg IV and, meeting resistance there, give rise to a pseudo- insertion. The number of pseudo-insertions performed during such pairings ranged from one to seventeen. Two males lacking one palp and having difficulty inserting the available palp showed pseudo-insertions, as well as courtship and tying down. Males having one or both palps fixed dorsad did not perform pseudo-insertions with their treated palps. A behavior probably related to the mechanism underlying pseudo- insertions was also seen in the males which were paired with females having one or both copulatory pores sealed. It occurred while the male was resting on one side of the female after a series of insertion attempts. (Many of these attempts involved the male’s leaning steeply and scraping his palp much further posterior than the female’s epigynum, sometimes thereby contacting her spinnerets.) During this resting period, while the male held his palp near his face, slow and weak pulsations were evident in the genital bulb. During this “throb- bing” the hematodocha was not visible. Partial, synchronous leg spine erections accompanied the genital bulb pulsations. 158 Psyche [September Table I. Behaviors shown ; during copulation by males of treated and untreated pairs of Lycosa ra bida. (The first figure represents the number of males showing that behavior ; the next figure, the number Courtship Disorientation Both $ palps of males tested.) Tying Pseudo- down insertion lacking Both $ palps 7/10 6/10 3/10 fixed dorsad One $ palp 5/5 4/5 4/5 0/5 lacking One $ palp 4/10 1/10 2/10 2/10 fixed dorsad Both $ 4/5 1/5 1/5 0/5 pores sealed One $ 1/5 0/5 0/5 4/5 pore sealed $ and $ 2/5 0/5 0/5 4/5 untreated 0/10 0/10 0/10 0/10 Many of the above results are summarized in Table I. Data for males treated prior to the final molt are grouped with those for males treated similarly after the final molt. There did not seem to be a difference in performance between the pre- and the post-molt-treated males. Further reduction of the data is provided in Table II, in which experimental pairings are reduced to three classes of treatment. Details of palpal moistening behavior were studied in the various experimental pairs. Males entirely lacking one palp moistened the available palp only after using it in an insertion or an insertion attempt. They did not perform moistening of the available palp after visiting the palpless side. Some males of this experimental category possessed a vestigial palp (complete but smaller and lacking a genital bulb). Insertion attempts with the latter were accom- panied by synchronous leg spine erections, but were not followed by palpal moistening. On the other hand, after insertions or insertion attempts of the functional palp, both palps were involved in bouts of palpal moistening. In pairings of normal males with females having sealed copulatory pores, many of the groups of insertion at- tempts culminated in a bout of palpal moistening, which always began with the palp used in the attempts. Males having both palps fixed dorsad and males lacking both palps performed behavior associated with moistening of the unavailable palps. After adopting a medial position above the female’s carapace and tilting his body caudad, such a male alternately spread and closed his chelicerae for a period of time in a manner similar to that 1971] Rovner — W olf Spiders 159 Table II. Percentages of male Lycosa rabida performing various behaviors during copulation. (This summary was obtained by appropriate re-groupings of the data presented in Table I.) Both $ palps % Courtship % Disorientation % Tying down % Pseudo insertion unavailable One $ palp 80.0 66.7 46.7 unavailable One or both $ 53.3 13.3 20.0 13.3 pores unavailable $ and 9 30.0 0.0 0.0 80.0 untreated 0.0 0.0 0.0 0.0 seen during normal moistening. Likewise, those males in which only one palp was fixed dorsad alternated moistening of the available palp with periods in which the cheliceral movement continued without the available palp being drawn between the chelicerae. Males unable to insert one palp due to unilateral modification of themselves or their partners usually shifted over to that unavailable side after a successful insertion. After a bout of attempted insertions and palpal moistening or a period of inactivity on the unavailable side, the male shifted back to the functional side. In other words, throughout most of the copulation such males maintained a pattern of right-left alternation and did not remain on one side or the other for an extensive period of time. Towards the end of the mating, some of the males paired with females having one pore sealed did tend to perform an increased number of insertion attempts on the sealed side. Another irregularity in the right-left pattern which occurred occasionally in males of various experimental pairings was the attempted use of the “wrong” palp while on one side of the fe- male. For example, while on the female’s right side, the male would press his left palp against the base of the female’s right leg IV and achieve a partial (or, rarely, complete) hematodochal expansion. Females paired with males lacking both palps or males having both palps fixed dorsad showed appropriate abdominal swiveling when the male shifted from one side to the other. Abdominal swiveling was also elicited experimentally in females that had remained in the so-called cataleptic state after the male’s dismount, as well as in a few females which had already resumed an active state, e.g., a de- fensive posture. In the latter instances, the females returned to the passive copulatory condition when I pressed down on their posterior carapace with a probe. Abdominal swiveling was subsequently elicited in the below-described manner. It was also possible to elicit abdom- i6o Psyche [September Fig. 5. Female Lycosa rahida being tested after the male’s dismount. She has responded to tactile stimulation of the left posterior region of her carapace by swiveling her abdomen so as to raise the left side dorsad. (Note that the medial dark band on her abdomen is in line with the right dark band on her carapace.) inal swiveling in a few females that had initiated catalepsis prior to the male’s mount in apparent response to the male’s courtship dis- play alone. In all of these cases, tactile stimulation of various regions of the female’s cephalothorax and abdomen with an artist brush or a probe revealed that maximal abdominal swiveling was elicited by touching the posterior quarter of the carapace. (Tactile stimulation of the more anterior portions of the carapace or of the anterior lat- eral sides of the abdomen yielded weaker swiveling responses.) Touch- ing the right side of the posterior carapace resulted in a single ab- dominal movement which brought the right epigynal pore to a more dorsad position; touching the left posterior carapace elicited the cor- responding abdominal swiveling (Fig. 5). I could go from side to side with the probe in this manner and elicit appropriate abdominal movements in relatively rapid succession. If both sides of the fe- male’s carapace were pressed simultaneously, her abdomen was held medially and swiveled very slightly from side to side in alternation. Copulations in most experimental categories were similar in dura- tion to those observed in normal pairs (average duration for the lat- 1971] Kovner — Wolf Spiders 1 6 1 ter = i hr). Males having both palps fixed dorsad tended to remain on the female an average of 2 hr. (N = 5). The data presented in Tables I and II reflected in some cases the influence of variables other than those suggested by the type of ex- perimental treatment. A few of the males with one palp unavailable were unable to achieve insertion with the remaining palp. As the hematodocha approached maximal expansion, a drop of hemolymph issued from some point in or near the palpal tarsus. This drop, roughly having a diameter as great or greater than the width of the palpal tarsus, was then sucked up by the male. In such cases the male performed only a few insertions and thereafter behaved as did males lacking the use of both palps. (Although such a malfunction seemed to be related to the presence in such males of only one usable palp, I later observed such a “leaking palp” in a normal male.) Level of activity in the female was another variable in these ex- periments. In the majority of pairings, the females remained inactive (except for abdominal swiveling) throughout copulation. However, a few females initiated periods of locomotor activity (including car- rying the male to another part of the arena), particularly those fe- males whose partners had both palps unavailable. The frequency or duration of various behaviors in the male may have been influenced by such behavior in the female. Discussion Sensory information from the palps was important for maintenance of the male’s copulatory state. While mounted on the female, males unable to use one or both palps (due to modification of the male) performed courtship displays and bouts of tying down. In normal males proprioceptive feedback from insertions probably reduced the likelihood that the male would switch back to an earlier behavior, courtship display, or switch to an out-of-context behavior, tying down. (A behavior similar to tying down sometimes occurred in normal, solitary individuals in response to recently captured large prey such as adult field crickets, Gryllidae.) It is unlikely that tying down served to facilitate the male’s post-copulatory escape, in that : ( 1 ) fe- males were only slightly, if at all, hindered by the few threads of silk; and (2) normal males did not employ any such device during mating but did use a cheliceral pinch at the time of dismounting to aid in escape (Rovner, in press). Among normal males paired with females having one or both copulatory pores sealed, three males showed courtship and none, tying down. Thus, even though unable to insert one or both palps, these males generally did not switch to inappropriate behaviors nor Psyche [September 162 did they become disoriented. It is likely that: (1) their ability to achieve numerous, partial hematodochal expansions during insertion attempts provided sufficient feedback for maintenance of the copula- tory mode, and (2) the availability of the palps for sensory purposes provided input for orientation purposes. The early occurrence of courtship after mount by some of the males lacking one palp suggested that there was a relatively inde- pendent insertion tendency for each palp. Even though able to suc- cessively insert the remaining palp, such males were still motivated to insert the absent palp and, when unable to do so, switched back to pre-copulatory display behavior for a period of time rather than continue to use the functional palp. A relative independence of insertion tendency for each palp was also suggested by the change in behavior towards the end of copula- tion in males paired with females having one copulatory pore sealed. Such males continued their insertion attempts on the unavailable side but decreased their visits to the functional side. This indicated that the motivation underlying insertions of the functional palp had decreased to a minimal level as a result of numerous successful insertions. Peripheral input from the palps seemed important for the male’s positioning on the female. Two- thirds of the males in which both palps were unavailable showed disorientation at various times while above the female. Since disorientation usually involved a 180° mis- alignment, it seemed likely that input from other parts of the body, probably the sternum and legs, served for maintenance of a medial position (above and in line with the female) and that input from the palps determined the correct anterior-posterior orientation. This is quite different from the control system in a linyphiid spider studied in this regard (Rovner, 1967). Palpless male Linyphia triangularis maintain a normal position throughout “copulation” (both partners hanging inverted, facing in opposite directions, and the male’s head pivoting on or near the female’s chelicerae). In the latter case, in- formation from tactile hairs on the anterior dorsal point of the male’s pars cephalica (and probably the legs also) suffices for orientation. Most of the males copulating with females whose genital openings were sealed performed pseudo-insertions. This behavior was a re- sponse to sub-normal releasing stimuli, in that the entire cycle of events associated with a palpal insertion occurred without successful engagement of the embolus. It seemed that the threshold for release of complete hematodochal expansion gradually lowered during a succession of insertion attempts. At some point, an initiated expan- sion went to completion without the normally necessary engagement 1971] Rovner — W olf Spiders 163 of the embolus. One interpretation is that the motivation for palpal insertion built up under the maximal stimulation of repeated, un- successful insertion attempts and that the complete behavior finally was released. Theoretical considerations aside, pseudo-insertion be- havior may be of interest to workers studying the mechanics of palpal insertion in spiders. In a pseudo-insertion the entire genital bulb, which is only partly visible when pressed tightly against the female’s epigynum during a normal insertion, is brought into view during hematodochal expansion and collapse. In a previous paper on L. rcibida (Rovner, in press) 1 had em- phasized that the palp just used in an insertion was the palp em- ployed first in a bout of palpal moistening. The question remained as to what aspect of the insertion sequence served as the stimulus determining the latter. Three observations in the present study pro- vided evidence for an hypothesis: (1) Males lacking one palp moist- ened the remaining palp only after insertions or insertion attempts with the latter. (2) Males with vestigial palps (lacking genital bulbs) did not initiate moistening of these palps after using them in attempted insertions. (3) Normal males unable to insert one or both palps (due to closure of the female’s copulatory pores) moistened the palp just used in an insertion attempt. Apparently the stimulus normally releasing palpal moistening was associated with the onset of hematodochal expansion, which accompanied the palpal scrape of an insertion attempt. (According to Gering, 1953, hematodochal expansion is initiated as the palp swings downward at the outset of an insertion attempt.) Thus, the proprioceptive stimuli associated with the initial phase of an insertion probably were the ones deter- mining which palp would initiate the subsequent bout of palpal moistening. The occurrence in palpless males and males with their palps fixed dorsad of cheliceral movements like those seen during palpal moisten- ing indicated that the onset and maintenance of this behavioral pat- tern did not depend on stimuli resulting from the grasping of the palp by the chelicerae. The “chewing” movements occur in vacuo . A similar phenomenon was reported in palpless male Linyphia tri- angularis (Rovner, 1967). Persistence of the pattern of right-left alternation in males unable to insert one of their palps suggested that this pattern was not altered by the experience of repeated failures on the non-functional side. At no point in copulation did males begin to favor the function- ing palp by remaining on that side of the female. Males typically shifted, however briefly, to the non-functional side, a behavior which, in normal males, initiated another palpal insertion. 164 Psyche [September Various evidence gathered in this study indicated that abdominal swiveling was elicited in the female primarily by the male’s leaning against the side of the female’s posterior carapace. Palpal tapping, one of the most obvious aspects of the male’s behavior during his shift to the opposite side, had been suggested to be the stimulus re- leasing abdominal swiveling (Hallander, 1966). However, the latter seemed to play a minimal role, if any, in triggering the female’s response. (Perhaps palpal tapping, if it did indeed have a function, represented an exploratory behavior aiding the male in his orientation on the female.) The observation that simultaneous stimulation of both sides of the female’s carapace resulted in a medial abdominal position (with weak side-to-side swiveling) suggested that bilateral input resulted in a bilateral response, in which the motor mechanism for both sides was activated. Summary Studies involving experimentally modified Lycosa rabida suggested some of the mechanisms controlling various aspects of copulatory behavior. Both maintenance of the copulatory state and proper positioning of the male depended on input from the palps. There existed an independent insertion tendency for each palp. Males with modified palps showed courtship, tying down, and disorienta- tion. Pseudo-insertions occurred in normal males paired with females having sealed copulatory pores. Inability to insert one palp did not eliminate the pattern of shifting from side to side on the female. Events associated with partial hematodochal expansion provided adequate stimuli for eliciting moistening of the palp. Cheliceral movements typical of palpal moistening occurred in palpless males. Mechanical stimuli resulting from the male’s leaning against one side of the female’s carapace released her abdominal swiveling re- sponse. Literature Cited Gerhardt, U. 1924. Weitere Studien iiber die Biologie der Spinnen. Arch. Natur- gesch. 90: 85-192. Gering, R. L. 1953. Structure and function of the genitalia in some American age- lenid spiders. Smithsonian Inst. Misc. Collect. 121, No. 4. Hallander, H. 1967. Courtship display and habitat selection in the wolf spider Par- dosa chelata (O. F. Muller). Oikos 18: 145-150. Montgomery, T. H. Jr. 1903. Studies on the habits of spiders, particularly those of the mating period. Proc. Acad. Nat. Sci. Philad. 55: 59-149. 1971] Kovner — W olf Spiders 165 Rovner, J. S. 1967. Copulation and sperm induction by normal and palpless male linyphiid spiders. Science 157: 835. In press. Copulation in the lycosid spider Lycosa rabida Walckenaer: a quantitative study. Anim. Behav. In press. Copulation in the lycosid spider Schizocosa saltatrix (Hentz) ; an analysis of palpal insertion patterns. Anim. Behav. Wilson, R. S. 1970. Some comments on the hydrostatic system of spiders ( Cheliccratay Arancae ). Z. Morph. Tiere 68: 308-322. THE LARVA OF IIELIOCAUSUS LARROIDES (HYMENOPTERA, SPHECIDAE) By Howard E. Evans Museum of Comparative Zoology Heliocausus is one of several interesting genera of sphecid wasps wdiich has been known only from the adult stage. I therefore wel- come the opportunity to describe larvae of H. larroides (Spinola) sent to me recently by Manfredo A. Fritz of Buenos Aires, Argen- tina. My description is based on four fully grown larvae from El Salto, Valparaiso, Chile, collected by Dr. Fritz on 24 December 1969. I shall defer a discussion of larval characters until after the description. Body. — -Length 9 mm; maximum width 2.8 mm. Fusiform, somewhat curved anteriorly ; pleural lobes moderately prominent ; terga indistinctly divided into two annulets each ; apical abdominal segment rather slender and protuberant, the anus terminal (Fig. 2). Spiracles weakly pigmented, inconspicuous; atrium lined with weak, irregular polygons; opening into subatrium unarmed; subatrium abruptly widened and much folded just beneath atrium (Fig. 6). Integument mainly smooth and without setae or spinules, but ex- treme anterior part of prothorax, the “neck” region, densely spinulose, also thoracic venter somewhat spinulose and dorsum with sparse, minute setae, the largest about 20 /i long. Head. — Subcircular, width .75 mm; height (exclusive of labrum) about the same; front of head with a pair of longitudinal, welt-like elevations just mesad of the antennae; coronal suture and parietal bands weakly developed ; front surface of head weakly pigmented, light brown (Fig. 1). Antennal orbits circular, about 85 /x in diameter, antennal papillae barely longer than thick, 25 [i long (Fig. 7). Head very sparsely punctate, some of the punctures near the vertex, on the lower sides, and on the clypeus bearing short setae. Mouthparts. — Labrum with about 24 strong setae as well as a subapical row of 16-18 small sensilla; apical margin spinulose, especially laterally; epipharynx mainly papillose, but the papillae grading into spinules medio-basally and laterally, also with a. few small sensilla (Fig. 3). Mandibles approximately twice as long as their maximum width, with a single rounded tooth on the inner margin in addition to the apical tooth ; base with a single lateral sensillum (Fig. 5). Maxillae densely spinulose along the mesal 1971] Evans — Larva of I~I eliocausus 167 Larva of H eliocausus larroides (Spinola). Fig. 1. Head, anterior view. Fig. 2. Body, lateral view. Fig. 3. Labrum (left) and epipharynx (right). Fig. 4. Maxilla. Fig. 5. Mandible. Fig. 6. First thoracic spiracle. Fig. 7. Antennal orbit and papilla. Psyche [September 68 margin ; palpi strong, about 80 /jl long, terminating in three large sensory cones; galeae more slender, about .7 as long as palpi, ter- minating in a single large sensory cone (Fig. 4). Labial palpi slightly shorter than maxillary palpi, slightly exceeding the blunt, paired spinnerets; oral surface of prementum with a patch of small spinules. Discussion. — Despite the larrine-like appearance of adults of this wasp (giving rise to the specific name larroides) , the larvae bear no resemblance to those of Larrinae. The presence of an antennal papilla, the terminal anal opening, the form of the mandibles, and several other features indicate that Heliocausus belongs in or near the subfamily Nyssoninae. In fact, it runs readily to Nyssoninae in my table of subfamily characters (Evans, 1959, p. 168), and in my artificial key to genera (ibid, p. 1 7 1 ) the genus keys out near Sphecius and Gorytes (Nyssoninae, Gorytini). The mandibles, labrum, and maxillae are especially similar to those of Gorytini. It should be added that Dr. Fritz found II. larroides preying upon leaf- hoppers, a distinctive (although not exclusive) gorytine attribute. On the other hand, there are certain larval features not shared by any known Gorytini: the oral surface of the prementum is spinulose (as in Bembicini), the opening between the spiracular atrium and subatrium is unarmed (as in Philanthinae and a few primitive genera of Nyssoninae), and the apical abdominal segment is slender and protuberant (as in Philanthinae). I regard the Nyssoninae and Philanthinae as having had a common origin inde- pendent of other subfamilies (ibid, p. 183). While on the whole Heliocausus has many more larval characters in common with the Nyssoninae, the presence of some philanthine characters and some in common with generalized Gorytini suggests that the group may be a relict of a primitive nyssonine-philanthine stock. Manfredo Fritz informs me that he and Prof. H. Toro of Valparaiso have come to somewhat similar conclusions on the basis of adult structure ; their paper is soon to be published in the Ann. Mus. Hist. Nat. Valparaiso. Reference Cited Evans, H. E. 1959. Studies on the larvae of digger wasps (Hymenoptera, Sphecidae). Part V: Conclusion. Trans. Amer. Ent. Soc. 85: 137-191. FLIGHTS OF THE ANT FORMICA DAKOTENSIS EMERY* By Mary Talbot The Linwood Colleges, St. Charles, Missouri 63301 Formica dakotensis is a small ant of the rufa group whose red and black workers are easily recognized by the petiolar scale, which is thick and flat on top and has sides that are parallel along the upper half and taper inward only in the lower half. This is the first report of F. dakotensis in Michigan. However, its presence here is not surprising since it has been taken in Indiana and Wisconsin. Other records have been west of this range. Al- though ant collecting has gone on at the Edwin S. George Reserve in southern Michigan (Livingston County) since 1951, it was not until August 20, 1969 that a colony of this ant was found. Subse- quently, two other nests have been located on the Reserve and W. F. Buren discovered a colony in Branch County, near the Ohio border. The Edwin S. George Reserve is a two square mile inclosure of rolling country with woods, fields, swamps and marshes. The three colonies (in grids G22, N26, and Cio of the Reserve map) each lay near the border of a field, close to a good growth of shrubs, such as Spirea alba Du Roi or Cornis stolonifera Michx., which grew near a swamp edge. This paper is primarily concerned with one colony, which was studied during the summers of 1970 and 1971 to determine its flight activities, nest structure, development of brood and foraging habits. The colony was discovered because the grass border of a road had been cut, revealing the series of small grass mounds in the high grass just behind. This strip (6 to 9 feet wide) of uncut grass, scattered shrubs and forbs formed a field- wood border. In front of it, across the road, an open field stretched to the northeast and allowed continuous sun from shortly after sunrise until mid afternoon, when the nest was shaded by trees. Behind lay a small poplar (Populus tremuloides Michx.) wood, which extended back to a swamp 18 yards away. Plants in the nest strip were mostly grasses ( Poa pratensis L., Panicum oligosanthes Scribnerianum (Nash) Fern., Aristida purpurascens Poir.), together with shrubby meadowsweet ^Facilities of The University of Michigan’s Edwin S. George Reserve were made available by the Reserve’s administrators, Dr. F. C. Evans and Dr. N. G. Hairston. Manuscript received by the editor October 21, 1971. 169 170 Psyche [September ( Spirea alba Du Roi) and some not very flourishing berries: red raspberry ( Rubus idaeus L.), creeping blackberry ( Rubus fiagellaris Willd.) and common blackberry ( Rubus sp.). Just back of the nest more vigorous Spirea alba made an almost complete wood-field fringe and extended back into the wood for 6 to 14 feet, forming a shrub layer until the more moist ground near the swamp was taken over by a thick growth of sensitive fern ( Onoclea sensibilis L.). The visible parts of the nest consisted of a series of inconspicuous low grass-thatch mounds strung along parallel to the woods edge behind and to the road in front. In 1969 there had been seven major mounds along 30 feet, but in 1970 those of the northeast half of the line had been abandoned and the ants occupied 10 mounds along the south-west 15 feet. A main cluster, lying near the center of the line, was composed of five, which rose into peaks of three to six inches and intercommunicated by low thatch structures barely above ground level. This made a conglomerate which stretched out into a very irregular shape, which was four feet at its longest extension. The five other mounds had no above-ground thatch connections. Three of these extended to the north-east of the main group and were 16, 33 and 38 inches from it. A fourth was 9 feet to the south-west and the last was five feet back from the main group and lay at the base of a poplar tree at woods edge. The mounds varied in diameter from 8 X 10 inches to 18 X 21 inches and in height from two to six inches. Structure of the mounds. The mounds were built of grass thatch with bits of poplar and spirea leaves intermingled. Live grasses growing through them gave some reinforcement but major support was furnished by spirea and berry stems. Superficially they looked much like enlarged igloo mounds of Dolichoderus mariae but inside there was no large central chamber. Instead, the grasses formed floor and ceiling for two or three stories of chambers and galleries. These provided a surprisingly dry place for pupae to mature. Concealed runways at the bottom of the mounds extended out just under the compressed leaf layer and were cut into the ground for about one-fourth of an inch. They ran to other mounds and to foraging areas. In addition there were galleries and chambers cut beneath the ground surface down among the plant roots for three or more inches. Thus part of the nest was underground and the mounds served primarily for hastening the maturing of brood. The mound set back at the base of the poplar tree at woods edge had a special significance. It was large, 20 X 21 inches across, and was built up partly of soil. On September 24, 1971, considerable 1971] Talbot — Formica dakotensis 171 dug-out soil was noticed just in front of it and workers were begin- ning to carry other workers from various mounds to this one. When leaves were cleared away a labyrinth of pathways was found cut into the soil and going into the mound’s base. Each day hereafter, until observations ceased on October 1, ants were seen carrying other ants into the lower part of the mound. Evidently this was the hibernating place for the whole colony. On June 3, 1971 workers were carrying other workers out from this mound. Trails and foraging grounds. Among the grasses of the nest area there was a superficial trashy layer of loose tree leaves, stems, etc., and under this the usual matted layer of compressed leaves decaying into the soil. It was this soil-leaf stratum which the ants used for tunneling to their foraging grounds. Their food seemed to be aphid honey dew but very little attending of above-ground aphids was seen. In 1970 the only workers seen foraging above-ground went by tunnel for about 10 feet and then up into a small poplar tree. Here 5 to 15 ants at a time attended aphids on leaf petioles. Most of the hidden aphids were on bases of spirea stems. Only the young, green stems were used and all such stems investigated had the lowest inch or two covered with aphids. If the lower stem had leaves piled aboout it, no structure was built but if the aphids extended above the leaf layer, a very tight thatch shelter encircled them. These thatch collars were rather easy to see and were very abundant but no ants were seen upon them because runway tunnels took the ants directly into their bases. Perhaps the ants also attended root aphids. There were some tunnels extending into the soil be- neath the plants but no aphids were found on roots. In the grassy area in front of the nest ants were seen going into several holes at sides of clumps of grasses. One such clump was dug and aphids were found at the base of leaves, where stem and roots joined. The other two colonies were more typically field ants, being further away from shrubs and trees and more directly dependent on field plants to harbor their aphids. The larger colony, near Southwest swamp, had 8 major mounds spread over 19 feet. It lay in a field of Poa compressa grass, goldenrod ( Solidago nemoralis Ait.), iron- weed ( Verononia altissima Nutt), Queen Anne’s Lace ( Daucus carota L.) and small scattered dogwood (Cornus stolonifera Michx.). Runways radiated from the mounds to many of these plants which had small thatch shelters surrounding their bases. All of the aphids found were just below ground level where roots and stems met. The smaller colony, on a grassy slope above a cattail marsh, had only three main mounds spread over 27 feet. It had a number of 172 Psyche [September _c bJD .5? C CC CCCCCCCC *6 E £ E *£ E E 'E OOOOOOO’-h oooooooo vOOOmCM-*-000 Mi Fig. 2. The percentage of the total number of stinkbugs counted on a strip of blackberry bushes which were totally exposed to the sun. The counts took place at various times during the day on 5 October 1971. The arrow points to the time of appearance of the sun following a period of early morning fog. blackberry patch. This species took second and third instar nymphs of E. conspersus as well as the nymphs of other Heteroptera. A larger unidentified black wasp was seen once with an adult bug and on another occasion as it searched the stems and undersides of leaves of blackberry plants. I watched a yellowjacket, probably Vespulci penn- sylvanica , macerate a late stage nymph. The most conspicuously successful predator of the bug was the garden spider, Araneus diademata. Captured stinkbugs often ap- peared in the orb webs of this species in the late summer. Less com- monly spiders were seen feeding on a wrapped stinkbug. A number of other potential predators of stinkbugs, several in- sectivorous birds, were seen hunting in the lot. Some bird species are known to take pentatomids (Southwood and Leston, 1959; Orians 1971] Alcock — Behavior of Stinkbug 219 and Horn, 1969). I offered sixteen juvenile redwinged blackbirds (Agelaius phoeniceus) two live E. conspersus adults. Nine of the birds ate both bugs, although usually not before giving behavioral signs that they found the insects distasteful (Alcock, ms. submitted). Stinkbugs, as is well-known, possess thoracic glands which are capable of secreting a volatile substance with an odor somewhat to extremely offensive depending on the species of bug and human tester. E. conspersus readily discharged its glands when handled roughly, or pecked by a bird, and several times I detected the odor coming from bugs trapped in a spider’s web. This action may protect some bugs from some predators, particularly birds, which are not strongly mo- tivated to feed. However, the insect is not solely dependent upon its secretion as it engages in a variety of defensive maneuvers — re- maining immobile much of the time, scuttling under leaves or drop- ping from them to the ground when approached or touched, kicking at ants, buzz-flying and wing-whirring. The defensive function of the latter two behaviors is purely speculative. However, bugs flying some distance definitely produced a fairly loud and, to this observer at least, a very bee-like buzz which might deter some aerial predators from attacking. Wing-whirring consisted of lifting and rapidly vibrating the wings, producing a loud buzz. Five of a group of about fifty bugs which I picked up, handled, and returned to a leaf remained on the leaf and wing-whirred. Similar behavior has been reported for some species of stinkbugs occurring on cocoa plants (Callan, 1944) and might serve to startle or to warn a predator not to attack. Reproductive Behavior Only one generation of stinkbugs mated in Seattle although in California some populations have two reproductively active genera- tions in a single summer (Borden, Madsen and Retan, 1952). The adults which overwintered in leaf litter beneath the blackberries had emerged in large numbers by the time observations were begun (mid- May). Many adults and mated pairs were seen in the lot through late June but by the end of the first week in July very few adults could be found. By late July the second generation bugs were appear- ing in abundance with many feeding on the blackberries which were just beginning to ripen at that time. Mated pairs were rarely seen in the morning, never at midday, and often from 15.00 to dusk (21.00). I believe that the initiation of courtship and copulation occurred almost exclusively in the late afternoon and early evening. The mated pairs seen in the morning probably had coupled the previous evening. Hunter and Leigh 220 Psyche [December (1965) report that in the lab bugs remain in copulo from 3-35 hours. A total of six successful courtships was observed, largely in late May, with copulation occurring at 15.35, 18.39, 18.46, 19.27, 19.50, and 20.05. The description of courtship behavior which follows is based primarily on observations of one aggregation of between 10-20 bugs regularly present on one bracken fern in late May and early June. Courting males were active despite the cool evenings; the females present were pressed flat against the stem or leaves of the fern. Males ( 1 ) palpated the upper surfaces of females with their antennae, (2) appeared to attempt to raise the female’s abdomen by placing their heads under the side of the female and lifting up- wards, and (3) turned and pressed their aedaegus, which might be completely extruded and partially inverted, against the side, belly, or tip of the abdomen of the female (or occasionally against the stem of the fern or even against another male courting the same fe- male). These behavior patterns occurred generally in the order pre- sented but since females were often unresponsive might be repeated over and over again. Sometimes in response to a courting male a female would lift her abdomen slowly upward away from the surface on which it had been resting. The male would continue to court, particularly with activ- ities (2) and (3), and often (4) palpated the undersides of the female’s abdomen while standing directly behind her. This might induce the would-be mate to raise her abdomen still higher until her body formed an approximately 30° angle with the stem to which she clung. At this point the male turned away and with completely extruded and inverted aedaegus backed toward the female all the while facing directly away from her. The male’s abdomen was also raised and upon touching the female he pressed his aedaegus about the tip of the female’s abdomen until it entered the female genital opening. After a series of small movements copulation was firmly achieved. The bugs might then move a short distance before settling down to remain in copulo for many hours. Males were extremely persistent courters often attending to a single female for 15-30 minutes before copulation was accomplished or before the male left in search of a more receptive female. One male was watched for almost one hour as it courted without success. However, on return to the plant 45 minutes later a mated pair rested on the branch where the persistent male had been active. Unreceptive females often simply remained flattened against a stem and did not move. Rejection of a male could be more active how- 1971] Alcock — Behavior of Stinkhug 221 Fig 3. A stinkbug photographed in the morning with its back presented directly to the sun with the bug assuming the “tilt” position. 222 Psyche [December ever. Females occasionally moved their abdomens slowly from side to side or rocked back and forth as males pressed at them. One fe- male repeatedly placed a hind leg on a male and pushed it away over a period of 45 minutes. Two females lifted their abdomens upwards but refused to permit the male aedaegus to enter their genital open- ing. The Aggregation and Dispersal of E. conspersus This stinkbug species often forms groups. As previously noted the courtship fern usually carried several bugs (during the period from 27 May to 1 June) although the exact number fluctuated from o to 21. Although the bugs were not marked it is at least possible that the same individuals came and went returning to the fern several times. Six meters away another bracken fern was found with an- other group of bugs. This aggregation was a durable one lasting from late May to 1 July and probably several days more. Again the number of stinkbugs present varied a good deal reaching a maximum of 30 individuals on 21 June including 8 mated pairs at 20.30. Given the considerable amount of sexual activity which occurred in these first generation groups I initially felt that their function was exclusively sexual in nature. However, I later discovered that aggregations were also formed by nymphs and second-generation non- breeding adults. Groups of 5-6 late instar nymphs could be found basking on the same leaf together, some even with their heads under the side of a companion, an action reminiscent of the abdomen lifting behavior of courting males. As fall neared large numbers of bugs (up to 61) occurred on a single fern. Smaller contact groups of 5-10 individuals were common (Fig. 4) and one large tightly clumped cluster of between 15 and 35 bugs was found on exactly the same portion of a blackberry stem on various dates from 28 September to 17 October. At the same time not all individuals showed a tendency to clump together in semi-permanent groups. On the contrary many E. con- spersus were highly active and could be seen walking substantial dis- tances along blackberry stems from one plant to another. On warm days in the afternoon bugs were often seen flying five to ten meters. Of 30 bugs marked on 21 May, I could find only 2 the next day. The apparent mobility and dispersal of many stinkbugs contrast sharply with the seeming stability of some groups. Moreover, the bugs, despite sometimes forming contact groups, also could demonstrate a degree of anti-social behavior. Stinkbugs were seen kicking at each other when touched by a companion. Once one 1971] Alcock — Behavior of Stinkbug 223 A group of five stinkbugs in a typical fall cluster on a blackberry stem. 224 Psyche [December bug was observed repeatedly butting another with its head with what appeared to be aggressive intent. Discussion Basking Behavior Although there have been few reports of basking by pentatomids (however see Thomas, 1954) it nevertheless may be practiced quite commonly by them. Chlorochoa sayi , a less abundant stinkbug than E. conspersus in the study plot, also basked on blackberry leaves. This behavior is widespread among insects in general (particularly in deserts and northern latitudes where there are great temperature fluctuations). Some species, for example the desert locust (Waloff, 1963), may also orient the body to achieve maximum exposure to the sun. Basking is appropriate for the Northwest where summer nights are usually quite chilly and summer days often partly cloudy and cool. Feeding Behavior Although many pentatomids feed on a wide range of plant species, E. conspersus is the first stinkbug known to exploit a fern. Very few insects attack ferns (Brues, 1920; Soo Hoo and Fraenkel, 1964). The only other Heteroptera associated with ferns are some members of the mirid subfamily Bryocorinae (Southwood and Leston, 1959 ; Woodward, Evans and Easton, 1970). The stinkbug attacked the stems of adult bracken ferns avidly despite the fact that the plant when full grown is avoided by almost all herbivores and is reputed to be toxic to cattle (Muenscher, 1939). Moreover, bracken pinnae of at least some populations contain ana- logs of ecdysone (Kaplanis et al., 1967) although whether stems con- tain the substance is not proven. The bug’s ability to feed on bracken is all the more remarkable because, unlike other fern herbivores, E. conspersus is not a specialist limited to ferns. Reproductive Behavior The courtship behavior of very few stinkbugs has been reported in any detail (but see Kullenberg, 1947; Teyrovsky, 1949; Southwood and Hine, 1950; Leston, 1955; Kaufmann, 1966). Judging from these cases courtship among stinkbugs must be highly diverse. For ex- ample, the male of Calidea dregii first faces the female and then climbs forward onto her head (Kaufmann, 1966) ; the males of Dolycoris haccarurn first creep under the abdomen of the female (Teyrovsky, 1949). However, in every previously studied case the male eventually climbs onto the back of the female and faces in the same direction 1971] Alcock — Behavior of Stinkbug 225 as its partner. It is in this position that insertion of the aedaegus occurs. Usually the male then turns, dismounts and faces away from the female. Copulation proceeds in this end-to-end pose. This is a very common sequence of events among the Heteroptera in gen- eral (Grasse, 1951; Weber, 1930). E. conspersus is then the only stinkbug known to omit the male-above position and to effect a copulatory union directly end-to-end. How might such behavior have evolved? Weber noted (i93°> p. 307) that there were only two basic patterns of Hemipteran copu- lation — ( 1 ) the male and female facing in the same direction with the male by the side or above the female and (2) the male and female united end-to-end facing away from each other. At the time no species was known which copulated in position (2) without first assuming the side-by-side or male-above position. From this evidence Weber argued that the end-to-end position was an elaboration of the pre- sumably more primitive position ( 1 ) . Therefore it seems reasonable that the unusual courtship pattern of E. conspersus which omits the male-above component altogether is probably a relatively recent evolu- tionary invention derived from the second basic pattern. Representa- tives of all three copulatory patterns occur in the Orthoptera (and doubtless other insect orders); Alexander’s (1964) explanation for the evolution of direct end-to-end initiation of copulation in the Orthoptera is essentially the same as that outlined above for the Hemiptera. Omission of the back-climbing phase of courtship makes some sense for pentatomids given their bulky flattened shape which could make coupling awkward in a male above/female below position (N. T. Davis, pers. comm.). Indeed in type (2) copulations the function of the male’s dismounting and turning away from the female may be to achieve a more stable and easily maintained copulatory position. Initiation of copulation directly in the end-to-end position might have had its origin in premature turning behavior. After a number of unsuccessful attempts to copulate while on the back of a female, some males might tend to dismount and turn despite the fact that coupling had not yet taken place. Teyrovsky (1949) observed a male D. baccarum attempt a direct end-to-end union with another male it had been unsuccessfully courting. Presumably in the evolution of the sexual behavior of E. conspersus selection first favored males with a low threshold for dismounting and turning. Gradually selection has eliminated mounting the female entirely while favoring those males which act to induce the female to assume that position which makes direct end-to-end mating most easy. 226 Psyche [December The Formation of Aggregations Groups of stinkbugs, both adults and nymphs, are quite well-known (Thomas, 1954; Southwood and Le:ton, 1959; Canerday, 1965) and appear to have multiple functions. First, mating aggregations permit males to locate or attract females of their own species (Tey- rovsky, 1949; Southwood and Leston, 1959). The means by which reproductively active E. conspersus manage to form groups is un- known. The fact that non-reproductive individuals also clump to- gether means that specific sex pheromones are not a necessary basis for group formation. Second, clumping could be an anti-predator adaptation. A group of toxic, warningly-colored insects may more effectively advertize po- tential unpalatability than scattered individuals could. In addition should one member of a group be taken, the predator would be likely to avoid the others. However, the formation of groups by nymphs and adults of a cryptically colored species such as E. conspersus can hardly serve this function. As previously noted, the bug proved to be edible, although apparently distasteful, to redwinged blackbirds. The formation of groups must make these bugs more vulnerable, not less, to hungry redwings as well as to their wasp predators which can probably learn to return repeatedly to a productive searching area (Evans, 1966). The fact that the bugs are not terribly nimble also places a premium on avoiding detection by their enemies. Thus there must be some other advantage for group formation by second genera- tion adults and nymphs which outweighs increased risk of attack by predators. What this advantage is remains uncertain. However, given the death of the ferns it seems probable that the bugs were injecting toxic substances into their foodplants as Hemiptera are known to do (Nuorteva, 1958; Adams and McAllen, 1968). The salivary glands of some mirids contain pectinases (Laurema and Nu- orteva, 1961) as does as least one lygaeid (Adams and McAllen, 1958). The pentatomid D. baccarum possesses salivary proteases and amylases (Nuorteva, 1954). Thus it is possible that by feeding in groups individual bugs may extract plant juices more easily than otherwise. In any event, the relative advantages of group formation as opposed to dispersal must be rather closely balanced given the great variability in the tendency to aggregate exhibited by E. conspersus. Summary This paper describes the behavior of members of one population of Euschistus conspersus Uhler, a small brown stinkbug. These bugs characteristically bask for some time after sunrise, often on black- 1971] Alcock — Behavior of Stinkbug 227 berry leaves. Later in the day they feed on a wide variety of plants including bracken fern, the first pentatomid reported to attack a fern. Adults which have overwintered mate in the late afternoon and evening in May and June. Males engage in several tactile court- ship activities which may induce the female to raise her abdomen upward. Males then turn away from females and back toward them with aedaegus extruded. Copulation is initiated in this end-to-end position. In all other pentatomids and most Heteroptera copulation begins with the male above the female, although later the male often dismounts and faces away from its mate. Aggregations of adults and nymphs are quite common and may have two functions — ( 1 ) to bring reproductively active individuals together and (2) to facilitate extraction of plant juices. References Allen, J. B. and J. W. McAllen 1958. Pectinase in certain insects. Can. J. Zool. 36: 305-308. Alexander, R. D. 1964. The evolution of mating behaviour in arthropods. Roy. Entomol. Soc. Lond. Symp. 2: 78-94. Borden, A. D., H. F. Madsen and A. H. Retan 1952. A stinkbug, Euschistus conspersus , destructive to deciduous fruits in California. J. Econ. Entomol. 45: 254-257. Brues, C. T. 1920. The selection of food plants by insects with special reference to lepidopterous larvae. Amer. Nat. 54: 313-332. Callan, E. McC. 1944. Cocao stinkbugs (Hem., Pentatomidae) in Trinidad, B.W.I. Rev. Entomol. 15: 321-324. Canerday, T. 1965. On the biology of the harlequin bug, Murgantia histrionica (Hem- iptera: Pentatomidae). Ann. Entomol. Soc. Amer. 58: 931-932. Essig, E. O. 1929. Insects of western North America. Macmillan, New York. Evans, H. E. 1966. The comparative ethology and evolut on of the sand wasps. Har- vard University Press, Cambridge, Mass. Grasse, P.-P. 1951. Tra te de zoologie , anatomie, systematique, biologie. Tome X. Insectes superieurs et hemipteroides. Ordre des Heteropteres. Mas- son et Cie, Paris. Hunter, R. E. and T. F. Leigh 1965. A laboratory life history of the consperse stinkbug Euschistus conspersus (Hemiptera: Pentatomidae). Ann. Entomol. Soc. Amer. 58: 648-649. Kaplanis, J. N., M. J. Thompson, W. E. Robbins, and B. M. Bryce 1967. Insect hormones: Alpha ecdysone and 20-hydroxyecdysone in bracken fern. Science 167: 1436-1437. 228 Psyche [December Kaufmann, T. 1966. Notes on the life history and morphology of Calidea dreg'ri (He- miptera : Pentatomidae : Scutellerini ) in Ghana, West Africa. Ann. Entomol. Soc. Amer. 59: 654-659. Kullenberg, B. 1947. Uber Morphologie und Funktion des Kopulationsapparats der Capsiden und Nabiden. Zool. Bidrag Fran Uppsala 24: 217-418. Laurema, S. and P. Nuorteva 1961. On the occurrence of pectin polygalacturonase in the salivary glands of Heteroptera and Homoptera Auchenorrhyna. Ann. En- tomol. Fenn. 27: 89-93. Leston, D. 1955. The function of the conjunctiva in copulation of a shieldbug, Peizodorus lituratus (Fabricius) (Hemiptera, Pentatomidae). J. Soc. Brit. Entomol. 5: 101-105. Muenscher, W. C. 1939. Poisonous plants of the United States . Macmillan, New York. Nuorteva, P. 1954. Studies on the salivary enzymes of some bugs injuring wheat kernels. Ann. Entomol. Fenn. 20: 102-124. Nuorteva, P. 1958. Die Rolle der Speichelsekrete in Wechselverhaltnis zwischen Tier und Nahrungspflanze bei Homopteren und Heteropteren. Ent. Exper. Appl. 1 : 41-49. Orians, G. H. and H. S. Horn 1969. Overlap in foods and foraging among four species of blackbirds in the Potholes of central Washington. Ecology 50: 930-938. Soo Hoo, C. F. and G. Fraenkel 1964. The resistance of ferns to the feeding of Prodenia eridania larvae. Ann. Entomol. Soc. Amer. 57: 788-790. SOUTHWOOD, T. R. E. AND D. J. HlNE 1950. Further notes on the biology of Sehirus bicolor (L.). Entomol. Mon. Mag. 86: 299-301. Southwood, T. R. E. AND D. Leston 1959. Land and water bugs of the British Isles. Frederick Warne and Co., London. Teyrovsky, V. 1949. Praeconnubia and courtship in terrestrial bugs. Acta Acad. Sci. Nat. Hist. Moravo-Silesiacae, Brno 21: 1-16. Thomas, D. C. 1954. Notes on the biology of Hemiptera Heteroptera. Entomologist 87: 25-30. Waloff, Z. 1963. Field studies on solitary and transiens desert locusts in the Red Sea area. Anti-Locust Bull. no. 40. Weber, H. 1930. Biologie der Hemipteren. Julius Springer, Berlin. Woodward, T. E., J. W. Evans and V. F. Eastop 1970. Hemiptera. In The insects of Australia. Melbourne University Press, Melbourne. THE ORB-WEAVER GENERA SINGA AND HYPSOSINGA IN AMERICA (ARANEAE: ARANEIDAE)* By Herbert W. Levi Museum of Comparative Zoology, Harvard University The North American spiders commonly placed in Singa belong to four different genera. It was first thought best to publish on these species together with the small species of Araneus. However, the slowness with which the Araneus studies proceed makes it advisable to publish on Singa and Hypsosinga first. Most of the specimens of these genera belonging to the American Museum appear to be lost and only small collections other than those of the Museum of Comparative Zoology were available. I would like to thank the following for loan of additional specimens: Dr. J. A. Beatty; Mr. D. E. Bixler; Mr. D. Buckle; Dr. J. A. L. Cooke for collections of the American Museum of Natural History and Cornell University; Dr. R. Crabill for specimens from the U.S. National Museum; Dr. B. Cutler; Dr. C. D. Dondale of the Canada Dept, of Agriculture; Dr. B. J. Kaston; Dr. W. W. Moss for collections from the Academy of Natural Sciences, Phila- delphia; Dr. W. Peck for specimens from the Exline collection; Dr. J. Proszynski for collections of the Polish Academy of Sciences; Miss Susan Riechert; Mr. V. Roth; Dr. W. Shear; Dr. C. Triple- horn of Ohio State University; Prof. S. L. Tuxen for specimens from the Zoological Museum of the University of Copenhagen; Dr. J. D. Unzicker for specimens from the Illinois Natural History Survey; Prof. M. Vachon and Dr. M. Hubert for collections of the Museum National d’Histoire Naturelle, Paris; Dr. B. Vogel; and Dr. H. K. Wallace. I am also indebted to Mr. J. Denis, Mr. G. Puhringer, and especially to Miss Susan Riechert for helpful information. The research and publication were supported by Public Health Service Grant AI-01944 from the National Institute of Allergy and Infectious Diseases. Singa C. L. Koch Singa C. L. Koch, 1836, Arachniden, vol. 3, p. 42. Type species Singa hamata (Clerck) designated by Thorell, 1869. On European Spiders, p. 58. The name is of feminine gender. Diagnosis . The anterior median eyes are the largest, the posterior medians the same size or smaller, the laterals about 0.6 diameter of the anterior medians. In Singa the median ocular quadrangle is wider *Manuscript received by the editor February 8, 1972. 229 230 Psyche [December in front (Figs. 20, 30) ; in Hypsosinga it is rectangular or wider behind. The carapace of the female is shiny and has no thoracic depression (Figs. 22, 32) ; in the male it has a short longitudinal line. The first coxa of the Singa male, unlike that of the Hypso- singa male, has a hook on the side. The height of the clypeus is equal to or less than the diameter of the anterior median eyes (Figs. 20, 30) ; in Hypsosinga it is higher. The second tibia of the Singa male may be modified; in Hypsosinga the first tibia may be modified. In Singa the sides of the abdomen are almost parallel and the abdomen overhangs the spinnerets. There are two dorsal longi- tudinal black bands on the abdomen of Singa (Figs. 21, 22, 31, 32). The female genitalia differ from those of Hypsosinga in that Singa has a scape on the epigynum (Figs. 6, 11, 25). The palpus of the Singa male has an enormous terminal apophysis (A in Figs. 3, 5» 9), ending in a sclerotized spine and located in the contracted palpus on the outer side of a rather narrow tegulum (Fig. 5). Below the terminal apophysis, a heavily sclerotized hook, perhaps the subterminal apophysis (SA in Figs. 1, 3, 10), lies hidden in the contracted bulb. The embolus may have a lamella (Figs. 1, 9) ; it does not seem to have a part that breaks off in mating. The stipes ( I in Fig. 1 ) is a distinct sclerite in S. hamata , but is apparently fused to the embolus in S. keyseriingi (Fig. 9). The tegulum is widest near the base of the conductor (Figs. 2, 3, 10). Natural History. Surprisingly little is known about the habits of Singa. They make a complete orb. Nielsen (1932, Biology of Spiders, 2 ; fig. 330) has a picture of the retreat of S. hamata. Both American species prefer moist locations, and adults are found through- out the season. G. Piihringer (personal communication) told me that Singa phragmiteti Nemenz is common on reeds along the Neu- siedler See in Austria. It prefers a site above water and has to be collected from a boat. I suspect that American species have similar habits, which may account for the few specimens in collections. Distribution. Species are known only from Eurasia and temperate North America. All others described are probably misplaced. Misplaced American Species. (This list of names follows Roewer, 1942, Katalog der Araneae. The types of the species, unless indicated otherwise, have been examined. Species placed in Hypsosinga are not listed.) abbreviata Keyserling, 1879 — Theridiosomatidae. bengryi Archer, 1958 = Metepeira bengryi (Archer), new com- bination. 1971] Levi — Orb-W caver Genera 231 calix Walckenaer, 1841 — Alpaida calix, new combination. crewii Banks, 1903; the type is lost, the description is not recog- nizable. dotana Banks, 1914 — Theridion dotanum. duodecimguttata Keyserling, 1879 — Alpaida duodecimguttata (Keyserling) , new combination. erythrothorax Taczanowski, 1873 = Alpaida erythrothorax (Tac- zanowski), new combination. essequibensis Mello-Leitao, 1948 — ? Alpaida essequibensis, type specimens unavailable. flava O. P.-Cambridge, 1894 = Araneus flavns (O. P. -Cam- bridge) . floridana Banks, 1896 — Araneus floridana (Banks). leuco gramma White, 1841 — Alpaida leuco gramma (White), new combination. listerii McCook, 1893 = Araneus pratensis Emerton. longicauda Taczanowski, 1878 = generic placement uncertain. marmota Taczanowski, 1873 — Alpaida m arm or at a (Taczanow- ski), new combination. maura Hentz, 1847 = Alpaida calix Walckenaer. moesta Banks, 1893. The type has been destroyed, the descrip- tion cannot be recognized. mollybyrnae McCook, 1893 — Metazygia pallidula (Keyserling). new SYNONYMY. Type locality in error; not District of Colum- bia, probably Colombia. niveosigillaa Mello-Leitao, 1941 = ? Alpaida niveosigellata, type specimens unavailable. pratensis Emerton, 1884 = Araneus pratensis (Emerton). praticola Simon, 1895 — Araneus pratensis (Emerton). tremens Holmberg, 1876. The type has been destroyed. vanbruysseli Becker, 1879 = Cyclosa turbinata (Walckenaer), NEW SYNONYMY. vittata Taczanowski, 1873. The type is lost. The two common species north of Mexico, calix and pratensis , are obviously misplaced. Judging by the structure of the genitalia, Singa pratensis is close to Araneus sturmi (Hahn) of Europe. Dif- ferences are the shape of the abdomen and lack of body setae, but the shape, setation, and coloration of the abdomen are quite variable in the many species of these small Araneus. Araneus sturmi is the type species of the genus A tea. At present it does not seem wise or even feasible to fragment the genus Araneus. It would lead to 2 32 Psyche [December proliferation of names without meaning for relationships. They are obviously monophyletic. Singa calix belongs to a South American genus, one of the largest genera of orb weavers in the Americas. As far as I know at present, the oldest name is Alpaida O. P. -Cambridge, 1889. But numerous other generic names have been used for this genus; Lariniacantha Archer (1951, Amer. Mus. Novitates, no. 1487, p. 15) most re- cently. The genus is much closer to Acanthepeira than to Singa. However, I am still hestitant about the placement until I have more knowledge of the webs and habits of the species in the genus. Key to American Species of Singa 1 a. Base of epigynum trapezoidal, with sides sclerotized (Figs. 11- 14) ; median apophysis of palpus with one hook (Figs. 23, 24) keyserlingi ib. Base of epigynum with a lobe on each side (Fig. 25) ; median apophysis of palpus with two hooks (Figs. 33, 34) eugeni Singa hamata (Clerck) Figures 1-8 Araneus hamatus Clerck, 1757, Aranei Svedici, p. 51, pi. 3, fig. 4. Female type specimens from Sweden believed lost. Bonnet, 1955, Bibliographia Araneorum, vol. 2, p. 513. Singa hamata, — C. L. Koch, 1836, Die Arachniden, vol. 3, p. 42, figs. 197, 198, 9, $. Wiehle, 1931, in Tierwelt Deutschlands, vol. 23, p. 42, figs. 54-57, 9, $ . Roewer, 1942, Katalog der Araneae, vol. 1, p. 873. Locket and Millidge, 1953, British Spiders, vol. 2, p. 157, figs. 102b, 103c, 105c, 9, $. This species, very similar to the two American ones, is known only from Eurasia. Singa keyserlingi McCook Figures 9-24, Map 1 Singa keyserlingi McCook, 1893, American Spiders, vol. 3, p. 230, pi. 19, fig. 2, 9. Female holotype from St. Louis, Missouri, in the Academy of Natural Sciences, Philadelphia; examined and labelled as type. Singa campestris Emerton, 1915, Trans. Connecticut Acad. Sci., vol. 20, p. 153, pi. 3, fig. 3, $. Male syntype from Rat Portage, Ontario, in the Museum of Comparative Zoology; examined, new synonymy. Note. McCook described S. keyserlingi as a new species. How- ever, in the first paragraph of the description he finds it necessary “to propose a new name” for the species Keyserling illustrated and called erroneously Singa rubella (Hentz). The specimens McCook 1971] Levi — Orb-W eaver Genera 233 Figs. 1-8. Singa hamata (Clerck). 1-5. Left palpus. 1-3. Expanded. 4. Mesal view. 5. Ventral view. 6-8. Epigynum. 6. Ventral. 7. Posterior. 8. Dorsal. Figs. 9-10. Singa keyserlingi McCook, palpus, expanded. Abbreviations: A, terminal apophysis; C, conductor; DH, distal hema- todocha; E, embolus; H, basal hematodocha ; I, stipes; M, median apophysis; P, paracymbium ; R, radix; S, subtegulum ; SA, subterminal apophysis; T, tegulum; Y, cymbium. Size Indicators : 0.1 mm. 234 Psyche [December had on hand and figured were specimens of what is here considered to be S. keyserlingi , and not what Keyserling called rubella. In the United States National Museum there is a specimen marked Singa rubella Hentz, St. Louis, Mo., collected by Marx. Another label in the vial reads “Cotype 1688 U.S.N.M.” The specimen contained is a female of what is here called S. eugeni , and there is no evidence that it is a syntype of S. keyserlingi McCook. McCook reports having seen specimens from the District of Columbia in the Marx collection. Keyserling also reports that the Singa rubella he examined came from the District of Columbia from the Marx collection. Figure 13 was prepared from the type of 5. campestris. Description. Female from Wisconsin. Carapace orange; head region, clypeus black. Chelicerae, labium, endites dark brown. Ster- num orange. Legs orange, distal articles darker. Dorsum of abdo- men with two wide black bands separated by a narrow white band (Fig. 22). Sides of black bands have lateral white band (Fig. 21). Venter has a black patch which may have a white line on each side. Anterior median eyes slightly more than one diameter apart, one diameter from laterals. Posterior median eyes their radius apart, two and one-half diameters from laterals. Total length 5.5 mm. Carapace 2.4 mm long, 1.7 mm wide. First femur, 1.4 mm; patella and tibia, 2.1 mm; metatarsus, 1.3 mm; tarsus, 0.7 mm. Second patella and tibia, 1.9 mm; third, 1.3 mm; fourth, 1.7 mm. Male from Wisconsin. The coloration is like that of the female except that black on the head seems limited to the eye region. The chelicerae are black only distally, orange at the base. The dorsum of the abdomen is all black. The anterior median eyes one and one- third diameters apart, one diameter from laterals. Posterior median eyes their radius apart, two and one-half diameters from laterals. The palpal patella has two weak macrosetae. The first and second legs have strong macrosetae on prolateral surface, but are not bent or swollen. Total length 3.9 mm. Carapace 2.2 mm long, 1.6 mm wide. First femur, 1.5 mm; patella and tibia, 2.0 mm; metatarsus, 1.4 mm; tarsus, 0.7 mm. Second patella and tibia, 1.8 mm; third, 1.2 mm; fourth, 1.5 mm. Variation. In the female the scape of the epigynum varies greatly in length and shape (Figs. 1 1 - 1 5 ) ; the internal ducts may be either heavily sclerotized or transparent, sac-like and difficult to make out. Females vary from 5. 1-6.0 mm in total length, carapace 1 .5-1.7 mm 1971] Levi — Orb-W eaver Genera 235 Figs. 11-24. Singa keyserlingi McCook. 11-22. Female. 11-19. Epigynum. 11-14. Ventral. 11. (Illinois). 12. (Wisconsin). 13. (Ontario). 14, 15. (South Dakota). 15. Posterior. 16-19. Cleared. 16. Subventral. 17. Ventral. 18. Dorsolateral. 19. Posterior. 20. Face. 21. Abdomen, lateral. 22. Dorsal. 23-24. Left palpus. 23. Mesal. 24. Ventral. Size Indicators : 0.1 mm, except for Figs. 21-22, 1 mm. 236 Psyche [December wide. Males vary from 2. 3-4.0 mm in total length, carapace 1 .3-1.6 mm wide. Diagnosis. Females differ from those of S. eugeni in the shape of the base of the epigynum, trapezoidal with the lateral margins sclerotized (Figs. 11-14). The male differs from that of S. eugeni by having only one hook on the median apophysis (Figs. 23, 24). Natural History. Singa keyserlingi has been collected from open woods, in low shrubs and by sweeping grass on lakeshores. Mature males have been collected in all months between May and August. Females have been collected through September. Distribution. From Edmonton, Alberta, Smoky Falls, Ontario to Black Warrior National Forest (Winston Co.), Alabama (Map 1). Singa eugeni sp. n. Figures 25-34, Map 1 Singa rubella, — Keyserling, 1893, Spinnen Amerikas, vol. 4, p. 284, pi. 14, fig. 209, $ . Not Epeira rubella Hentz. Type. Male holotype and female paratype from T8N, R5E, S9NWJ4, Iowa County, Wisconsin (Susan Riechert), in the Mu- seum of Comparative Zoology. The species is named after Count Eugen Keyserling. Description. Female. Carapace orange with a wide black band covering eye region (Fig. 32), narrowing behind. Clypeus black. Chelicerae brown-black. Labium black. Sternum yellowish with dark brown margin. Legs yellow. Dorsum of abdomen with two longi- tudinal dark bands; at each end the bands are darker and approach each other (Fig. 32). The bands are separated by a white pigment line. The sides are white ( Fig. 31); the venter is yellowish with an indistinct dark area in the middle. The anterior median eyes are one and one-quarter diameters apart, less than one diameter from laterals. Posterior median eyes are less than one-quarter diameter apart, one and one-half diameters from laterals. Total length 4.6 mm. Carapace 2.0 mm long, 1.4 mm wide. First femur, 1.3 mm; patella and tibia, 2.2 mm; metatarsus, 1.2 mm; tarsus, 0.6 mm. Second patella and tibia, 1.9 mm; third, 1.2 mm; fourth, 1.7 mm. Male. The coloration of the male is like that of female. The carapace is narrower in front than in the female. The anterior median eyes overhang the chelicerae. The anterior median eyes are more than one diameter apart, their radius from laterals. The pos- 1971] Levi — Orb-W eaver Genera 2 37 Map. 1. Distribution of S.nga keyserlingi (McCook) and S. eugeni sp. n. 238 Psyche [December terior median eyes are one-quarter diameter apart, one and one-half diameters from laterals. The second tibia is thick, slightly curved, with macrosetae on prolateral side, but only very slightly modified. The palpal patella has one strong and one very weak seta. There are eight black, sclerotized dorsal muscle attachments on the abdo- men. Except for the narrower carapace, the male looks like the female. Total length 5 mm. Carapace 2.2 mm long, 1.6 mm wide. First femur, 2.2 mm; patella and tibia, 3.4 mm; metatarsus, 2.3 mm; tarsus, 0.9 mm. Second patella and tibia, 2.9 mm; third, 1.4 mm; fourth, 1.9 mm. Variation. In some individuals only four black spots remain of the two abdominal bands, two anterior and two posterior. If present the soft projection from the terminal apophysis, seen in ventral view (Fig. 34), may be either a flap or a rod. Females are from 4.3- 6.5 mm in total length, carapace 1.1-1.6 mm wide; males are from 3. 6-5. 4 mm in total length, carapace 1.2- 1.8 mm wide. Diagnosis. Females are distinguished from those of S. keyserlingi by the smaller epigynum with a lobe on each side of scape (Fig. 25), as opposed to a sclerotized diagonal margin seen in S. keyserlingi. The median apophysis of the palpus has two hooks, one on each end (Figs. 33, 34). In the related S. nitidula of Europe the median apophysis is of different shape and the embolus narrower. Natural 'History. Wisconsin specimens came from open bottom- land forest, along backwaters of river edges, and edge of marsh ; Georgia specimens came from Spartina stems. Barrows (1918, Ohio J. Sci., 18: 310) reported that a Singa from Cedar Point, Ohio, almost certainly this species, made “a small orb in tops of dune grass (Andropogon) . During the day it stays in the hollow stems of dead grass.” The spiders have also been collected in Penn- sylvania by the wasp Episyron quinquenotatus (Say). The males are mature in September and October in the north. Adult females have been collected from May to October. Localities collected. Pennsylvania ; Erie Co. : Presque Isle State Park. Ohio. Erie Co.: Cedar Point. D.C. Washington. Georgia. McIntosh Co. : Sapelo Island. Michigan. Clinton Co. : Rose Lake. Eaton Co. : Calumet. Livington Co. : George Reserve. Midland Co. Wisconsin. Iowa Co.: $, g c ^ G g •- , c ^ 2 § 0 Cl, ‘-a | £• £ o £ g ° « T3 MH G qj 0 QJ O iu t: 7 G G 2 1 ’-5 Z ^ Mc'T' £.2 6, T s — T3 G ~- G g G G 0 G © u^tnininMOO'OM^ONoo OOOOt^Or^OC^CT'^ON HNto(\)tn + N+ ^ + ^ N 'O 'O M O On 00 CNJ O 00 u~i 2 £j3 G T) — Nil NO MO h h N N N N M 1T| Ov 'O (S N N co O00V0lflNl,>t'''0'O + 0\ ^orac^>n_cooscs'OLr,o> conOOnCNJ'j~>u~>1j-ico^nOno OOO'+OOOOOCO'OcOt^ i-Ht— ico^-nooOOnOCnJco ininvovONNoooo NO NO NO NO NO 'O NO 'O OnOnONONOnOnONOn >> . >N 03 D-i >0 . 03 Oh > ■S 2^ £ £ H, £ u e •-h £ G i“! s © 'o * Q > > * * * CM co On co co NO ON CO rj- 26q Psyche [December markers placed at ICO meter intervals throughout the field (see Fig. i). The lines on Fig. i deliniate the limit of eastward ex- pansion of 7. humilis colonies at the time the survey was taken. The new area added was then calculated by subtracting the total occupied area at the previous sampling date from the new total area occupied by 7. humilis. This area was then divided by the number of colonies in the new added territory to get the mean area per colony values. At the same time, quadrat sampling with 30 randomly placed 2 meter by 2 meter quadrats were carried out to determine the vegeta- tion characteristics. Weather data were taken from a station 3 miles northeast of the field site and averaged to get monthly mean tem- peratures and precipitation. Results The displacement of the three other ant species by 7. humilis started slowly in October, 1963, but increased to an almost constant rate from 3 May 1964 to 4 October 1968 (Table 1, Fig. 1). The new area added by 7. humilis during each displacement interval of six months was approximately 14000 m2 ranging from 8318 m2 to 19988 m2 in the study field. The mean area per colony of 7. humilis for the whole occupied portion of the field increased during each sampling interval, whereas the mean area per colony in the newly displaced land was almost a constant 1400 m2 per colony. The num- ber of colonies of 7. humilis increased during each displacement in- terval as the displacement proceeded whereas the number of colonies of P. calif ornicus decreased except for a minor fluctuation due to flooding between 3 October 1965 and 1 May 1966 (Fig. 2). By October 1968, not a single colony of P. grallipes was observed in the study field, and by 5 March 1969 all colonies of P. calif ornicus and V. pergandei were located outside the boundaries of the study field. In their place remained 57 colonies of 7. humilis. Discussion To explain this phenomenon of displacement, some comparison is necessary of the basic biology of the ant species involved. The nests of 7. humilis are situated wherever there is sufficient moisture and where light is excluded, as under rocks and logs (Wood word, 1905, 1910; Eckert and Mallis, 1937, Smith, 1947) or in shallow nests in the soil (Cook, 1953). These ants occur in a wide variety of habi- tats — swamps, beaches, lawns and gardens, roadsides, houses, and various woodlands (Crowell, 1968). 7. humilis are exceptionally rest- less ants and normally emigrate one or more times a season in search NUMBER OF COLONIES 1971] Erickson Displacement of Native Ant 261 Figure 2. The number of colonies in the entire field of P. calif ornicus (o) and 1. humilis (®) from October 1963 to October 1968. of more favorable habitats (Wilson, 1971). Colonies of I. humilis contain a large number of queens with thousands of workers ( Smith, 1947) and proliferate by swarming of detachments of workers who accompany secondary queens out of the nest (Wheeler, 1933 ; Wynne- Edwards, 1963; Crowell, 1968). They are highly omnivorous but tend to seek sweet or fatty foods (Eckert and Mallis, 1937,’ Creigh- ton, i960; Cook, 1953), and tend aphids and scale insects in orchards and gardens (Skaife, 1961). In contrast with I. humilis , the California harvesters are large ants (4-6 mm long) which are primarily seed gatherers, but are also known to be slightly omnivorous (Van Pelt, 1966). Colonies of P. calif ornicus are small in comparison to I. humilis and contain only one queen. Proliferation takes place by large swarms of winged reproductives. The California harvester ant tends to nest in dryer semi-desert habitats and can tolerate much higher temperatures than I. humilis (Wheeler, 1926; Cole, 1932, 1968; Michener, 1942; Erickson, 1972). The relative reproductive potential of I. humilis is probably much higher than P. calif ornicus. This is most likely due to the large num- 262 Psyche [December ber of queens in each colony, the method of colony proliferation, and the omnivorous habits of these ants. These factors may also account for the great and rapid spread of I. humilis throughout the temperate regions of the world. Basic differences in food resources limit to some extent the amount of competition between 7. humilis and the three harvester ant species (Table 2). 7. humilis is highly omnivorous whereas the three har- vester ant species are only slightly omnivorous, being basically seed gatherers. I. humilis not only monopolizes the proven food sources but attempts to control the remaining foraging areas (Wilson, 1971). In the main study field, food, especially seeds, were very abundant. The food chambers of P. calif ornicus and I. humilis were always full when the colonies were excavated. In fields 2 A and 2B, the area was supplemented with approximately five pounds of mixed grass seed per month to determine the foraging characteristics and distances for P. calif ornicus. The seeds, colored with common food coloring, were fully acceptable to the ants, making up 43 to 59% of the P. calif ornicus food stores and 9 to 17% of the I. humilis food stores. The variously colored seeds were spread in concentric circles from a nest of P. calif ornicus every 5 meters to a distance of 30 meters. The maximum foraging distance for P. calif ornicus was about 10 meters except in areas where there was an I. humilis colony in which case the harvester ants foraged no farther than 5 meters even though the I. humilis colony was 20 meters away. Even though both fields were supplemented with a little over 50 pounds of mixed grass seed per year, in one year P. calif ornicus was displaced 76 meters (2A) and 109 meters (2B) by I. humilis. It does not appear that this displacement is due to any overlap of a fundamental food dimension. At each sampling interval the mean area per colony of 7. humilis in the newly displaced territory was approximately 1400 m2 whereas the mean area per colony for the entire field increased from 1400 m2 to approximately 2600 m2 during the five year period (Table 1). There thus appears to be a minimal area for a colony of I. humilis in the newly acquired areas and as these colonies become established and increase in population density, the colony requires a larger area. Michener (1942) working with P. calif ornicus encountered a similar displacement by 7. humilis. He described in detail how in- dividual harvester ants would be set upon and killed by groups of 7. humilis. When temperatures are cool, Pogonomyrmex species tend to be sluggish and it is at this time that the Argentine ants torment the harvester ants as they forage around the nest (Michener, 1942). 1971] Erickson — Displacement of Native Ant 263 Table 2. Food resources of I. humilis compared with species it has displaced throughout the world. Main Degree of Species Food Source Omnivory Reference Iridomyrmex humilis Mayr sweet or fatty foods, tends aphids scale insects, grains + + + Wheeler, 1910 Eckert & Mallis, 1937 Creighton, 1950 Skaife, 1961 *Pogouomyrmex calif ornicus Buckley seed gatherers + Wheeler, 1910 Forel, 1928 Wildermuth & Davis, 1931 Cook, 1953 Van Pelt, 1966 Cole, 1968 Phe'.do'e megacephala F. sweet or fatty foods + + + Wheeler, 1910 Forel, 1928 *Pheidole grallipcs Wheeler seed gatherers + Eckert & Mallis, 1937 Cook, 1953 *V eromessor pergandci Mayr seed gatherers + Eckert & Mallis, 1937 Cook, 1953 Solenopsis saevissima Forel insects, fruits, grains, flowers, vegetables + + + Creighton, 1950 Cook, 1953 Solcnopsis geminata F. insects, fruits, grains + + Creighton, 1950 Fluker & Beardsley, 1970 ^Displaced in present study. + d~ T zzr highly omnivorous; + + = moderately omnivorous; + =: slightly omnivorous Should a harvester ant come upon an Argentine ant during the warmer parts of the day, the former grasps the smaller ant with its mandibles and stings it to death (Michener, 1942). At dawn, sun- set, or on a cloudy day the Argentine ants will attack and cling to the mandibles, legs, and antennae of the harvester ants and attempt to kill the larger ant. Observations made in the present study con- firm Michener’s discussion of the aggressive actions between the spe- cies. There were no significant differences in the mean monthly tem- perature or precipitation from month to month (i.e. — all the Janu- arys, etc.) over the course of the study. The vegetation studies 264 Psyche [December similarly showed that there was no significant difference in the order of dominance of the six plants mentioned. It does not appear that P. calif ornicus ameliorates the habitat as it does not clear vegetation as many harvester ant species do. I. humilis does not utilize the same nest sites as the displaced species and in fact, not a single I. humilis colony was found within two meters of an abandoned harvester ant colony. Pasfield (1968) found I. humilis displaced its neighbors at a max- imum rate of 274 meters (300 yards) per year in Australia. This value is higher than the 100 to 200 meters per year at Fort Shafter on the island of Oahu between 1940 and 1944 (Pemberton, 1944) or the average of 100 meters per year in the present study (Fig. 1). Fluker and Beardsley (1970) observed /. humilis displace P. megace- phala in Hawaii at about 66 to 100 meters per year. All these values seem low when compared to the displacement rate of 8 kilometers (5 miles) per year for native species by the fire ant S. saevissima in the Gulf states (Wilson and Brown, 1957). The effectiveness of competition in nature is best demonstrated by the impact of an invading species on the native fauna. It appears that here, there is a tremendous competition for nest space, which is the general case for highly aggressive territorial ant species such as Pheidole , Solenopsis, and Iridomyrmex (Wilson, 1971). Three as- pects of the populations biology of I. humilis gives this species a dis- tinct competitive advantage over the native harvester ants. The general aggressive nature of I. humilis as well as the large number of queens and method of proliferation allow these ants to move in and establish new colonies in a very short time. Raiding columns of workers clear the way and pioneer groups of workers and queens follow into freshly opened nest areas. Acknowledgements I would like to thank my uncle, Mr. Carl Erickson, for the gen- erous use of his property and to my cousins, Bob, Tom, David, and JoAnn for their assistance in various aspects of the study. Thanks are also due to Mrs. Mudie of the Department of Botany at San Diego State College for determination of the plant species and Dr. A. C. Cole, Jr. of the University of Tennessee for identification of the ant species. Mr. F. Slansky, Jr. and Drs. W. L. Brown, Jr., R. G. Helgesen, and P. P. Feeny read the manuscript and gave many helpful comments. A grant from the Grace Griswold Fund assisted with the publication costs. 1971] Erickson — Displacement of Native Ant 265 Literature Cited Brun, R. 1924. Das Leben der Ameisen. B. G. Teubner, Leipzig. 211 pp. Cole, A. C., Jr. 1932. Notes on the ant Pogonomyrmex calif ornicus Buckley (Hym: Formicidae). Entomol. News 43: 113-115. 1968. Pogonomyrmex harvester ants. A study of the genus in North America. University of Tennessee Press, Knoxville. 222 pp. Cook, T. W. 1953. The ants of California. Pacific Books. Palo Alto, California. 462 pp. Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool. Harv., 104. Cambridge, Mass. 568 pp. Crowell, K. L. 1968. Rates of competitive exclusion by the Argentine ant in Bermuda. Ecology 49: 551-555. Eckert, J. E. and A. Mallis 1937. Ants and their control in California. Cal. Agric. Exp. Sta. Circ. 342. Erickson, J. M. 1972. Mark-recapture techniques for population estimates of Pogono- myrmex ant colonies: An evaluation of the P32 technique. Ann. Ent. Soc. Amer. 65: 57-61. Fluker, S. S. and J. W. Beardsley 1970. Sympatric associations of three ants: Iridomyrmex humilis, Phei- dole megacephala and Anoplolepis longipes in Hawaii. Ann. Ent. Soc. Amer. 63 : 1290-1296. Forel, A. 1928. The social world of the ants compared with that of man. G. P. Putnam’s and Son’s, Ltd. London. 445 pp. Haskins, C. P. and E. F. Haskins 1965. Pheidole megacephala and Iridomyrmex humilis in Bermuda — Equilibrium or slow replacement? Ecology 46: 736-740. Michener, C. D. 1942. The history and behavior of a colony of harvester ants. Sci. Monthly 55: 248-258. Morley, D. W. 1953. The ant world. Penguin Books, Ltd. London. 191 pp. Newell, W. 1908. Notes on the habits of the Argentine ant or “New Orleans” ant, Iridomyrmex humilis Mayr. Jour. Econ. Ent. 1: 21-34. Pasfield, G. 1968. Argentine ants. Aust. Natur. Hist. 16: 12-15. Pemberton, C. E. 1944. (Notes and exhibitions). Proc. Hawaiian Entomol. Soc. 12: 25. Shapley, H. 1920a. Preliminary report on Pterergates in Pogonomyrmex californi- cus. Proc. Nat. Acad. Sci. 6: 687-690. 1920b. Notes on Pterergates in the California harvester ant. Psyche 27: 72-74. 266 Psyche [December Skaife, S. H. 1961. The study of ants. Spottiswoode, Ballantyne, and Co. Ltd. Lon- don. 178 pp. Smith, M. R. 1947. A generic and subgeneric synapsis of the United States ants, based on the workers (Hym: Formicidae). Amer. Mid. Natl. 37: 521-647. Stoll, O. 1898. Zur kenntnis der geographischen verbreitung der maeisen. Mitt. Schweiz. Entomol. Ges. 10: 120-126. Titus, E. S. G. 1905. Report on the “New Orleans” ant Iridomyrmex humilis Mayr. U. S. Dept. Agr. Bur. Ent. Bull. 52: 79-84. Tulloch, G. S. 1930. An unusual nest of Pogonomyrmex. Psyche 37: 61-70. Van Pelt, A. F. 1966. Activity and density of old field ants on the Savannah River Plant, South Carolina. J. Elisha Mitchell Sci. Soc. 82: 35-43. Wheeler, W. M. 1906. On certain tropical ants introduced into the United States. En- tomol. News 17: 24. 1926. Ants: their structure, development and behavior, ed. 3. Colum- bia Univ. Press. New York. 566 pp. 1933. Colony founding among ants. Harvard Univ. Press. Cambridge, Mass. 179 pp. WlLDERMUTH, V. L. AND E. G. DAVIS 1931. The red harvester ant and how to subdue it. U. S. Dept. Agr. Farm. Bull. No. 1668. 21 pp. Wilson, E. O. 1951. Variation and adaptation in the imported fire ant. Evolution 5: 68-79. 1971. The insect societies. Harvard University Press. Cambridge, Mass. 548 pp. Wilson, E. O. and W. L. Brown, Jr. 1957. Recent changes in the introduced population of the fire ant Solenopsis saevissima. Evolution 12: 211-218. Wilson, E. O. and R. W. Taylor 1967. The ants of Polynesia. Pacific Insects Monogr. 14. Bishop Museum, Honolulu, Hawaii. Woodword, E. W. 1905. The Argentine ant in California. Cal. Agr. Exp. Sta. Circ. 38. 1910. The control of the Argentine ant. Cal. Agr. Exp. Sta. Bull. 207. Wynne-Edwards, V. C. 1963. Animal dispersion in relation to social behavior. Oliver and Boyd. London. 653 pp. Zimmerman, E. C. 1940. The Argentine ant in Hawaii. Proc. Hawaiian Ent. Soc. 11: 108. ADDITIONAL INSECTS IN PENNSYLVANIAN CONCRETIONS FROM ILLINOIS By F. M. Carpenter1 and Eugene S. Richardson, Jr.2 The ironstone nodules from the Francis Creek Shale (Mid- dle Pennsylvanian) of Illinois continue to yield many interesting and significant insects. The specimens described in this paper were obtained in former mine pits in Grundy, Will and Kankakee Counties3, and have been made available to us by the following col- lectors, who have been unusually successful in finding insects: Mr. Jerry Herdina, Berwyn, Illinois; Mr. Joseph Makowski, Chicago; Helen and Ted Piecko, Chicago; Mr. Paul Tidd, Mendota, Il- linois; and Mr. and Mrs. Francis Wolff, Park Forest, Illinois. We are most grateful to them for their cooperation in loaning their specimens to us for study and their patience in waiting for the re- results. Special thanks are extended to Mr. Jerry Herdina and to Helen and Ted Piecko for allowing us to photograph and to make a thorough examination of all the insects in their collections. Sub- sequent papers in this series will deal with additional specimens which they and other local collectors have found.4 The insects discussed in this paper belong to four orders: Palae- odictyoptera, Megasecoptera, Prodonata and Protorthoptera. All of these specimens are of unusual interest for one reason or another, but two of the specimens in Mr. Herdina’s collection are of excep- tional significance; one is the first unquestioned nymph of the order Palaeodictyoptera that has been found and the other is a brachy- pterous adult of a protorthopteron. Order Palaeodictyoptera Family Lycocercidae Handlirsch Among the Palaeodictyoptera in the collections at hand there are two species referable to the family Lycocercidae. One of these is in the Herdina collection and the other is in the Field Museum col- lection. harvard University, Cambridge, Mass. 02138 2Field Museum of Natural History, Chicago, Illinois 60605 3Pit Six is about three miles northeast of Coal City, about on the Will- Grundy County line; Pit Eleven is about three miles south of Braidwood, on the Will-Kankakee County line, in northern Illinois. 4Partial financial support of this research is gratefully acknowledged to the National Science Foundation: Grant No. GB27333, F. M. Carpenter, Harvard University, principal investigator; and Grant No. GB8266, R. G. Johnson, University of Chicago, and E. S. Richardson, Jr., Field Museum of Natural History, principal investigators. 267 268 Psyche [December The Herdina specimen is a nymph with its wing pads held ob- liquely away from the body (figure i ). In this respect the wing pads resemble those of the megasecopterous nymph, Mis chapter a doug- lassi , described from the same formation a few years ago (Carpenter and Richardson, 1968). However, the venation in the present nymph is sufficiently well indicated in one of the wing pads to show that the insect is a member of the Palaeodictyoptera and that it prob- ably belongs to the family Lycocercidae. This is of unusual signifi- cance, since all of the nymphs which have previously been assigned to the Palaeodictyoptera actually belong elsewhere or have very du- bious palaeodictyopterous affinities. Although until recently the family Lycocercidae has been known only from the Upper Car- boniferous of Europe, it has lately been found in Pennsylvanian de- posits of New Mexico (Carpenter, 1971). Genus Lycodemas, new genus5 It seems advisable to refer this nymphal form to a separate genus rather than to assign it to one of the three genera of the family al- ready recognized (Kukalova, 1969) ; the venation of the nymphal wing is not fully developed and does not provide a satisfactory con- cept of the adult venational pattern for comparison with the other known lycocercid species. Since the posterior media (MP) is much less developed in the nymph than it is in the known adults of Lyco- cercus and the related genus Apopappus, we consider that this (i.e., the less developed MP) should be the diagnostic feature of the genus Lycodemas. The more obvious peculiarities of the nymphal wing, such as the narrow proximal region, are aspects of the immature state of its development. Type species: Lycodemas adolescens, n. sp. Lycodemas adolescens, n.sp. Figures 1-3 Length of fore wing pad, 11 mm.; width, 3 mm.; length of body from front of mesothorax to end of abdomen (as preserved), 26 mm. ; width of first abdominal segment, 6 mm. The venation, as faintly indicated in a fore wing pad, is represented in figure 2. The wing sheath (w) is conspicuous around the wing except in the ap- ical region, where it is narrow. The subcosta (SC), which extends nearly to the apex of the wing, has a series of blunt projections or tubercles near the middle of the wing; these may possibly have been setal bases in the living insect. The precise origin of Rs from R is 5The generic name is derived from a combination of lykos (wolf) and demas (body) and is considered neuter. 1971] Carpenter & Richardson — Pennsylvanian Insects 269 Figure 1. Lycodemas adolescens, n.sp. Photograph of holotype, No. H413, Herdina collection. 270 Psyche [December Figure 2. Lycodemas adolescens, n.sp. Drawing of holotype. W, wing sheath ; other abbreviations are the conventional venational symbols. not visible in the wing pad but it presumably arises near the basal part of the wing, as indicated in the drawing. Rs has ten primary branches, with one or two secondary branches; the area between MA and the preserved part of MP presumably had some anterior branches from MP but they are not discernible in the specimen ; in any event, MP seems to be less developed than in other genera of this family. CuP is extensively branched and conforms to the usual pattern in the Lycocercidae. The narrow form of the wing basally is probably due to the immature nature of the wing pad. Cross veins are discernible in a few small areas; they are apparently numerous and reticulate. The general venational pattern, as far as it can be determined, is shown in figure 2.6 The venation of the hind wing appears to be very similar to that of the fore wing. The wing pads on one side of the body seem some- what broader in the photograph (figure i) than the other pair, but this is deceptive; the matrix was chipped away for some distance from these wing pads, giving the impression that the exposed area was actually part of the wing. The wing pads on both sides are 3 mm. wide. Holotype: No. Hd.i3a, b; collected in Pit Eleven; in the collec- tion of Mr. Jerry Herdina, Berwyn, Illinois. The most interesting feature of this nymph is the position of the wings with respect to the body. The arrangement is very similar indeed to that which we have described in the megasecopterous nymph, Mischoptera douglassi Carpenter and Richardson. As in the latter, the wing pads of Lycodemas have no contact with the body fiWe are indebted to Dr. Jarmila Kukalova-Peck for the preparation of this figure. 1971] Carpenter & Richardson — Pennsylvanian Insects 27 1 Figure 3. Lycodemas cf. adolescens. Drawing of specimen No. H110, Herdina collection. except in the articular area of the wings themselves, in contrast to the structure of the wing pads in the nymphs of Recent insects, in- cluding those of the Ephemeroptera. A discussion of the evolution- ary significance of this position of the wing pads is contained in our previous paper cited above and in Dr. Kukalova’s account of the Permian may-fly nymphs ( 1968). The type of L. adolescens is the first nymph known that can with- out question be assigned to the Palaeodictyoptera ; the nature of the venation in the wing pad seems to provide conclusive evidence of the palaeodictyopterous nature of the insect, especially in view of the homonomous condition of the wings. It is regrettable, of course, that more of the body structure is not preserved. What little is visible gives no indication that the nymph was modified for an aquatic existence; this is consistent with the more extensive evidence pro- vided by the nymphs of Mischoptera for the Megasecoptera. In the Herdina collection there is also a single wing (No. Hi 10, Pit Eleven) ; it is obviously a nymphal wing, since it is included in a sheath (figure 3). It has a length of 27 mm. and a maximum width of 7.5 mm. and is therefore three times the size of adolescens. In all probability, although no proof can be given, this fossil is an older nymph of adolescens or of another species of the genus. The wing has the slightly falcate shape that is present in the mature wings of the Lycocercidae (see Kukalova, 1968, figs 33 and 34). The specimen in the Field Museum collection is being assigned tentatively to the Lycocercidae. Assignment to this family is based on what little is preserved of the venation of one wing, probably about half the entire wing; MP is extensively branched, much more so than in the Dictyoneuridae; CuA is unbranched but CuP is well developed; the anal veins are apparently numerous. On the basis of the wings alone, especially in view of their fragmentary na- ture, this insect would hardly warrant formal description and nam- 272 Psyche [December Figure 4. Notorachis cwolfforum, n.sp. Drawing of holotype, No. PE 21699, Field Museum; ant, antenna; hind wing represented by dotted lines. Black circles on abdomen mark the locations of spines on the posterior edges of the tergites ; the probable lateral margins of the abdomen are indicated by broken lines. ing. However, the prothoracic lobes of this species are extraordi- narily modified, nothing comparable to them having previously been observed in the Palaeodictyoptera. Since this insect is of unusual interest, generic and specific names are being assigned. Genus Notorachis, new genus7 This genus is apparently related to Lycocercus. The outer mar- gins of the pronotal lobes are heavily sclerotized and form a series of long spines; the more basal portion of each lobe is less sclerotized but there is a double row of blunt setae or tubercles extending trans- versely across each lobe (preserved as pits in the obverse). In Lyoo- 7The generic name is derived from a combination of noton (back) and rachis (spine) and is considered feminine. 1971] Carpenter Richardson — Pennsylvanian Insects 273 cercus, although the lobes are prominent, they are membranous or nearly so with a radiating series of vein-like structures, as in most other Palaeodictyoptera ; paranotal spines are unknown in Lycocercus. Type species: Notorachis wolff drum, n.sp. Notorachis wo Iff orum, n.sp. Figures 4-6 Length of wings, as preserved, 22 mm.; maximuum width of fore wing, 10 mm.; width across pronotal lobes, including spines, 20 mm. The fossil consists of the dorsal aspect of the head, pronotum, and meso- and metathorax; a few segments of the abdomen are in- dicated and the wings are outstretched in the palaeopterous position. The fore and hind wings on each side overlap but the basal part of the venation of one wing is discernible for the most part. The costal space seems to be of uniform width ; the precise origin of the radial sector is not preserved ; MA obviously arises before the ori- gin of the radial sector and is probably unbranched, although its distal part is not preserved; MP has at least five terminal branches and CuP is even more extensively branched. The hind wing is almost completely covered by the fore wing and only a few veins can be distinguished. The most conspicuous of the body structures are the pronotal lobes (figures 5 and 6), which are not sharply set off from the rest of the prothorax. The margins of the lobes are heavily sclerotized and bear seven prominent spines on each side, the first and last of these being somewhat shorter than the others. The two posterior spines of each side are directed somewhat dorsally as well as laterally. Extending along the median axis of each lobe is a double row of setae (preserved in the reverse of the fossil as fine pits). The sur- face of the slightly elevated margin of the pronotal lobe is smooth along the posterior edge of the lobe; on the anterior edge it is occu- pied by scattered setae or fine tubercles. Similar setae are present on the proximal third of all the spines. Somewhat before mid-length of each spine the setae are smaller and directed posteriorly; in the distal potion of each spine, the setae are succeeded by strong, longitudinal ridges. The head is visible just anterior to the lobes but there is no sharp separation in the fossil between the head and thorax; two relatively large, circular structures are visible on the sides of the head, pre- sumably the compound eyes. A four-millimeter portion of one an- tenna is preserved; it appears to arise from above the center of the 274 Psyche [December Figure 5. Notorachis wolfforum , n.sp. Photograph of holotype (reverse), showing overlapping of wings and the general form of the body. eye and curves laterally. Its proximal segments are less than half as long as wide; the distal segments are relatively longer and are clad with fine prostrate setae. The preserved part of the antenna is surprisingly thin and delicate. The meso- and metathorax seem to form a compact unit, each of the segments being relatively short. Portions of two segments, prob- ably tibia and tarsus, of a weakly sclerotized fore leg, are preserved anterior to one of the pronotal lobes. The portion of the tibia is about three millimeters long and the tarsus nearly two millimeters. The tibia is armed with short spines directed distally, and the tarsus with setae. There is no reason to suppose that this leg was adapted for anything but slow walking. The abdomen is indicated by a series of faint, transverse depres- sions that mark the segmentation. It is preserved for a length of 1971] Carpenter & Richardson — Pennsylvanian Insects 275 about 23 mm., to the edge of the nodule, and it undoubtedly orig- inally extended beyond that point. Pairs of strong, flat, setiferous spines project dorsally and somewhat posteriorly from the abdomen, arising directly on the posterior margins of the tergites. The first spine is short; the second, third and fourth are nearly as long as the width between them; the fifth is short; a very slight projection marks the position of the sixth and there are none in the more posterior positions. These spines do not appear to mark the lateral edges of the tergites, which do not have their full widths preserved. The spines that are preserved in the fossil are about 3.5 mm. apart laterally and were almost certainly a little to each side of the median longitudinal axis of the abdomen. Other spines along the posterior tergal margins were probably present near the sides but can only be assumed since the lateral portions of the segments are broken away. In all probability the tergal spines in Notorachis were similar to those already described in the megasecopterous Mischoptera douglassi , though the strong, dorsally projecting spines just described are flat- tened parallel to the body axis rather than parallel to the surface of the tergites. (See Carpenter and Richardson, 1968, p. 306). It is pertinent to note that prothoracic spines were also present in Mis- choptera. Holotype: No. PE21699, Field Museum of Natural History; from the collection of Mr. and Mrs. Francis A. Wolff, Park Forest, Illinois, who collected the unique specimen in Pit Eleven. The spe- cies is named for both Mr. and Mrs. Wolff. This insect is unusual, among the Palaeodictyoptera so far known, in having elaborately modified pronotal lobes; the only other species that may approach Notorachis wolff orum in this respect is Stilbocro- cis heeri (Goldenberg) (Upper Carboniferous of Germany), which is, however, a member of the Dictyoneuridae. In Stilbocrocis the lobes are smaller than in Notorachis and possess fewer spines.8 The most puzzling aspect of this fossil is the shortness of the pterothorax and the consequent overlapping of the fore and hind wings. Since this overlapping is symmetrical for both pairs of wings, it is almost certainly not due to distortion during preservation. Fur- thermore, the thorax itself is so short that the overlapping would 8 Stilbocrocis has consistently been represented in the literature as having large pronotal lobes, bearing a series of radiating veins within the lobes themselves. From a recent examination of the type specimen (on deposit in the Natural History Museum at Bonn), I am convinced that the radiat- ing ridges are actually spines which project beyond the edges of the lobes. A detailed account of this fossil will be published elsewhere. (FMC). 276 Psyche [December Figure 6. Notorachis 'wolfforum, n.sp. Photograph of pronotum and portions of some other body struc- tures; width across pronotum, including spines, 20 mm. (Holotype). 1971] Carpenter & Richardson — Pennsylvanian Insects 277 Figure 7. Homaloneura dabasinskasi Carpenter. Photograph of speci- men in Tidd collection. The arrow points to the cross-section of the beak. seem to have been unavoidable in the living insect. On the other hand, such an arrangement of the wings would seem to be very in- efficient mechanically. It is in this connection interesting to note that the only other palaeodictyopteron known that shows a similar overlapping of the fore and hind wings and a comparable, short thorax, is Stilbocrocis heeri, already mentioned. This condition of the wings is readily seen in the photograph of the specimen given by Guthorl (1934), as well as in the figures by Goldenberg (1854), Schlechtendal (1912) and Guthorl (1934); it is not shown in Handlirsch’s highly imaginative reconstruction of the insect (1920), in which the wings are represented in normal position. As noted above, Stilbocrocis has the wing venation characteristic of the Dic- tyoneuridae, whereas Notorachis has a very different vena.tional pat- tern, allying it to the Lycocercidae. 278 Psyche [December Figure 8. Homaloneura dabasinskasi Carpenter. Drawing of specimen in Tidd collection. Family Spilapteridae Brongniart Homaloneura dabasinskasi Carpenter Figures 7-9 Homaloneura dabasinskasi Carpenter, 1964: 1 1 7 A second specimen of this species, also in an ironstone concretion from the Francis Creek Shale, was collected in 1971, by Mr. Paul Tidd of Mendota, Illinois, in Pit Eleven and is now contained in Mr. Tidd’s collection (figure 7) It consists of the basal part of a fore wing, portions of the head and thorax and traces of a hind wing. Determination of the species is based on the fore wing; except for minor points, such as a deeper fork on CuP, the venation is like that of the holotype. The wing markings, characteristic of the spe- cies, are faintly preserved. Some of the thoracic structures are of interest, since they are known in only a few genera of the order. The thoracic segments are unequal (figure 8) the prothorax being slightly the smallest of them. There are no indications of the prothoracic lobes, which have previously been described in Homa- loneura (Kukalova, 1969). The fore legs are preserved in the ob- verse half of the specimen; they are robust and possess numerous 1971] Carpenter & Richardson — Pennsylvanian Insects 279 Figure 9. Homaloneura dahasinskasi Carpenter. Photograph of cross- section of beak; specimen in Tidd collection. The arrow points to the median stylet. setae, as in other Palaeodictyoptera. The coxae are prominent and in the obverse half of the fossil extend anteriorly from the pro- thorax. The tarsal segmentation is not discernible, except for the first tarsomere. The head is only vaguely indicated in the fossil (reverse half) but just anterior to the head itself there is a transverse section of the sucking beak, this being visible at the point where the two halves of the concretion separated. In order to appreciate the sig- nificance of this section, it must be borne in mind that conclusive evidence has now been provided, in the publications of Crampton (1927), Lameere (1933), Laurentiaux (1952 and 1953) and Ku- kalova (1969 and 1970), showing that all of the Palaeodictyop- tera possessed a haustellate beak. According to observations pre- viously made (and summarized by Kukalova, 1970), the beak consisted of four slender stylets, apparently supported ventrally by a long and somewhat broader labium, usually transversely ridged. 28o Psyche [December The beak was not directed ventrally from the head but anteriorly and ventrally. The section of the beak in the new specimen of dcibasinskasi is about 3 mm. in front of the anterior margin of the head (figures 7 and 8) ; it presumably shows the structure of the beak close to the head, not at a point further along the beak. As shown in the photo- graph (figure 9), there is a vague circular area of discoloration which appears to mark off the area of the beak. Within this can be seen sections of two pairs of stylets; these are very dark and distinct. The anterior pair, which are somewhat the larger, are presumably the mandibles, and the other pair, the maxillae. These four stylets are symmetrically arranged within the beak and with respect to each other. However, in addition to these there is a fifth stylet, midway between the maxillae and slightly more posterior. The sym- metry of the pattern formed by these five structures, as seen in sec- tion, is very striking. It is virtually certain that the fifth stylet is one of the mouthparts, presumably derived from the hypopharynx; the latter, of course, is a median structure and it does function as one of the stylets of the haustellate mouth-parts of some Diptera and Hemiptera. The labium, which as noted above is broader and thicker than the stylets in the Palaeodictyoptera, is missing from the section, as are the large maxillary palpi. In any event, it now seems clear that in the spilapterid Palaeodictyoptera, at least, there were five stylets in the beaks, with an arrangement which closely parallels that in some Recent insects. Genus Spilaptera Brongniart In the Herdina collection there are two spilapterids that appear to belong to the same species. Only two representatives of this family have been previously reported from the Francis Creek Shale, or, in fact, from all Pennsylvanian deposits in North America: Homaloneura dabasinskasi Carpenter and Mcluckiepteron luciae Richardson. The new specimens belong to an undescribed species, which we are assigning to Spilaptera , otherwise known only from the Commentry shales of France. Spilaptera differs from Homaloneura in having several long, sig- moidal and oblique cross veins between Ri and Rs, beyond mid- wing. In Homaloneura these cross veins are straight or nearly so and they are transverse, or only slightly oblique. CuP is usually unbranched in Homaloneura , although it may be forked distally, very near its termination. In Spilaptera CuP ranges from unbranched to deeply forked. The new species described below has the sigmoidal 1971] Carpenter & Richardson — Pennsylvanian Insects 281 sc - Figure 10. Spilaptera americana, n.sp. Drawing of holotype, No. H 463, Herdina collection. Length of wing as preserved, 38 mm. and oblique cross veins present and it is accordingly placed in the genus Spilaptera. The several other known genera of Pennsylvanian Spilapteridae, including Mcluckiepteron , have very distinctive fea- tures, such as numerous cross veins, more extensive branching of veins, pronounced tapering of wing, etc. Spilaptera americana, n.sp. Figure 10 Length of wing (as preserved) 38 mm.; maximum width, 15 mm. Costal margin distinctly concave, slightly more so than in the type species of the genus, S. packardi ; Sc extending to wing apex; Rs with 6 terminal branches, none forked in types; MA with 4 branches, CuA with 5 ; CuP deeply forked, almost to its origin. Cross veins not numerous, about as many as in S. packardi , forming poorly de- fined rows. The sigmoidal cross veins between Ri and Rs are long and more oblique than in S. packardi. The wing markings consist of darkened spots at most of the cross veins. Holotype: No. H463, in the collection of Mr. Jerry Herdina, Berwyn, Illinois; collected in Pit Six. This is a very well preserved fore wing, lacking the apex and the basal region ; the wing was ob- viously broken along the line of the cuticular thickenings, as in the type of H. dahasinskasi. Paratype: No. H459, in the Herdina collection; collected in Pit Six. This consists of a less clearly preserved and more fragmentary wing; the preserved part is 33 mm. long and 15 mm. wide. Although these two specimens have virtually an identical venation, they do show one difference. As Dr. Kukalova has pointed out ( 1969, p. 166), the spilapterids often have some kind of supporting struc- 282 Psyche [December Sc Figure 11. Eubrodia dabasinskasi Carpenter. Composite drawing of entire wing, based on specimen HTP 433, Piecko collection, and the holo- type. tures in the basal third of the wings. There is usually a thickened ridge, resembling a vein, between M and Rs, close to the origin of the latter and in an oblique position. From her study of the Com- mentry spilapterids, Dr. Kukalova concluded that the ridge was present in all Homaloneura but was absent in Spilaptera. In S. packardi (type species of the genus), the oblique ridge was absent, although normal transverse cross veins were present in the corres- ponding area of the wing. In the only other species of Spilaptera previously known (libelluloides) the nature of this particular area of the wing is unknown because that part of the wing is not pre- served. In the holotype specimen of S. am eric ana > described above, the cuticular, oblique ridge is present (see figure 10), but in the para- type there is no oblique structure, only the normal cross veins. In view of the very close similarity of these two fossils in other re- spects, we are inclined to infer from this scant evidence that the oblique ridge is, in fact, a modified cross vein and that the degree of its inclination may vary within the species. Order Megasecoptera Family Brodiidae Handlirsch Eubrodia dabasinskasi Carpenter Figures 11-13 Eubrodia dabasinskasi Carpenter, 1967:73 In the collection of Helen and Ted Piecko, there is an interesting specimen of Eubrodia dabasinskasi , a species originally described from a single wing, lacking only the base (Carpenter, 1967). The new specimen (HTP433, Pit Eleven) consists of the body, which is 1971] Carpenter & Richardson — Pennsylvanian Insects 283 Figure 12. Eubrodia dabasinskasi Carpenter. Photograph of specimen No. HTP 433, Piecko collection; the arrow points to the filamentous struc- tures associated with the insect. 284 Psyche [December poorly preserved, and the basal portions of three wings. Photographic enlargement of these wing bases shows that they fit perfectly the wing of the type specimen of E. dabasinskasi. It is now certain that the wing of this insect was petiolate, much as in Brodia. In figure 11 the wing of the holotype of dabasinskasi is combined with that of the Piecko specimen to show the entire wing. As shown in the figure, the serrated margin extends to the base of the petiole, and a definite group of reticulated cross veins occurs between iA and CuP.9 The body structures of the new specimen of dabasinskasi are vaguely preserved, showing the general outline of the head, thorax and part of the abdomen (figure 12). The head and the thoracic segments are each 5 mm. long. What makes the specimen of special interest is the presence of sev- eral very long, filamentous structures that rest along one side of the abdomen and extend anteriorly to about the level of the fore wing (figure 13). We assumed at first that these were of plant origin, but two paleobotanists, Dr. Sergius Mamay of the U. S. Geological Survey, and Professor Elso Barghoorn of Harvard Uni- versity, both well acquainted with Upper Carboniferous plants, have expressed their convictions, after examination of the specimen, that the filaments were not parts of plants. As preserved, these structures are not carbonized but have the same surface texture, including the rugosity, as the brodiid integument. Similar but fewer filaments were associated with the megasecopterous nymph ( Mischoptera doug- lassi), already described from the ironstone nodules (Carpenter and Richardson, 1969). Furthermore, Dr. Jarmila Kukalova-Peck in- forms us that such filaments are associated with the specimens of Megasecoptera in the Upper Carboniferous shales of Commentry, France, and with Palaeodictyoptera which she has collected in the Permian deposits of Moravia. Although it is difficult to see how these filaments can be part of the insects concerned, judgment on this possibility should wait until Dr. Kukalova-Peck has published on the evidence obtained from the study of the European material at her disposal. Order Protodonata A specimen of this order was collected in an ironstone nodule by Mr. Joseph Makowski, of Chicago. It is of unusual interest, since the Protodonata have previously been represented in the Francis The cross veins in the rest of the wing of Eubrodia are shown in the photograph of the holotype (Carpenter, 1967, p. 67). 1971] Carpenter & Richardson — Pennsylvanian Insects 285 Figure 13. Eubrodia dabasinskasi Carpenter. Photograph of filamentous structures associated with specimen No. HTP 433, Piecko collection. 286 Psyche [December Figure 14. Oligotypus mako'wskii, n.sp. Drawing of holotype, No. IWJM-I-C, Makowski collection. Length of wing as preserved, 65 mm. Creek shales by only a single species, Paralogopsis longipes Hand- lirsch. The new fossil clearly belongs to the family Paralogidae. Family Paralogidae Handlirsch This family is characterized by having the subcosta terminate at about mid-wing, instead of extending to the wing apex, as in other families of protodonates ; and by having R.2 + 3 and R4 + 5 widely divergent. Two genera are known: Paralogus Scudder, from Upper Carboniferous shales (Allegheny Series) of Rhode Island, and Oligotypus Carpenter from Lower Permian deposits in Kansas and Oklahoma. The new fossil is referable to Oligotypus. Oligotypus makowskii, n.sp. Figure 14 Length of wing, as preserved, 65 mm.; maximum width, 18 mm.; estimated complete length, 90 mm. Similar to the type species of the genus, tillyardi , but with the first forking of Rs occurring basal of the termination of Sc; with the area between MA and CuP about twice as wide as it is in tillyardi; and with the hind margin of the wing more strongly curved than in tillyardi. The cross veins are not so numerous as in tillyardi and the cellules formed by the cross veins are a little larger than, and not so numerous as, in tillyardi. Other venational details are shown in figure 14. Holotype: No. IWJM-I-C, in the collection of Mr. Joseph Makowski, Chicago; collected in Pit Eleven. The specimen consists of about three-fourths of a wing; it is very well preserved, with the convexities and concavities of the veins very distinct. The applica- tion of water or alcohol brings out the cross veins very distinctly. It is not possible to determine with certainly whether the fossil is a fore or hind wing; the strongly convex margin suggests the latter. 28? 1971] Carpenter & Richardson — Pennsylvanian Insects Handlirsch’s Paralogopsis longipes , also from the ironstone nod- ules, may belong to the family Paralogidae, as suggested by Hand- lirsch ( 1 9 1 1 ) . However, since the species is known only from a small fragment that lacks the areas of the wings including the struc- tures indicative of the family, we consider any attempt to make fam- ily assignment really futile. In fact, Handlirsch’s publication of generic and specific names for such a small fragment is regrettable, for it is doubtful that even a good specimen of the insect, if one were ever found, could be rcognized as conspecific with his type. In any event, should longipes turn out to belong to the Paralogidae, obvious differences in the origins of Ri, Rs and MA, and in the branching of iA, separate that species from makowskii. Order Protothoptera Included in the Herdina collection is a specimen of a remarkable protorthopteron, which appears to be a brachypterous adult. Its venational pattern, which is similar in both fore and hind wings, resembles very closely that of the Cacurgidae, but in its total struc- ture the insect represents an undescribed family. Family Herdinidae, new family This is related to the Cacurgidae on the basis of the following venational features: CuA and MP are fused basally and CuP is forked at least once, with its more anterior branch coalescing with CuA. The distinctive characteristics of the new family are as fol- lows: the base of the costal area is strongly sclerotized and is with- out veins; Sc and Ri teminate well before the apex of the wing; Rs arises at about mid-wing, with at least two main branches; MP diverges from the stem of CuA well before mid-wing and gives rise to at least one main branch. Cross veins are numerous and they form, with longitudinal veins, a coarse net- work over the entire wing. The hind wing is much smaller than the fore wing but with the exception of the narrow costal area the venational pattern is similar to that of the fore wing; there is no anal fan or enlarged anal area. Small tubercles are distributed over the longitudinal and cross veins of both the fore and hind wings. The prothorax was apparently large and possessed a cordate shield. This family is readily distinguished from the Cacurgidae by the reticulate nature of the cross veins and by having Rs much less ex- tensively developed, in addition to the obvious differences in the form of the fore wing 288 Psyche [December Figure 15. Herdina miri ficus, n.sp. Photograph of holotype, No. H 412a, Herdina collection (obverse). 1971] Carpenter & Richardson — Pennsylvanian Insects 289 Figure 16. Herdina mirificus, n.sp. Photograph of holotype, No. H 412b, Herdina collection (reverse). 290 Psyche [December Figure 17. Herdina mirificus, n.sp. Drawing of holotype, No. H 412a, b, Herdina collection. 1971] Carpenter & Richardson — Pennsylvanian Insects 291 Genus Herdina, new genus Fore and hind wings short, the fore wing extending to about the middle of the abdomen.10 Fore wing: longitudinal veins, including Sc and Ri, somewhat irregular, cross veins unusually strong, nearly as thick as the longitudinal veins; costal area moderately broad, with two rows of irregular cells along most of the area; Rs forking into two main branches shortly after its origin at about mid-wing; secondary branching irregular, with short rows of convex intercalary veins between the normal branches of Rs; MP apparently with one prominent branch arising at about mid-wing; CuA apparently un- branched; CuP forked shortly after its origin, one branch directed towards the hind margin ; between CuP and CuA is a short, irreg- ular intercalary vein; anal veins very weakly developed and irregu- lar. Hind wing: costal area narrow; venational pattern basically similar to that of the fore wing, including CuA and CuP. Type species: Herdina mirificus , n.sp. The genus is named for Mr. Jerry Herdina, of Berwyn, Illinois, in recognition of the contribution which he has made to insect paleon- tology through his extensive and remarkable collection of arthropods, preserved in the Mazon Creek nodules. Herdina mirificus, n.sp. Figures 15-20 Fore wing: length, 8 mm.; width, 3.5 mm.; hind wing: length, 5.5 mm.; width, 2.3 mm.; length of prothorax, 5 mm.; length of abdomen, 10 mm. It is virtually impossible to assign other specific characters for this species; venational details are shown in figures 17 and 18. As can be seen from these figures, there are some differ- ences in the venation of the two right wings, such as in the amount of the irregularity of the longitudinal veins and the shapes of the various cells of the wings; but these are within the normal range of variation in the species of Recent Orthoptera. Holotype: No. H 412a and b (obverse and reverse) in the col- lection of Mr. Jerry Herdina, Berwyn, Illinois; it was collected at Pit Eleven. The specimen consists of two fore wings and one hind wing, as well as parts of the thorax and abdomen.* * There are several remarkable and puzzling features about this fossil. As can be seen from comparison of the obverse and reverse 10The brachypterous condition is here treated as a generic characteristic but it could equally well be a specific or an individual trait. *A second specimen of Herdina has just been found (5/27/72) in the ironstone nodules (Wolff collection). It will be discussed in another article. 292 Psyche [December Figure 18. Herdina tnirificus, n.sp. Drawing of fore and hind wings of holotype. halves of the specimen, (figures 15 and 16), the counterparts appear to be very different. The reverse (figure 15, with the subcosta con- vex) shows the wings and thorax clearly; there is a large pronotum expanded into a dorsal shield, not unlike that known in some other Protorthoptera ; only one side of the shield is preserved in the speci- men. The pronotum and the rest of the thorax are apparently strongly sclerotized; the other thoracic and the abdominal segments are apparently preserved in a somewhat twisted position, so that parts of a pleuron and of the tergum of each segment are visible. The full size of the abdomen is not indicated in the reverse but is shown in the obverse (figure 16). The wings are very clearly preseved in the reverse. They are not so distinct in the obverse but their convexities or concavities are very clear. In the reverse half, all of the veins, including the cross veins, possess prominent pits or depressions, these being almost con- tiguous along the veins; they are especially evident in the reverse half, where the pits are filled with a white matrix (figure 20). In the obverse, which represents the wing as seen from above, the struc- 1971] Carpenter £sf Richardson — Pennsylvanian Insects 293 Figure 19. Herdina mirificus, n.sp. Photograph of fore and hind wings of holotype, No. H 412a, Herdina collection (obverse). tures are in the form of bluntly rounded projections or tubercles; they do not seem to be setae.11 At first glance, one might consider the type specimen of Herdina mirificus , with its short wings, to be a nymphal form, rather than an adult. However, there are several major points against this in- terpretation. The most important of these is the obvious nature of the wings, themselves; they are not wing pads, included in a wing sheath, but are fully sclerotized, with a definite venational pattern, including all cross veins. Furthermore, the wings themselves are not like those of any neopterous nymph known; they are independent of each other and not attached to the adjacent part of the thoracic wall of the insect, as they are in all Recent nymphal forms. It might be argued that the insect represents a newly hatched adult in which the wings had not yet reached full development. If this were the case, it seems most unlikely that the veins in the wings would be so dis- tinctly and clearly preserved. Actually, the occurrence of brachyp- tery among the Protorthoptera is not surprising, in view of the 1JIf figure 20 is held in an inverted position, the veins will appear to most readers as they are in the obverse half of the fossil. 294 Psyche [December Figure 20. Herdina mirificus, n.sp. Photograph of central portion of fore wing of holotype, No. H 412a, Herdina collection, showing the details of veins and the tubercles along the veins. frequency with which this condition is found among existing orthop- teroids, such as the Orthoptera, Phasmatodea., Blattodea, etc. Although the slender band of sclerotization at the base of the cos- tal margin is unusual, it is by no means unique; sclerotized areas, diversely located, have been found in other orthopteroids, such as Nacekomia rossae Richardson (also from the Francis Creek Shale). The venation of the wings in Herdina has many of the same fea- tures as those of the Cacurgidae, as noted above. The most distinctive characteristic is the irregularity of the main veins ; this is undoubtedly associated with the thickness of the cross veins, which are only slightly less heavy than the main veins. The relatively small size of the hind wing of Herdina is also not surprising in it- self; its most striking and significant feature is the similarity of its venation to that of the fore wing. In all of the Protorthoptera in which the hind wing is known, and in the Orthoptera as well, the venation of the hind wing is distinctly different from that of the fore wing; Rs arises much nearer the base of the wing; CuA is unbranched ; and the anal area is enlarged, with several additional anal veins. Herdina is the first instance known of an otherwise typ- ically protorthopterous insect in which the fore and hind wings are virtually homonomous in venational pattern. Unfortunately, nothing is known of the hind wings of those Protorthoptera which appear to 1971] Carpenter & Richardson — Pennsylvanian Insects 295 be closely related to Herdina , i.e. the Cacurgidae and Omaliidae. H ence, we do not know if the hind wing venation in these families was like that of other Protorthoptera or if it was like that of the fore wings, as in Herdina. Such a condition does occur, however, among several existing orders of Neoptera (Isoptera and Embiop- tera, for example). In these the homonomous condition is apparently derived from a previous heteronomous one; the same was almost cer- tainly true for the Herdinidae. References Carpenter, F. M. 1964. Studies on North American Carboniferous insects. 3. A spilap- terid from the vicinity of Mazon Creek, Illinois (Palaeodictyop- tera). Psyche 71: 117-124. 1967. Studies on North American Carboniferous insects. 5. Palaeodicty- optera and Megasecoptera from Illinois and Tennessee, with a discussion of the Order Sypharopteroidea. Psyche 74: 58-84. 1971. Fossil insects from New Mexico. Psyche 77: 400-412. Carpenter, F. M. and E. S. Richardson, Jr. 1968. Megasecopterous nymphs in Pennsylvanian concretions. Psyche 75: 295-309. Crampton, G. C. 1927. Eugereon and the ancestry of the Hemiptera, psocids, and Hymenoptera. Bull. Brook. Ent. Soc. 22: 1-17. Goldenberg, F. 1854. Die fossilen Insekten der Kohlenformation von Saarbriicken. Palaeontogr. 4: 17-38. Guthorl, P. 1934. Die Arthropoden aus dem Karbon und Perm des Saar-Nahe- Pfalz-Gebietes. Preuss. Geol. Landes., Abhandl. N.F. 164: 73- 76. Handlirsch, A. 1911. New Falaeozoic insects from the vicinity of Mazon Creek, Il- linois. Amer. Journ. Sci. (5) 31: 374. 1920. Palaeontologie. In Schroder, Handbuch der Entomologie. 3: 129. Kukalova, J. 1968. Permian mayfly nymphs. Psyche 75 : 310-327. 1969-1970. Revisional study of the Order Palaeodictyoptera in the Upper Carboniferous shales of Commentry, France. Psyche 76: 163-215 ; 439-486; 77: 1-44. Laurentiaux, D. 1952. Decouverte d’un rostre chez Stenod ctya lobata Brgt. (Paleodic- tyoptere stenodictvide) et le probleme des Protohemipteres. Bull. Soc. Geol. France, 2: 233-247. 1953. Classes des Insectes. In Traite de Paleontologie (Piveteau), Paris. 3 : 415. Lameere, A. 1933. Precis de Zoologie. 4: 255-256. SCHLECHTENDAL, D. V. 1912. Untersuchung iiber die karbonischen Insekten und Spinnen von Wettin unter Berucksichtigung verwandter Faunen. Abhandl. Kaiserlichen Leopold. -Carol. Deutsch. Acad. Natur. 98: 119-124. THE MALE GENITALIA OF BL ATT ARIA. VIII. PANCH LORA , ANCHOBLATTA, BIOLLEYA , PELLOBLATTA , AND AGHROBLATTA. (BLABERIDAE: PANCHLORINAE) . By Louis M. Roth Pioneering Research Laboratory U. S. Army Natick Laboratories Natick, Massachusetts 01760 McKittrick (1964) placed three blaberid genera, Panchlora Bur- meister, Achroblatta Saussure, and Capucina Saussure, in the Panch- lorinae. Princis (i960) listed Panchlora , Phortioecoides Rehn, and Proscratea Burmeister under the Panchloridae. My studies of male genitalia of Blaberidae have shown that Capucina and Phortioecoides are members of the Zetoborinae (Roth 1970a). The genitalia of Proscratea differ markedly from those of genera I believe should be placed in the Panchlorinae and I do not consider it in this paper. The male genitalia of species of Panchlora , Achroblatta , Ancho- blatta Shelford, Biolleya Saussure, and Pelloblatta Rehn are basically similar and I place these five genera in the Panchlorinae. Unfor- tunately the male genitalia of an undescribed species of Pelloblatta were lost in preparation, due principally to their marked reduction and light sclerotization. Princis (1965) placed Pelloblatta in the Oxyhaloidae but the male genitalia of members belonging to the Oxyhaloinae (Roth, 1971) differ markedly from those of the Panch- lorinae. Princis (i960) placed Achroblatta in the Laxtinae, a sub- family which he considered provisional (Roth, 1970a), and Biolleya in the Latindiidae. McKittrick (1964) placed Laxta in the Epilam- prinae, tribe Laxtini. The genus Latindia Stal is an oviparous poly- phagid genus whereas Biolleya is an ovoviviparous member of the Blaberidae. Princis (i960) placed Anchoblatta in the Brachycolinae with a ( ?) but included it under this subfamily in his 1963 Catalogus. I consider Princis’ Brachycolinae to be a tribe in the Blaberinae (Roth, 1970b). Anchoblatta signifera (Scudder) was originally placed in Panchlora , but Kirby assigned it to Achroblatta. Princis (1963) listed Anchoblatta signifera with a query under Achroblatta , but Gurney (personal communication) regards it to be an Ancho- blatta. Materials and Methods The genitalia were treated with 10% KOH and mounted in Per- mount. Considerable care must be taken with these specimens be- cause they are so lightly sclerotized and small that they can be readily 296 1971] Roth — Blattaria 297 Fig. 1. (978 L). Male genitalia (dorsal view) of Panchlora vosseleri Shelford. Usambara-Berg, Tanganyika (det. Princis). (Ll = first sclerite of left phallomere; L2d = dorsal sclerite of L2 ; L2vm = ventromedial sclerite; R2 = hooked sclerite of right phallomere. To conserve space the photographs of the phallomeres have been placed closer together than they were arranged on the slide. lost in preparation. Because of light pigmentation, Polaroid Type 51, high-contrast film usually was used to illustrate them (cf. Figs. 21-22). The source of each of the specimens illustrated is given using the following abbreviations: (BMNH) = British Museum (Natural History), London; (CUZM) = Copenhagen University, Zoological Museum, Denmark; (L) — Zoological Institute, Lund, Sweden; (N) = U. S. Army Natick Labs.; (USNM) = United States National Museum, Washington, D.C. Geographical collection data and the names of specialists who identified the specimens, if known, follow these abbreviations. The number preceding the abbreviations 298 Psyche [December refers to the number assigned the specimen and its corresponding genitalia (on a slide) which are deposited in their respective museums. Results and Discussion The most characteristic feature of the male genitalia of most Panchlorinae is the absence of the genital hook and light pigmenta- tion and marked reduction of the remaining 2 phallomeres when they are present. McKittrick (1964, p. 72) noted the marked reduction in the genitalia of Panchlora nivea and pointed out that the only sclerotized structures are the cleft region of Li and the lightly pig- mented median sclerite (L2vm). However, a study of several species of Panchlora shows that some forms have all three phallomeres (Figs. 1-4) ; males that lack certain phallomeres probably evolved from species that had them. Based upon the presence or absence of genital phallomeres, the Panchlora studied here can be divided into the following Species Groups: Group 1 ( Panchlora vosseleri) . Though lightly sclerotized, all three basic phallomeres are present, including an L2d (Figs. 1-4). Group 2 (Panchlora stanleyana) . L2d is absent; Li, L2vm, and R2 are present (Figs. 5-7). Group 3 (Panchlora nivea , P. thalassina) . L2d and R2 are absent; Li, and L2vm, are present (Figs. 8-15). Group 4 (Panchlora bidentula , P. minor , P. sagax, P. dumicola, P. peruana). L2d, R2, and apparently L2vm are absent; only Li is present (Figs. 16-20). Group 5 (Panchlora exoleta Burmeister). Apparently no sclero- tized structures are present. Groups 1 and 2 are African and 3 to 5 are found in Central and/ or South America. In Groups 4 and 5 it is quite possible that L2vm was so lightly sclerotized and Li so small, and poorly defined, that they were lost in preparation of the genitalia. The male phallomeres L2vm of Achroblatta (Figs. 24-25), Biolleya (Figs. 21-23), and Anchoblatta (Figs. 26-29) are very lightly pig- mented. Li is also reduced as in most Panchlora , and L2d and R2 are absent. The marked reduction in the male genital phallomeres of this sub- family may be related, in some way, to the mating behavior of these genera. In most species of cockroaches the receptive females respond to male courtship by palpating his dorsum, which in many genera possesses specialized glands which persumably produce a secretion at- 1971] Roth — Blattaria 299 Figs. 2-7. Male genitalia of Panchlora spp. 2-4. (978 L). P. vosseleri (same specimen as shown in Fig. 1). 5-7. P. stanleyana Rehn. 5-6. (977 L) Cameroon Republic (det. Princis). 7. (N). Ivory Coast (det. Roth), (scale = 0.1 mm). 300 Psyche [December Figs. 8-16. Male genitalia of Panchlora spp. 8-9. (N). P. nivea (det. Gurney). 10-11. (56 BMNH). P. thalassina Saussure and Zehntner. Villa Ana, F.C.S.F. Argentina (det. Hebard). 12-13. (173 USNM). P. sp. Palenque, Chiapas, Mexico. 14-15. (172 USNM). P. sp. Palenque, Chiapas, Mexico. (15 is a ventral view). 16. (57 BMNH). P. bidentula Hebard. Mosqueiro, Rio de Para (det. Hebard). (scale = 0.1 mm). 1971] Roth — Blattaria 301 Figs. 17-25. Male genitalia of Panchlorinae. 17. (976 L). Panchlora peruana Saussure. Bolivar, Batatal, Colombia (det. Princis). 18. (57 CUZM). Panchlora minor Saussure and Zehntner. Brazil (det. Princis). 19. (973 L). Panchlora sagax Rehn and Hebard. St. Thomas, Lesser An- tilles (det. Princis). 20. (975 L). Panchlora dumicola Albuquerque and Gurney. Brazil (det. Princis). 21-23. (174 USNM). Biolleya alaris Saus- sure. Paratype. La Palma, Costa Rica. (Figs. 21 and 22 are the same L2vm taken with Polaroid Type 51 and 52 film respectively). 24-25. Achro- blatta luteola (Blanchard). Darien Province, Sante Fe, Panama (det. Fisk), (scale — 0.1 mm). 302 Psyche [December Figs. 26-29. Male genitalia of Anchoblatta. 26-27. (170 USNM). Signi- fera (Scudder). San Martin, Peru (det. Gurney). 28-29. (53 USNM). Rio Ucuyali, Pucallpa, Peru (det. Gurney). (L2vm was accidentally broken [arrows indicate the two parts of the sclerite]). (scale = 0.1 mm). 1971] Roth — Blattaria 303 tractive to the female (Roth, 1969). When the female is in the proper position above the male he extends the genital hook (R2) and seems to use this structure to pull down the female’s subgenital plate so that he can insert his genitalia and grasp her genitalia. Ac- cording to Khalifa (1958) in Blattella germanica (L.), when the female is palpating the male’s tergal glands, the male fully extends the hooked phallomere, which is on the left side in this genus (not the right as in all Blaberidae) directs it upwards, and inserts it into the female’s genital chamber ; there it clasps a sclerite situated in front of the ovipositor. Once a secure hold is obtained, the pair assumes an end-to-end position, with their heads facing in opposite directions, and the hook “. . . acquires a hold on the ovipositor.” In Periplaneta americctna (L.) the male . . prior to getting an actual hold on the female pulls down the gynovalvular portion of the seventh ster- nite of the female by the tip of its protruded titillator.” and then inserts its genitalia into the female’s genital pouch. (Gupta, 1947). Zabinski (1933) showed that the titillator, in Blatta orientalis , is used to seize the female in the initial phase of copulation. The male apparently cannot mate if the titillator (genital hook or R2 in Blaberidae) is surgically removed (Zabinski, 1933, Roth and Willis, 1952). The mating behavior of Panchlora differs markedly from the above species. In Panchlora nivea (Roth and Willis, 1958) and P. irro- rata Hebard (Willis, 1966) the female does not assume a position above the male prior to mating, but the males simply back into the female. Possibly stridulation plays a role in mating behavior of these species (Roth and Hartman, 1967). The difference in precopulatory positions may have had some role in the marked reduction of male genitalic structures, especially the loss of the genital hook. Nothing is known of the precopulatory positions of the genera of Achroblatta, Anchoblatta, and Biolleya , and it would be of interest to see if the mating behavior of these genera is similar to that of Panchlora. It should be pointed out that in Groinphadorkma portentosa (Schaum) (Oxyhaloinae) the male also backs into the female to assume the copulatory position (Barth, 1968) but the males of this genus have a well-developed, but relatively short, genital hook (Roth, 1971). Summary Five genera, Panchlora , Anchoblatta , Biolleya , Pelloblatta, and A chroblatta are included in the blaberid subfamily Panchlorinae. Cer- tain male genitalic phallomeres are usually reduced or absent and structures that are present are very lightly sclerotized. One African 304 Psyche [December species of Panchlora has all the phallomeres characteristic of most male blaberid genitalia. Five groups are erected for species of Panchlora , based on the number of phallomeres which are missing. The genital hooks (R2) are absent in Panchlora (from Central and/ or South America), Achroblatta , Biolleya, and Anchoblatta. The mating behavior of Panchlora differs from most other Blat- taria, and it is discussed in relation to the loss of the genital hook. Acknowledgments I thank the following for the loan of museum specimens: Dr. David Ragge (British Museum), Dr. Ashley Gurney (U. S. Na- tional Museum), Dr. Karl Princis (Lund), Dr. S. L. Tuxen (Copenhagen), and Dr. Frank Fisk. I am grateful to Mr. Samuel Cohen for taking the photographs. References Barth, R. Jr. 1968. The mating behavior of Gromphadorhina portentosa (Schaum) (Blattaria, Blaberoidea, Blaberidae, Oxyhaloinae) : an anomalous pattern for a cockroach. Psyche 75: 124-131. Gupta, P. D. 1947. On copulation and insemination in the cockroach Periplaneta americana (Linn.). Proc. Nat. Inst. Sci. India 13: 65-71. Khalifa, A. 1950. Spermatophore production in Blattella germanica L. Orthoptera: Blattidae). Proc. Roy. Entomol. Soc. London 25(A): 53-61. McKittrick, F. A. 1964. Evolutionary studies of cockroaches. Cornell Univ. Agr. Exp. Sta. Mem. No. 389, 197 pp. Princis, K. 1960. Zur systematik der Blattarien. Eos 36: 427-449. 1963. Orthopterorum Catalogus. Subordo Polyphagoidea : Fam. Homoe- ogamiidae, Euthyrrhaphidae, Latindiidae, Anacompsidae, Atti- colidae, Attaphilidae. Subordo Blaberoidea: Fam. Blaberidae. s’-Gravenhage, pp. 76-172. Roth, L. M. 1969. The evolution of male tergal glands in the Blattaria. Ann. En- tomol. Soc. Amer. 62: 176-208. 1970a. The male genitalia of Blattaria. III. Blaberidae: Zetoborinae. Psyche 77: 217-236. 1970b. The male genitalia of Blattaria. IV. Blaberidae: Blaberinae. Psyche 77: 308-342. 1971. The male genitalia of Blattaria. VI. Blaberidae: Oxyhaloinae. Psyche 78: 84-106. Roth, L. M. and H. B. Hartman. 1967. Sound production and its evolutionary significance in the Blat- taria. Ann. Entomol. Soc. Amer. 60: 740-752. 1971] Roth — Blattaria 305 Roth, L. M. and E. R. Willis. 1952. A study of cockroach behavior. Amer. Midland Nat. 47: 66-129. 1958. The biology of Panchlora nivea with observations on the eggs of other Blattaria. Trans. Amer. Entomol. Soc. 83. 195-207. Willis, E. R. 1966. Biology and behavior of Panchlora irrorata , a cockroach ad- ventive on bananas. Ann. Entomol. Soc. Amer. 59: 514-516. Zabinski, J. 1933. Fonctionnement des differentes parties des appareils copulateurs chitines males et femelles de la Blatte ( Periplaneta orientalis L.). Soc. de Biol, de Varsorvie 112: 598-602. THE STRUCTURE OF DUN BARI A ( PALAEODICTYOPTERA ) * By Jarmila Kukalova— Peck Department of Geology, Carleton University, Ottawa, Ontario, Canada The extinct order Palaeodictyoptera. is the most abundant paleop- terous group of insects found in Paleozoic deposits. During its geological range, from the Upper Carboniferous (Namurian) to the Permian (Leonard), the order radiated into extremely diverse lines with adaptations to many environments and with various specializa- tions. In contrast to this speciation, fossils are relatively scarce in spite of their wide distribution. Furthermore, the aerial way of life virtually eliminates the finding of specimens deposited in assemblages of their original and natural communities. These circumstances have greatly reduced the chances of collecting many specimens belonging to one species, with the result that there are almost no data available for intraspecific variations in size, wing venation, and especially sec- ondary sexual characters. The single exception to this is the case of Dunbaria fasciipennis Tillyard ( Spilapteridae) from Permian (Leonard) deposits of Elmo, Kansas. Tillyard based his original description of this species (1924) on three specimens; later (1925) he discussed another three specimens and dealt with the wing venation (in the light of Lameere’s vena- tional concepts) and with the terminal abdominal appendages. All six specimens are in the collection of the Peabody Museum, Yale University, and are numbered as follows: holotype 1001 ab, allo- type 1002 ab, paratype 1050, and specimens 5020, 5021 and 5022. 1 During my stay at Harvard University in 1968, Professor Car- penter encouraged me to review all of the specimens of fasciipennis and he placed at my disposal three more specimens from his collection in the Museum of Comparative Zoology, Harvard University, num- bered 3056 ab, 3057 ab, and 3058. This series of nine specimens is *This research has been aided by grant number GB-7308 and number GB-27333 from the National Science Foundation (F. M. Carpenter, principal investigator, Harvard University). Specimen number 5020 in the Peabody Museum was designated by Till- yard in 1925 (page 335) as a paratype of fasciipennis. However, since this particular specimen was not mentioned or even seen by Tillyard at the time of his original description of the species (1924), it has no status as a paratype. 306 1971] Kukalovd-Peck — - Dunbaria 307 the most extensive aggregation of Palaeodictyoptera belonging to one species so far known. To avoid confusion, I will first of all discuss several misinterpreta- tions of morphological features included in Tillyard’s two papers. With respect to the wings , Tillyard (1924, P- 205 ; 1925, p. 329) mentions the presence of a very delicate archedictyon extending over the wings. I have been unable to see any traces of this in the fossils. The wing membrane is very thin and is delicately wrinkled in sev- eral of the specimens, probably as a result of the process of preserva- tion; I suspect that this wrinkling was interpreted by Tillyard as the archedictyon. A series of strong macrotrichia was described by Tillyard as being present on R, Ri, and basal part of Rs, but I find that these macrotrichia are present on all veins, not just these few. With respect to the thorax , Tillyard (1924, p. 203, 205) describes the prothorax as being very short and as lacking prothoracic, lateral lobes. Actually, as shown especially well by specimen 3057 MCZ, the prothorax is of normal length for the Palaeodictyoptera and it possesses a pair of sclerotized, prothoracic lobes. In specimen 1002 of the Peabody Museum, which was studied by Tillyard, there is a small fragment of a wrinkled prothoracic lobe, but this was inter- preted as the head (Tillyard, 1924, p. 205) and the remnant of the prothorax was considered its full length. So far, no representatives of Palaeodictyoptera have been found without prothoracic lobes. In highly modified species, the lobes may be reduced in size and sclero- tized, forming scale-like structures, as in Homalonenra lehmani Kuka- lova (1969, p. 182, fig. 8) belonging to the family Spilapteridae. With respect to the abdomen , Tillyard (1924, p. 205) mentioned ten visible segments but actually the eleventh vestigial segment is also present. Subsequently (1924, p. 207), Tillyard implied that males in the Palaeodictyoptera might not have the long cerci; how- ever, such cerci appear to be present in both sexes. The holotype (Peabody Museum 1001) was considered by Tillyard to be a male, because of having a narrower abdomen than the “female” (no. 1002) (1924, p. 207). However, the lateral edges of all abdominal tergites in the holotype are broken off so that their widths are not measur- able. Since the genital structures are not preserved in the fossil, the holotype does not provide any information about its sex. Specimen 1002, with the complete abdomen and terminal appendages, was de- scribed by Tillyard as a female (1924, p. 207; 1925, p. 334, fig. 3). Tillyard’s conclusion was based on a misinterpretation of the male claspers, which he thought were the ovipositor; this is discussed more 308 Psyche [December Fig. 1. Dunbaria fasciipennis Tillyard, 1924. Lower Permian of Elmo, Kansas; fore wing length 17 mm, width 5.8 mm; hind wing length 17.1 mm, width 7.6 mm; specimen 1002, male. Peabody Museum, Yale Uni- versity. 1971] Kukalova-Peck — Dunbaria 309 Fig. 2-A. Dunbaria fasciipennis Tillyard, 1924, detail of anterior margin in proximal half of the wing; S — serrated precostal strip; C — broadened band-like costa with serrated anterior ridge. 2-B. Specimen 1002, enlarged end of the abdomen; K — male claspers; P — slender appendages, with striations, probably parameres. 2-C. The same specimen 1002, enlarged end of the abdomen, after Tillyard (1925, fig. 3); ad-appendix dorsalis; c — cercus; gcx — gonocoxite; st — style; IX, X — abdominal segments. Peabody Museum, Yale University. 3io Psyche [December Fig. 4. Specimen 1050. Left fore wing; length 16.3 mm, width 5.2 mm. Peabody Museum, Yale University. Fig. 5. Specimen 1001. Holotype. Left fore wing; length 17.2 mm, width 5.5 mm. Peabody Museum, Yale University. Fig. 6. Specimen 1001. Right fore wing; length 17.2 mm, width 5.5 mm. Peabody Museum, Y’ale University. 1971] Kukalova-Peck Dunbaria 3 ii Fig. 7. Specimen 3057. Right fore wing; length 17.8 mm, width 5.5 mm. Museum of Comparative Zoology, Harvard University. Fig. 8. Specimen 5020. Right fore wing; length 18.1 mm, width 5.4 mm. Peabody Museum, Yale University. Fig. 9. Specimen 3056. Right fore wing; length 17.3 mm, width 5.3 mm. Museum of Comparative Zoology, Harvard University. 312 Psyche [December fully below. Consequently, the nine specimens of Dunharia fasciipen- nis do not provide conclusive information about sexual dimorphism. Mention should also be made of Tillyard’s reference to the pres- ence of a “short appendix dorsalis similar to those of some Recent Plectoptera” (1925, p. 335, fig. 3). This structure turns out to be a narrow piece of matrix, accidentally included between the cerci of specimen no. 1002. New Observations on the Structure of Dunbaria Wings: In Palaeodictyoptera with a very delicate wing mem- brane, additional supporting structures often occur, these being con- centrated in the basal third of the wing and along the anterior margin, much as in the wings of the Odonata (Kukalova, 1969-1970, Pt. I- III). In the wings of Dunbaria , of which the variation will be discussed later, there are several elements that strengthen the mem- brane : a series of strong cross veins connect the anal stem to the origin of Rs, supporting the basal third of the wings; the anal stem is unusually wide, flattened and heavily sclerotized ; the proximal part of Ai is provided with a conspicuous, convex cuticular thicken- ing, apparently also sclerotized ; the postcostal area is not flat as in most other Palaeodictyoptera, but forms a triangular, concave fold with 1-3 twigs; the precostal strip in both fore and hind wings is serrated, recalling the condition in the Odonata (Tillyard, 1924, p. 206) (see fig. 2 A-C) ; the costa is very strong, broadened and band-like in appearance to beyond the middle of the wings, and is provided also by a serrated anterior margin (Fig. 2 A-C). The en- tire apical and posterior margins are dentate. The prothoracic paranota are heavily sclerotized, cordate, and cov- ered by dense, short hairs; they do not overlap the fore-wings. Radiating veins are only vaguely indicated in the lobes; the margin is bordered by a thickened ridge. The sclerotization of the paranota is known to be correlated with the reduction of veins in the spilap- terid Homaloneura , which is closely related to Dunbaria. The presence of the thickened ridge has already been observed in Eublep- tus (Carpenter, 1965, p. 181, fig. 3) of the closely related family Eubleptidae. Body Structures: The head is small, eyes prominent; antennae thin, multisegmented, composed of long segments which are slightly broadened distally (fig. 15); thoracic segments almost equal; meso- thorax slightly broader than the prothorax and narrower than the metathorax; abdomen relatively slender, composed of 11 visible seg- 1971] Kukalovd-Peck — Dunbaria 313 ments, the first abdominal segment shorter than the following ones; the eleventh segment vestigial, divided by a deep median incision; cerci originating from the eleventh segment, markedly robust and annulated. The external genitalia, which are well-preserved in specimen 1002, are of special interest. At first sight, they are reminiscent of the valves of an ovipositor and they were interpreted as such by B ill- yard (1925, p. 334, fig. 3). This appearance, however, is mostly due to preservation, which must be discussed next. The abdomen is dorsoventrally flattened in the fossil, but the very end is twisted so that the terminal segments show a lateral view in part. The sternites and the genital appendages are slightly super- imposed on the 10th and nth tergites. As preserved, the abdomen could be considered comparable to a cylindrical body, flattened by pressure which is directed obliquely to the dorsoventral level, while the under surface of the cylindrical body splits. As a consequence, the genital structures appear more or less in ventral view (fig. 2B). The genital structures (fig. 2B-K) are interpreted here as male claspers. They originate at the posterior margin of the 9th seg- ment, are paired and diverge slightly distally and extend beyond the length of the body. They are not sclerotized and their surface is covered with scattered stiff, short hairs that are irregularly distrib- uted and directed posteriorly. Beneath the claspers there is another pair of slender processes, terminating in two thin, posteriorly curved, sclerotized projections; these bear a dense covering of transverse striae. The whole structure resembles the parameres (Fig. 2 B-P) of insects by their position and morphology. Several features suggest that the processes described above are male structures and not valves of an ovipositor as Tillyard assumed: their general morphology, characteristic attachment to the 9th ster- nite, their unsclerotized nature and the presence of posteriorly di- rected stiff and solitaiy hairs. These features are contrary to what is known about the ovipositor in Palaeodictyoptera and about ovi- positors in general. The female genitalia are known in the family Spilapteridae, to which Dunbaria belongs, in the closely related spe- cies Homaloneura ornata Brongniart (Kukalova, 1969, p. 179, fig. 7) ; the ovipositor does not differ from that of other Palaeodictyop- tera, Megasecoptera and Diaphanopterodea. The valves in all of these related orders are curved, heavily sclerotized and resemble in shape and their broad attachment the valves of some dragonflies (Kukalova, 1969, p. 449, fig. 32). If they are provided with hairs, 314 Psyche [December the latter are always directed anteriorly. As far as I know, there is no evidence in any insect ovipositor of hairs directed posteriorly, which would presumably prevent the valves from penetrating the substrate. The male claspers of Palaeodictyoptera have previously been known in only one specimen belonging to Stenodictya spinosa (Brongniart) (Kukalova, 1970, Pt. Ill, p. 11, fig. 53, p. 13, 54)* Their structure resembles that of the claspers of Permian Megasecoptera (Carpenter 1939, p. 31, fig. 2 A) and those of Permian and some Recent Ephemeroptera. They are relatively more primitive, however, composed of small, equal segments, not unlike cerci, and directed towards each other beyond the 9th segment instead of at about the middle. In the light of this evidence, the claspers of Dunbaria may be regarded as very advanced in structure, actually approaching in general respects those of some dragonflies. This circumstance is of great interest, since besides the significant similarity of the female ovipositor to that of dragonflies, Dunbaria provides the first evidence of the similarity of the male genital structures as well. The conver- gence of wing structures to dragonflies repeatedly occurs in different evolutionary lines of Palaeodictyoptera (Calvertiellidae, Eugereonidae, Archaemegaptilidae, etc.). In spite of the closer relationship which is indicated between the Palaeodictyoptera and Ephemeroptera, all of the evidence, including that of the genitalia, supports the assumption of single ancestral stock of all palaeopterous orders. Variability of Wings of Dunbaria In current studies on palaeodictyopterous wing venation, the sec- ondary branching of veins is generally considered to be not a specific feature but rather one of individual variability. Much emphasis is given by some authors to the level of first branching of main veins, especially R and M, and to the widths of wing areas. Little informa- tion is known of the individual variability of wing shape and size. The following analysis of wing morphology of nine specimens of Dunbaria fasciipennis provides some data on this subject. The fore wing length ranges from 16.3 mm to 18.1 mm and the width from 5.2 mm to 5.8 mm. The hind wing length varies from 17 mm to 17.9 mm and width from 6.6 mm to 7.6 mm. A nterior Margin : It is concavely curved in more or less all wings ; this feature in Palaeodictyoptera is often affected by preservation and needs to be considered very carefully. Costa: The bandlike broaden- ing of C tends to be more pronounced and longer in the larger wings. Sc: The length of this vein is variable within the anterior part of 1971] Kukalova-Peck — Dunbaria 315 Fig. 10. Specimen 1001. Holotype. Right hind wing; length 17.2 mm, width 7.6 mm. Peabody Museum, Yale University. Fig. 11. Specimen 1001. Holotype. Left hind wing; length 17.2 mm, wddth 7.6 mm. Peabody Museum, Yale University. Fig. 12. Specimen 1002. Right hind wing; length 17 mm, width 7.6 mm. Peabody Museum, Yale University. Fig. 13. Specimen 5021. Right hind wing; length 17.2 mm, width 7.2 mm. Peabody Museum, Yale University. Fig. 14. Specimen 3058. Right hind wing; length 17.9 mm, width 6.6 mm, Museum of Comparative Zoology, Harvard University. 3 1 6 Psyche [December the apical margin; additional terminal branches leading to Rs may occur (specimen 1001, fore and hind wings). Rs: The level of origin of this varies; in extreme cases Rs diverges almost at the very base (specimen 1001, right hind wing) ; the number of branches of Rs branches varies in fore wing from 7 to 10, in the hind wing from 7 to 9. Branches of Rs are simple with a single exception (specimen 1002, 4th branch forked in hind wing). M : It is much alike in all specimens with one exception (1001, left hind wing has an additional twig on MP). Cu: CuP is always simple, CuA with 3-4 simple branches in fore wing and with 4-5 simple branches in the hind wing (specimen 1001 has additional twig in the hind wing). An: Anal veins vary from 6 to 9 in the fore wing and from 7 to 8 in the hind wing. Cross veins: The number is very variable; this feature in the Palaeodictyoptera is strongly affected by preservation. Supporting, strong cross veins near the base are highly variable in number and arrangement. Other supporting sclerotized structures at the base (broadened anal stem, cuticular thickening on Ai) are different in size but always present. Wing areas: Postcostal area is highly vari- able in size and shape, with 1-3 twigs. All other areas vary in size and width (e.g., sc-r area). Anal area varies both in length and width, especially in the hind wings. Shape of the wings: This is fairly constant (not considering the secondary deformations of the anterior margin) except for the narrow basal third in the fore wing of the male 1002 and specimen 1050. The posterior margin of the hind wing retains the characteristic undulation. Color pattern: This varies as shown in figures 3-15. In conclusion, we can say that all of the morphological features in the wings of Dunbaria f asciipennis are variable to some extent, in- cluding the level of origin of Rs and the size of the anal area, which may be considered as providing a specific character but not a narrowly defined one. Supporting structures, such as strong cross veins or occa- sional twigs of main veins, are variable, definitely on an individual level. The outline of the wings presents a rather reliable specific character but the effect of preservation should be thoroughly consid- ered. Within the 9 specimens of Dunbaria f asciipennis, two fore wings are markedly narrow in the basal third (specimen 1002 and 1050). Since specimen 1002 is a male, there is a possibility that this feature is a character of sexual dimorphism. This cannot be resolved until more specimens are found with well-preserved body parts combined with the wings. 1971] Kukalova-Peck — Dunbaria 317 3 1 8 Psyche [December Relationships of Dunbaria. Tillyard (1924, p. 205) correctly re- lated Dunbaria with another spilapterid genus Homaloneura, occur- ring in the Upper Carboniferous of Europe and North America (Carpenter, 1964; Kukalova, 1969). Of the species known so far, Homaloneura ornata Brongniart, from the Commentry Shales in France, seems to be the closest. Dunbaria differs in possessing a very long Rs, with more numerous branches and also in possessing a prom- inent odonate-like serration. References Carpenter, F. M. 1939. The Lower Permian Insects of Kansas. Part 8. Additional Megasecoptera, Protodonata, Odonata, Homoptera, Psocoptera, Protelytroptera, and Protoperlaria. Proc. American Acad. Arts Sci. 73 (3) : 29-70. 1964. Studies on North American Carboniferous Insects. 3. A Spilapte- rid from the Vicinity of Mazon Creek, Illinois (Palaeodictyop- tera). Psyche, 71(3): 117-124. 1965. Studies on North American Carboniferous Insects. 4. The Genera Metropator, Eubleptus, Hapaloptera and Hadentomum. Psyche, 72(2) : 175-190. Kukalova, J. 1969. Revisional Study of the Order Palaeodictyoptera in the Upper Carboniferous Shales of Commentry, France, Part I. Psyche, 76(2) : 163-215. 1969. Ibid., Part II. Psyche, 76(4): 439-486. 1970. Ibid., Part III. Psyche, 77(1): 1-44. Lameere, A. 1922. Sur la nervation alaire des insectes. Bull. Classe Sci. Belgique 1922(4) : 138-149. Tillyard, R. J. 1924. Kansas Permian Insects. Part I. The Geologic Occurrence and the Environment of the Insects with Description of a New Palae- odictyopterid. Amer. Journ. Sci., (5) 7: 203-208. 1925. Kansas Permian Insects. Part 4. The Order Palaeodictyoptera. Amer. Journ. Sci., 9 (52): 328-335. THE MATING BEHAVIOR OF PARCOBLATTA FULVESCENS (SAUSSURE AND ZEHNTNER) ( BLATTARIA, BLABEROIDEA, BLATTELLIDAE, BLATTELLINAE ) 1 By Peter Wendelken and R. H. Barth2 Department of Zoology The University of Texas at Austin This communication is the sixth in a series of largely descriptive papers dealing with the mating behavior of cockroaches (see Barth, 1961, 1964, 1968a & b, 1970; Roth and Barth, 1967). The aim of this series is twofold : first to provide background information for experimental studies, and second to provide the detailed comparative information necessary for a study of the evolution of mating be- havior within the Blattaria. A more general introduction to the series may be found in Barth (1964). The mating behavior of the Fulvous wood cockroach, Parcoblatta fulvescens (Saussure and Zehntner), forms the subject of this communication. Materials and Methods Stock cultures of P. fulvescens were maintained as described by Barth (1964) for Byrsotria fumigata. The observations on mating behavior were made in the evening (the normal activity period for these animals) under red illumination in specially designed observa- tion chambers constructed of wood (13" X 9" X 5" deep) with a removable partition dividing the chamber into two equal parts (for details, see Barth, 1964). In each observation 2 to 3 males and 2 to 3 females were employed. The ethological terms employed in the description have been previously defined by Barth (1964). Results and Discussion Parcoblatta fulvescens is a small (11 to 17 mm in length) cock- roach generally found in wooded areas under leaf litter and other debris and is widely distributed in eastern, southern, and central areas of the United States. It shows marked sexual dimorphism. The females are robust and wingless with reduced tegmina that extend over the first abdominal segment. They are orange-brown *No. 6 in a series of papers entitled “The Mating Behavior of Cock- roaches.” Supported by N.S.F. Research Grant GB-4614 and NIH Training Grant 2T01-GM-008 37-07. 319 320 Psyche [December dorsally with a darker abdomen. The males are more slender and winged, and their tegmina extend beyond the abdominal tip. They are light brown in dorsal coloration. There seem to be no previous accounts of the mating behavior of this species in the literature although a brief account of the court- ship behavior of Parcoblatta virginica (Brunner) is given by Roth and Willis (1958). According to their description, males of P. virginica raise their wings after contacting the female with their antennae. The female, attracted by the secretion of the male tergal gland, mounts and feeds until she reaches the first abdominal tergite of the male at which time genital connection is achieved ; this is followed by assumption of the opposed position. Description of Normal Mating Behavior The following description is based on observations of 5 sequences resulting in successful copulation and numerous unsuccessful copu- lation attempts. 1. Behavior of males triggered by olfactory reception of female sex pheromone Upon olfactory reception of the volatile female sex pheromone, males exhibit sexual arousal by assuming an alert posture and increasing the rate of antennal waving. Oriented locomotion to the pheromone source ensues. Without reference to contact with females the behavior of sexually aroused males is characterized by rapid locomotion, frequent flying, and wing raising. The manner in which the male flies is variable. Some flights cover more distance than others. For example, a male may fly across the mating chamber or upward to an inverted landing on the underside of the lucite covers atop the chamber. Other flights are more cir- cumscribed, the male flying several inches upward and then return- ing to the substratum. During some of these flights, the male pivots to face in the opposite direction and then lands; this often results in the male landing on his back. In addition to rapid running and flying, sexually excited males show a great deal of wing raising of a quite variable nature. When the wings are raised, the angle formed by the wings and the abdomen varies between 20 and 80 degrees. The wings may be raised and lowered quite rapidly in what is essentially a pumping motion. This cycle of wing raising and lowering may be repeated a number of times in quick succession. On the other hand, the wings may remain in the elevated position for a brief period. And 1971] W endelken & Barth — Parcohlatta 3 21 in some cases a male will run about with his wings continuously raised. All of these variations in wing raising are frequently per- formed while the male is engaged in forward locomotion. The occurrence of wing raising while the male locomotes forward has also been observed in Periplaneta americana (Barth, 1970; Simon and Barth, in prep.). As the wings are raised, they are frequently spread laterally. Wing fluttering usually accompanies lateral spreading. The wings are fluttered at the high point of the wing raise and during the flutter the tegmina are spread laterally from the sagittal plane to an angle of 10 to 50 degrees (but usually about 45 degrees) and their lateral edges are directed forward. The wings are slightly less spread laterally and are not elevated as much vertically. For instance, if the tegmina are raised to 80 degrees during a flutter, the wings are only raised to about 60 degrees. One observation may be cited which underlines the amount of variation possible with regard to wing raising. In this case a male ran around very excitedly with his wings continuously elevated to about 20 degrees and then periodically raised them completely, very rapidly, with fluttering at the point of maximum elevation. During the wing raising displays, the abdomen is flexed so that the dorsal surface is convex and the tip contacts the substratum. 2. Male displays in the vicinity of the female The majority of male-female contacts are very brief. Unreceptive females most frequently decamp rapidly immediately after coming into contact with a male. When a sexually aroused male makes contact with a female, he immediately raises his wings, turns away from the female, and backs. The wing raising display, turning, and backing are all released by the initial momentary antennal contact; no further contact with the female is required. The elevation of the wings in the male’s display varies from display to display and is any- where from 15 to 80 degrees. The amount of turning varies between 90 and 180 degrees. The male’s abdomen is arched so that its dorsal surface is convex and the tip touches the substratum. The male’s backing movement may be oriented toward the female from any direction. Two cases were observed, one of which led to a successful copulation, in which the female contacted the male from behind (without contacting his antennae) resulting in the male wing raising and backing but without any turning. It seems that in these in- 322 Psyche [December MATING BEHAVIOR of PARCOBLATTA FULVESCENS ALTERNATIVE PATHWAY 9 MOTIONLESS Olfactory Reception of c? Pheromone 9 ORIENTED LOCOMOTION O IN CLOSE PROXIMITY of cf (no contact) DOMINANT PATHWAY cf MOTIONLESS Olfactory Reception of 9 Pheromone cf ALERT POSTURE and ANTENNAL WAVING cf ORIENTED LOCOMOTION cf Olfactory Reception of 9 Pheromone ? TOUCHES a" with ANTENNAE ' CLOSE PROXIMITY of 9 (no contact) 1 cT TOUCHES 9 with ANTENNAE Tactile Stim. Contact Chemoreception Tactile stim. cf FULL WING RAISING *• 1 DISPLAY with Contact Chemoreception TURNING Cf COPULATORY THRUSTS Figure 1. A summary of the mating behavior of Parcoblatta fulvescens indicating the possible releasers for each step in the sequence. For explana- tion of alternative pathways, see text. 1971] JVendelken & Barth — Parcoblatta 323 stances olfactory reception of the female sex pheromone combined with tactile stimulation of the male’s hindparts was sufficient to release the wing raising display and that tactile stimulation of the male’s hindparts inhibited turning. There were twelve observations in which a male on coming close to a female, but without contacting her, wing raised, turned, and backed. This represents approximately 20 percent of all observa- tions for which the events preceding display were recorded. In these cases the wing raising display, turning, and backing were all apparently released merely by olfactory reception of an intense con- centration of volatile female sex pheromone in the absence of any contact chemoreception or tactile stimulation. One of these twelve displays led to a successful copulation. If the female does not respond to the male’s display after a brief period, the male frequently will flutter his raised wings or pump and flutter them. This possibly serves to disseminate the male sex pheromone to a female who is not responding to the male’s display. Occasionally, after leaving the site of an unsuccessful copulation attempt, a male will locomote around with his wings still partially raised (10 to 30 degrees) for 30 seconds to one minute. 3. Terminal events in the copulation sequence A receptive female responds to the male’s display with active mounting and feeding, moving in a forward direction over the male’s exposed abdominal tergites. If the male’s backing is poorly oriented, the female adjusts her position accordingly. When the female is about two-thirds forward over the male’s abdomen, the male begins probing extensions with his abdomen which is now concave on the dorsal surface, the abdominal tip contacting the female’s under- surface. The female advances with her feeding activities to the region of the first tergite at which point genital connection is achieved. The female then performs a turning movement which results in the animals facing away from each other in the 180 degree opposed position which is maintained for the duration of copulation. In this position the male’s wings slightly overlie the abdominal tip of the female, covering her cerci. 4. Behavior of copulating pairs Five accurately timed copulations lasted 54.5, 55, 58, 59, and 67 minutes. The duration of a sixth copulation was less than 53 minutes. Copulating pairs were generally quiescent, showing little antennal activity for most of the copulation period unless disturbed 324 Psyche [December by other animals. The female is entirely responsible for the pair’s locomotion which can be quite rapid. In three copulations, an area of moisture was noticed on the paper towel liner beneath the male’s head. It appeared that this was due to the male extruding water or some other fluid from his mouth. In one instance, a sudden surge of this moisture on the paper coin- cided with a movement of the male’s head. These wet spots ap- peared within the first few minutes of copulation and were visible for about 4 minutes. The females of copulating pairs assume an arched posture in which the body is held rather high above the substratum and flexed sharply ventrally. The posterior part of a copulating female’s abdomen curves downward to where it joins the male’s abdominal tip which is very close to the substratum. This arched posture may be pronounced enough to cause the male’s wings (which overlie the female’s abdomen) to be raised somewhat. This posture may be observed throughout the copulation, but it varies in extent; periods of very marked arching alternate with periods during which the arching is much less noticeable. In two copulating pairs, rhythmical movements were observed for which the female appeared responsible. This entailed a pivoting of the female’s body about a transverse axis such that her abdominal tip moved upward, pulling the male’s abdominal tip upward with it. In both pairs these movements were observed toward the end of copulation and occurred in a series which ceased and then was later resumed. The Role of Various Releasers in the Courtship Sequence A diagram illustrating the various avenues courtship behavior may take is presented in Figure 1. Possible releasers of various events in the courtship sequence are indicated. 1. Release of the male’s behavior Olfactory reception of the volatile female sex pheromone, the primary releaser of the male’s courtship behavior, is sufficient to release all of the male’s courtship behavior up to and including backing. In addition to the mating chamber observations, tests using filter papers removed from the female side of the chamber were conducted prior to the observation period. The males responded with vigorous antennal waving, oriented locomotion, flying and wing raising with fluttering. The rapid locomotion became random after a short while. Tactile stimulation from other males was not in- 1971] W endelken & Barth — Parcoblatta 325 volved in eliciting these responses. The males of this species are small and no more than three were ever employed ; in the mating chamber, contact between males was infrequent in this situation. Probably because of the low density of males, neither homosexual or pseudofemale behavior was observed in the mating chambers al- though one instance of a male mounting a displaying male was observed in the more crowded breeding culture. That males have a considerable ability to orient to a pheromone source was shown in several instances during the observation periods when a male precisely followed the “trail” of a female that had previously decamped. Turning and backing were not released during the filter paper tests nor in the behavior observations except when a male was very close to a female. Turning and backing in the majority of these cases were released by contact chemoreception and/or tactile stimuli when the male’s antennae contacted a female. However, in a signifi- cant number of cases the release of these activities was triggered solely by the apparently intense concentration of sex pheromone immediately surrounding the female. Tactile stimuli are necessary for the release of copulatory thrusts and phallomere extension. 'Copulatory thrusts begin when the mount- ing female’s mouthparts have progressed about two-thirds of the way forward over the male’s abdominal tergites. Whether phallomere extension occurs at this point or not until the female reaches the region of the first tergite is uncertain. Predominance of the female sex pheromone in the release of male courtship behavior has also been reported for the distantly related species, Periplaneta americana (Blattinae) (Barth, 1970; Simon and Barth, in prep.). As mentioned above, both species show wing raising in the absence of tactile stimuli and during forward loco- motion. However, turning and backing (in addition to the full wing raising display) only rarely occur in the absence of tactile stimulation in P. americana (Barth, 1970) but are not infrequently observed in P. fulvescens. In this respect, the female sex pheromone plays a more prominent role in courtship behavior in P. fulvescens than in P. americana. Backing in P. americana (as well as in four other species of Periplaneta and also Blatta orientalist often occurs without any tactile stimuli in addition to those which release wing raising with turning (Simon and Barth, in prep.). This is also true of P. fulvescens in those cases in which tactile stimuli release the wing raising display. P. americana males differ from 32 6 Psyche [December P. fulvescens in the exhibition of phallomere extension without the stimuli derived from female mounting and feeding; contact of the abdominal tip with the female suffices to release this response. The release of flying by males in sexual situations has previously been observed in Epilampra azteca and Epilampra columbiana. In E. columbiana , females as well as males fly, and for courtship activity to occur it appears necessary for both the male and female to have flown just previously. The most frequent stimulus releas- ing courtship in a male which has just flown is a female landing next to him (Barth, unpublished data). 2. Release of the female’s behavior The mounting and feeding behavior of females is released by the male sex pheromone in many cockroach species (Barth, 1968c). Such a male sex pheromone, “seducin,” was extracted by Roth and Dateo (1966) from males of Nauphoeta cinerea. There is some evidence for the existence of a volatile male sex pheromone in P. fulvescens. The frequent wing fluttering by isolated males and particularly the wing fluttering that follows unsuccessful copula- tion attempts suggests the function of dissemination of a male sex pheromone. This function has been suggested for wing fluttering in P. americana (Barth, 1970; Simon and Barth, in prep.) and for various vibration and trembling movements in various species of coackroaches (Roth and Hartman, 1967; Barth, 1968c). The function of flying in courtship situations remains a mystery, but male sex pheromone dissemination is a possibility. During the observation periods, there were occasions in which females approached males in a manner which appeared to be non- random and suggestive of an awareness of the male’s presence. The following procedure was followed to test for oriented locomotion in females in response to a source of volatile male sex pheromone. Prior to an observation period, a filter paper from a beaker contain- ing a single male was placed into the female side of the mating chamber on the side opposite to the location of the two females. Before the filter paper was introduced, the females were relatively quiescent showing some locomotion and slight antennal waving. After introduction of the paper, one female showed increased an- tennal activity and the other female started to locomote in the general direction of the paper, palpating the substratum as she moved. When she had progressed to within two inches of the paper, she turned directly toward it and came into antennal contact with the paper. She then stroked the paper lightly with her antennae, 1971] W endelken & Barth — Parcoblatta 327 drummed it rapidly with her maxillary palps, and walked across it. This first female initially made contact with the paper about 1.5 to 2 minutes after its introduction and had remained upon it for about a minute when the other female arrived. The second female touched the paper with her antennae and showed the same behavior toward it as the first female. Then there was some aggressive behavior between the two females. The second female drove the first one away and then proceeded to move around the paper, palpating it and waving her antennae gently. More work is clearly needed to confirm the hypothesis of volatility of the male sex pheromone in this species. Aggressive Behavior Two examples of male-female aggression were observed. In one, a female approached a male from in front of him. The male, in what appeared to be an aggressive gesture, jerked his head toward the female and she decamped. In the second case, a male and female made antennal contact, facing each other. The female lunged toward the male and then ran off. The male gave chase for a short distance. In addition to the aggressive female-female encounter described above, an observation was made in which two females were facing each other and antennal fencing. One female lunged toward the other and chased it away. Female aggression directed toward a copulating pair was observed in two cases. In the first case, a female (with protruding egg case) twice in rapid succession approached and jumped on top of the copulating pair — primarily on the dorsum of the copulating female — • and then glanced off rather rapidly. Later this female twice butted into the side of the copulating female but did not jump on it; the copulating female moved the pair several inches away. In the second case, a female antennally contacted a copulating pair and then, about a second later, charged toward the center of the pair and bumped them. The fact that aggression was never observed between males may very likely be due to the fact that only 2 or 3 were ever employed during observations, greatly decreasing the chances of interaction. Summary In Parcoblatta fulvescens , the volatile female sex pheromone plays a very prominent role in the release of the male’s courtship behavior. Olfactory reception of the female sex pheromone releases in males 328 Psyche [December an alert posture, increased antennal waving, and oriented locomotion toward the female. Flying and wing raising are also released by the female sex pheromone, no contact with females being required. Wing raising is quite variable in nature and is frequently accom- panied by lateral spreading with fluttering. Wing raising is fre- quently performed by males engaged in forward locomotion. The majority of male-female contacts are quite brief, unreceptive females rapidly decamping. When a male contacts a female, he raises his wings, turns away from the female, and backs. About 20 percent of the time, an intense concentration of female sex pheromone is apparently solely responsible for the release of wing raising, turning, and backing when the male has come close to a female but without contacting her. A receptive female mounts and feeds in a forward direction over the male’s exposed abdominal tergites. Tactile stimuli release the male’s copulatory thrusts when the female is two-thirds forward over the male’s abdomen. When the female reaches the vicinity of the first abdominal tergite, genital connection is achieved. The female then turns, resulting in the assumption of the opposed copulatory position. Evidence for the existence of a volatile male sex pheromone is presented. The function of pheromone dissemina- tion is suggested for the male’s wing fluttering and flying. Also included in this communication are some observations on aggressive behavior and the behavior of copulating pairs. References Barth, Robert H., Jr. 1961. Comparative and Experimental Studies on Mating Behavior in Cockroaches, Ph.D. Thesis, Harvard University, Cambridge, Massachusetts, 274 pages. 1964. The mating behavior of Byrsotria fumigata (Guerin) (Blat- tidae, Blaberinae). Behaviour 23: 1-30. 1968a. The mating hehavior of Gromphadorhina portentosa (Schaum) (Blattaria, Blaberoidea, Blaberidae, Oxyhaloinae) : an anomalous pattern for a cockroach. Psyche 75: 124-131. 1968b. The mating behavior of Eurycotis floridana (Walker) (Blat- taria, Blattoidea, Blattidae, Polyzosteriinae) . Psyche 75: 274- 284. 1968c. The comparative physiology of reproductive processes in cock- roaches. Part I. Mating behavior and its endocrine control. Advances in Reproductive Physiology 3 : 167-207. 1970. The mating behavior of Periplaneta americana (Linnaeus) and Blatta orientalis Linnaeus (Blattaria, Blattoidea, Blattidae, Blattinae) with notes on the mating behavior of three additional species of Periplaneta. Z. Tierpsychol. 27: 722-748. 1971] W endelken & Barth — Parcoblatta 329 Roth, Louis M. and R. H. Barth, Jr. 1967. The sense organs employed by cockroaches in mating behavior. Behaviour 28: 58-94. Roth, Louis M. and G. P. Dateo 1966. A sex pheromone produced by males of the cockroach, Nauphoeta cinerea. J. Ins. Physiol. 12: 255-265. Roth, Louis M. and H. B. Hartman 1967. Sound production and its evolutionary significance in the Blat- taria. Ann. Ent. Soc. Amer. 60: 740-752. Roth, Louis M. and E. R. Willis 1958. The biology of Panchlora nivea with observations on the eggs of other Blattaria. Trans. Amer. Ent. Soc. 83: 195-207. Simon, David and R. H. Barth, Jr. 1972. Sexual behavior in the cockroach genera Periplaneta and Blatta: Descriptive aspects, ms. in preparation. BARRIERS TO GENE FLOW IN NATURAL POPULATIONS OF GRASSHOPPERS II. MAINTENANCE OF NARROW HYBRID-ZONES BETWEEN MORPHS OF ARPHIA CONSPERSA ON BLACK MESA, COLORADO1 By Robert B. Willey and Ruth L. Willey2 In the previous publication of this series (Willey and Willey, 1967), we described the zoogeography of Arphia conspersa on the two sides of the Black Canyon of the Gunnison River in southwestern Colorado. North of the canyon the populations are monochromatic for red-orange wing color and south of the canyon the populations show a steep cline from west to east culminating in nearly 100% yellow-winged demes even though they often are less than one aerial mile from the orange-winged populations on the North Rim. We concluded that the sides of the canyon were an effective barrier to gene flow between the two rim populations and that the sedentary behavior and social cohesiveness of the adults (Willey and Willey, 1967 and 1969) probably slowed down lateral gene diffusion between the nearly adjacent demes of the South Rim. We have completed a preliminary survey of a narrow hybrid-zone on Black Mesa, an adjacent area which we previously have discussed briefly (Willey and Willey, 1967). The barriers seem to be related to suitable habitat and, in this case, we hope to show that the mixed demes are subject to periodic extermination, and perhaps are main- tained as a hybrid-zone by reinvasion from the neighboring mono- chromatic populations. Methods The census method is that of Willey and Willey (1967). Briefly, we walked in a non-repeating spatial pattern through the habitat and scored each insect as it flew up as either orange or yellow. An effort was made to count 100 individuals in each contiguous deme (200 preferably), but counts as low as 10 are reported in the protocol (Fig. 1 and 2). Nearly all the populations on the mesa were cen- sused at least once during the six years of study; several were cen- sused as many as ten times. If we felt the population could withstand Research was based at the Rocky Mountain Biological Laboratory, Crested Butte, Colorado; and X Lazy F Ranch, Crawford, Colorado. Address of the co-authors: Department of Biological Sciences, University of Illinois at Chicago Circle, Chicago 60680. Manuscript received by the editor January 18, 1972 330 1971] Willey & Willey — Populations of Grasshoppers 331 Plate 1. Oblique aerial view of Black Mesa from 30 miles south and 13,000 ft elevation, 15 September 1970. Route 92 (white line) marks the south edge of the mesa-top. Black Canyon is hidden behind Cimarron Hill and Fitzpatrick Mesa. the predation, we sampled it, capturing 50 individuals in a random manner for more careful analysis and as specimens for future ref- erence. A mark-recapture program on two adjacent zoogeograph- ically strategic populations served as a check on census results and for analysis of individual movements during the adult period of the life cycle (Willey et al., in preparation). In 1970, total capture, character scoring and release were instituted, because of the low population level in that year. The following United States Geological Survey 7-1/2-minute Topographic maps were used for the survey, plates and figure-codes: Big Soap Park, Cathedral Peak, Cimarron, Crawford, Curecanti Needle, Little Soap Park, Sapinero, and X Lazy F Ranch, extending from ioj° 37*30" West and 38°37'3o" North to io7°i5/ West Longitude and 38°22'3o" North Latitude. Observations Black Mesa. This mesa is a well-defined plateau capped by volcanic rocks, principally andesitic breccias and rhyolitic welded tuffs of Oligocene age (Hansen, 1971, and pers. comm.). It slopes upward from 9,000 ft on the southern rim to over 11,000 ft at the north- eastern end. Its southern rim forms part of the North Rim of the 332 Psyche [December Black Canyon of the Gunnison River. This area is approximately 35 square miles and the mesa top stands 1000 to 2000 ft above the surrounding valleys. Intermittent streams dissect the surface into four major north-south gulches draining into the Gunnison River and a major gulch drains each of the east and west sides of the mesa (Plates i and 2). Private ranches make up about 10% of the area; 10% is the Black Mesa Experimental Forest and Range, adminis- tered jointly by the United States Forest Service and Colorado State University at Fort Collins; the rest is in the Gunnison National Forest. According to William Knott, a lifelong resident and forest ranger, the phytotopography of the mesa has not changed since settlement in 1880, except that the common dandelion (Taraxacum officinale) was not present until 1920. Boundaries of grasslands and forested tracts are largely unchanged, except for the results of the Indian Wars in 1870-80 during which several extensive forest tracts were burned by the Indians as a “scorched earth” policy. These burned areas and a few aspen groves with burned or sawed spruce stumps are easily distinguished from the virgin forests. The Indians used the mesa primarily as a hunting ground during the summer and several localities have numerous artifacts scattered over the surface as evidence of repeated summer encampments. Although the United States Geological Survey Topographic Maps were made from aerial photographs taken in 1955, the boundaries of chaparral, forest, and grasslands are accurate to minute detail even at the time of this writing (1971). The only changes are a small acreage of timber sales, construction of the Morrow Point and Blue Mesa Dams and Reservoirs within the Black Canyon itself, and the clearing for high tension power lines across the southwest corner of the Mesa. The major forest component is Engelmann spruce (Picea engel- manni), alpine fir (Abies lasiocarpa) and quaking aspen (Populus tremuloides) in mixed or monotypic stands (Tietjen et al ., 1967). Upland meadows, which are the primary habitats for Arphia , contain the bunch grasses Festuca thurberi , F. idahoensis and Stipa letter- mani plus the forbs Geranium fremontii , Chrysopsis villosa, Erigeron macranthus , Lathyrus leucanthus , Agoseris spp. and the shrubs Chrysotkamnus parryi and Potentilla fruticosa as the dominant plants in terms of herbage production (Paulsen, 1969). As can be seen from the map (Plate 2), the grasslands follow the drainages and were probably maintained as grasslands by the activities of the once 1971] Willey & Willey — Populations of Grasshoppers 333 numerous beaver. The forests form a barrier between each drainage. There are many upland meadows surrounded by forest forming a natural system of isolated habitats for grassland-dependent animal species such as Arphia conspersa. Below 9300 ft elevation, we have determined that sage brush (Artemesia tridentata) becomes a domi- nant shrub and mixed with oak brush (Quercus gamhellii) , service- berry ( Amelanchier pumila) , mountain mahogany ( Cercocarpus mon- tanus) and chokecherry (Prunus melanocarpa) form a dense chapar- ral on the edges and slopes around the mesa. The climate is subalpine with heavy and continuous snow cover 5 to 6 ft deep, usually from November through April (William Knott, weather station records and personal communication). An- nual precipitation is quite variable; X from 1956 to I971 is 29.7 inches (S.D. = 6.6), two-thirds falling as snow. The snow pack is variable in length; X from 1956 to 1971 is 203 days (S.D. ^ 17.5). Insolation is high during May, June, and July with our Belfort pyrheliograph reading as high as 1.6 cal/cm2/min. Air tem- peratures during the summer months range from freezing at night to 25 °C during the day. The area is subject to unseasonal snow- storms, periods of heavy rainfall and alternating saturated and dried-out soil. For example, on 26 June 1969, there was a wet snow- fall of 6 to 12 inches. On 12 October 1969, a wet snowfall of 24 inches followed a week of heavy rains wdiich saturated the soil. This snowfall formed the basis for a continuous dense snowpack above the 9000 ft level until 15 May 1970. This early and continuous snow- pack (216 days), just within one standard deviation, and long water saturation of the soil was unique in the memory of William Knott, whose family has kept precipitation and temperature records on the mesa since 1905. The snow pack of 1 970-71 was equally long, but started as a dry, cold snowfall. Again on 12 June 1970, there was a snowfall of 14 inches, which melted within a couple days. How- ever, the snowpack records of the climatological station show that these last two winters were not the longest packs on record ; that of 1956-57 lasted 247 days until 17 June and was equivalent to 30 inches of liquid water on 12 April. That year also was the wettest on record since 1905, 48 inches of precipitation (William Knott, personal communication). The effect on Arphia populations is un- known, since we began our studies in 1964. The vagaries of climatic conditions of this high plateau may be an important factor in the maintenance of steep polymorph dines in the A . conspersa populations. 334 Psyche [December Arphia conspersa. For a full discussion of the life history and be- havior of this oedipodine species, refer to our previous papers (Wil- ley and Willey, 1967 and 1969). The species is spring-brooded with the nymphs (hoppers) overwintering during the third or fourth in- star. The populations seem to be formed of loosely interacting social colonies which tends to reduce vagility. Their habitat preference seems to be a short narrow-leaved grassland with enough bare ground for courtship. Their altitudinal limit depends on slope and exposure and not yet understood limiting factors, but is between 970010,300 ft at the latitude of Black Mesa (38°3o' =+= 7'3o" N.). The species is seldom found in dense tail-grass meadow, thick aspen or spruce-fir forest, dense chaparral or in irrigated pastures. Nor is it likely to be found on extensive bare rock or freshly disturbed areas. It seems to prefer serai edges and, despite its apparent lack of vagility, seems to be an opportunistic species, often appearing in regrown road cuts and chained (cleared) chaparral/grasslands within 5 years if estab- lished adjacent populations are available (Locality #6ob, from Buck- horn Gulch #5ia). However, Alexander and Hilliard (1969) list this species as a non-adventive species, seldom occurring in zones where it is not resident as nymphs, e.g., alpine zones and the timber- line ecotone. Zoogeography. Plate 3 shows the location and graphed wing-color proportions of each major deme on and adjacent to Black Mesa. Figure 1 is the protocol of each deme. We must emphasize that the proportions refer only to phenotypes, since the genetic analysis is still under investigation. Nevertheless, it is striking that phenotype proportions are quite similar within a gulch and may differ by 25% from the adjacent gulch. From the west, the only possible path of invasion to the mesa top is along the slopes of Long Gulch on the southwest or up Crystal Creek on the northwest. The nearest nearly 100% orange-winged population is #52(a) [Fig. 2] which is sep- arated from the nearest Long Gulch demes (60% orange-winged) by a virgin spruce-fir forest only one-half mile thick. Northward, the nearest Crystal Creek deme is #25 on the 9200 ft contour. The passage onto Black Mesa rises to 10,500 on Powell Ridge and is, from all appearances, a suitable meadow and cut-over forest habitat, but presently that area is uninhabited by Arphia. Between Long Gulch and Mesa Creek are thin strips of aspen- spruce forest (some of which were lumbered 30-40 years ago) and very dense growths of Festuca thurberi. This latter is such a sufficient barrier that demes #53a (the Mounds) and #53b (the Forks) maintain proportions of 35-40% yellow and 60-75 % yellow re- 1971] Willey & Willey — Populations of Grasshoppers 335 spectively, even though they are only one-half mile apart. These demes were intensively studied during 1968-71 by mark-recapture methods (Willey et al., in preparation) and no interchange was noted nor were any individuals found in the interspace. This patch- iness of habitat utilization is characteristic of the Black Mesa demes. Figure 1 is an accurate appraisal of the number of demes we were able to find on Black Mesa during eight years of study. From Mesa Creek eastward, a slight drop in average proportions is seen in Corral Gulch and then a sudden drop to i%-4% in Myers Gulch. This phenotypic cline is two miles wide and there are several grassland corridors between the gulches through which gene flow could be accomplished. We have analyzed by census, line transect, and sampling those demes and subdemes which occur in the trans-mesa corridors. These are best exemplified by # 54a, and b in one series (Experimental Pastures #5 and 6) and # 54c, 54d, and 55a (the Transect). It is noteworthy that the corridor demes are, among themselves, a co- hesive unit and, instead of showing a gradual cline from low yellow in the west to high yellow in the east, the break is between the east- ernmost corridor demes and the adajacent Myers Gulch deme. This may be the result of a zone of uninhabited grassland on the western portion of Myers Gulch, and the only demes we have found in the gulch are those shown in Plate 3 on the eastern side (#450, 55b, 64a, and 64b). Several peculiar relationships are indicated by Figs. 1 and 2. Although the major demes seem relatively stable in wing-color pro- portions, the smaller subdemes of the trans-mesa corridors differ in proportions from one census to another. For example, Pasture 6 (#54b) varied from 9% to 34% orange over three years. There is no pattern in this variation, and the total numbers are nearly the same for each census. There is no geographic pattern (geographic subdeme records are in Fig. 2), nor is there a pattern of morning and afternoon differences in the censuses. On 29 June 1968, a morning census showed a concentration of six orange males in one area of #54b. These males could not be found on 1 July. The evidence suggests a transitory clustering of orange individuals in small areas which could skew censuses drastically if a group is missed, or if there is a single case of differential predation. However, a. mapping program on the “Mounds” ( #53a) disclosed no differ- ence in the dispersion pattern of distribution among any of the pheno- type classes (Willey et al ., in preparation). 336 Psyche [December Plate 2. Collage of CJ. S. Geological Survey 7 1/2 Minute Topographic Maps (Cathedral Peak, Cimarron, Curecanti Needle and X Lazy F Ranch), showing Black Mesa and environs. Compare with PI. 3 on the facing page to see the relationships between A. conspersa deme structure and phyto- topography. 1971] Willey & Willey — Populations of Grasshoppers 337 Plate 3. Same collage as in PI. 2 showing most of the censused localities and their relative proportions. For explanation see text, page 348. 338 Psyche [December The “Transect” (#44a, 54c, 55a) also exhibits this clustering phenomenon. In this case, we were able to map the orange-winged clusters in 1968 and again in 1969. They were present in the same positions both years. We then ran two transects, each 1 mile long, through the area (Hill, 1969, unpublished data) to ascertain any dispersion from these clusters. None seemed to occur. Uniformly, the clusters of orange-winged insects occurred on mounded elevations of pasture with Fesluca thurberi providing slightly greater cover than in the areas more densely populated with yellow-winged individuals. These results indicated three possible causes : 1 ) the orange-winged phenotype has some pleimorphic preference for denser habitat, 2) there is a low vagility of individuals during the entire life span and they seldom wander far from the original hatching point, or 3) the pheno- types have reached some level of behavioral or ecological separation. We currently are investigating the clustering patterns of these pop- ulations. It is clear that the only invasion route for yellow-winged pheno- types is from southern Curecanti Canyon and from the slopes of the South Rim of the mesa via the major gulches. All other avenues are blocked by the forest. We should emphasize the habitat type, the size, and the elevation of the northernmost denies on the mesa. Num- ber 45b is a tiny deme of 8 to 20 individuals which survives around an old sawdust pile — all that remains of a lumber mill. A few more individuals can be found in an adjacent burned-over and logged area, but this area is primarily an early sere characterized by broad- leaved plants, brambles, and few grasses. Number 35 is slightly larger and has several small subdemes. The deme inhabits a cleared private pasture at 10,300 ft, which enjoys a southeastern exposure. The floral phenology is similar to that of areas 500 ft lower. The “Burn Area” contains large subdemes (#45d) which have colonized an extensive burned-over area. After 85 years, this soil still can support only scattered bushes ( Ribes lacustre and S ambucus sp .) and small, sparse patches of Festuca idahoensis , F. thurberi , Stipa columbiana , S. lettermani and Blepharoneuron tricholepis, which pro- vide good though discontinuous habitat for A. conspersa. However, the floral phenology is much delayed over that in other habitats and indicates continuous snow cover until 15 May or later. Demes in higher altitudes farther north have been indicated in some years by sighting one or two males, a female, or hearing some crepitations. Whether these are founding colonies which succeed only for a year or two is not known. In two of these cases they were found in old 1971] Willey & Willey — Populations of Grasshoppers 339 lumber mill sites near sawdust heaps and in one case the male was found in an old homestead foundation. In 1970, populations over the entire mesa above 9200 ft crashed. In most areas, no Arphia could be found. In denies #5 3a> b and 54a, between 8 and 41 individuals were found in a total capture survey, about 1 to 15 percent of the expected population level. Of these, nearly one-half had blebs on the pronotum, an anomaly found in only 2% of normal populations. Lower altitude populations were very nearly normal in density and high counts were made of # 5°> 51a, 82 and the North Rim of the Black Canyon National Monu- ment (over 600), all occurring between 7700 and 9100 ft elevation. Similar crashes in usually dense populations occurred 40 miles north at Gothic (9500 ft, Gunnison County), and Jack’s Cabin Cut-Off (9000 ft, Gunnison County). On the other hand, we have noted and collected sufficient nymphs of A. conspersa (over 200 in several areas) in August and September, 1970, to convince us that the diapausing eggs of the 1969 adult brood were not adversely affected by the winter and hatched normally in the summer of 1970- In 1971, the populations were normal in abundance, though not as high in number as those of 1969. The mark-recapture program on the Mounds (#53a) showed a population of 400, whereas in 1970 only 15 adults were found. In September, 197 1, only one nymph was found in this area, two nymphs in the Burn Area (#45d, 100 nymphs in 1970), and 16 nymphs in Pasture 5 (#54a, 41 adults in 1970, over 200 adults in 1971). This small number of nymphs in 1971 indicates no recovery of the 1970 brood. Discussion Black Mesa is subject to severe vagaries of weather. Since the snow packs of 1969-70 and 1970-71 were nearly the same length (216 and 215 days respectively), and the 1971 adult brood was only a little below normal in density, we think that conditions before the snowpack of 1969-70 or after it had melted probably caused the pop- ulation crash of 1970. These conditions could have been 1) the snowfall of 26 June 1969, which may have killed hatching nymphs; 2) the wet snowfall and rains of early October, 1969, which could have harmed cold-immobilized nymphs; 3) the snowfall of 12 June 1970, which could have killed emerging adults, and 4) an undetected late freeze which could have decimated freeze-sensitive nymphs emerg- ing from hibernation in May. The fact that half of the surviving adults had developmental anomalies, especially blebs on the prono- tum, indicates some sort of post-dormancy damage. However, the 340 Psyche [December specific causes for the widespread failure of the 1970 adult brood are still uncertain and probably were cumulative. In a simpler case, Ehrlich et al. (1972) have noted a similar series of extinctions of the butterfly Glaucopsyche lygdamus at these altitudes in this same general area which they prove was due to destruction of oviposition sites and larval food plants by the snowfall of 26 June 1969. Consequences of the failure of the 1970 A. conspersa brood are the problem of subsequent recruitment and the entire question of the maintenance of the Black Mesa hybrid zone. It will be very in- structive to follow the reestablishment of the Arphia demes from the few surviving centers of the 1970 brood, from the off -mesa populations, and perhaps from the odd-year brood. Certain propor- tion discrepancies, such as those shown by #53b (the Forks), #J2b (Corral Gulch), and #45d(c) (southeast meadow of the Burn Area) seem related to alternate years. Since we have proved in the present results that the Black Mesa populations overwinter as egg and nymphs, there may be two geneticalty independent populations, one reaching adulthood on even-numbered years and one on odd years. However, further yearly censusing of strategic populations is necessary to determine the validity of this assumption and to deter- mine the amount of temporal crossing-over by any non-diapausing individuals. The narrowness of the Black Mesa hybrid-zone has puzzled us, but now it seems clear that periodic extermination of the mesa-top populations could afford a considerable setback to any extensive gene flow. Indeed, the presently inhabited portion of the mesa probably has been colonized by A. conspersa only within the past 150 years since the beginning of the last climatic warming trend after the Neoglaciation (Remington, 1968, p. 350; and Richmond, 1965) which probably had made most of the mesa-top uninhabitable. Any long term cooling trend resulting in increased precipitation, numbers of late frosts or snows, etc., could again cause the mesa to become an altitudinal barrier. It is also probable that the 1970 population crash is not a unique occurrence on the mesa, even during this present hypsothermal period. If the present zoogeographic pattern of wing-color variation is the result of recent colonization, it is probably safe to assume that the connecting corridors and forest barriers are at least as old and we can describe original invasion patterns in terms of the present topography. However, with current pressures on the National Forest Service to release tracts for lumbering, the original proportions of vegetational associations, the climatic patterns and even the water-holding capacity 1971] Willey & Willey — Populations of Grasshoppers 341 of the mesa will be changed. It is well to have a baseline for evaluat- ing the changes which will occur in the zoogeography of this zone of hybridization. It should be instructive to examine this area soon for evidence of other species- and morph-pairs in the process of hy- bridization, since climatic conditions and geography at present seem ideal for the development of a common “suture-zone” between spe- cies-pairs of diverse organisms (Remington, 1968), especially between those which are spring-brooded, ground-inhabiting poikilotherms. Conclusions On Black Mesa, Colorado, the grasshopper /lrphia conspersa ex- hibits a phenotypic cline in wing-color variation which separates 100% orange-red denies from 100% yellow demes by two to five miles. The cline seems to be maintained by three factors at least : 1 ) limited vagility of the individuals in a deme, 2) unsuitable habitat between the geographically discontinuous populations, and 3) periodic, vir- tually complete, exterminations of the hybrid demes by climatological catastrophes related to the fact that the mesa-top is at the altitudinal limit for the species. Summary Arphia conspersa varies in wing color on Black Mesa in southwest- ern Colorado. The phenotypes exhibit an orderly but steep step-wise cline from 100% orange and red to 100% yellow over a distance of only two to five miles, ascertained by sight-census and mark-recapture methods over a period of eight years. Black Mesa slopes gradually from 9000 ft elevation in the south to 11,000 ft in the north. Since A. conspersa has successfully colonized only a few areas above 1 0,000 ft, this also affords an opportunity to observe the ecology of a species at its altitudinal limit. In 1970, 50 demes which had been located on the mesa were vir- tually exterminated, probably by unseasonal weather conditions prior to and after a prolonged winter snow pack lasting more than 8 months. However, these high-altitude populations have a two-year life cycle with both eggs and nymphs overwintering. The alternate- year brood, which had over-wintered eggs in 1969-70, was unaffected and matured normally in 1971. Census of nymphs in the fall of 1971 indicated no recovery of the nearly exterminated brood. This catas- trophe strongly suggests that maintenance of this narrow hybrid-zone depends not only on the barriers produced by unsuitable habitat, but also on periodic extermination of the mixed populations. It also indi- cates the survival value of a two-year life cycle at the altitudinal limits of a species. 342 Psyche [December OO rf- 00 ON ON VO vo vO O O 0)0 0 QJ ZL >>>>>>>>>> >> >>> >>>>> >> OOOONOOOCMtOoot^i/Nm moo N N K -a- in no !>. 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PJ « bfl “ 2 6 Oh -re _o -d c 03 J3 V 33 i-s? rg cs C <-P • - ° ^ bC ° .2 Corrected from Protocol Fig. 2, Entry #31, Willey and Willey (1967). *Area censused in 1970, but total sighting < 10. **Area recensused > 2 times, never more than 5 sightings. 348 Psyche [December Acknowledgements We acknowledge the help of our assistants, undergraduate research participants and volunteers in censusing: David Ballestas, David Borg, William Davis, Sharon Ginsburg, Anne Hill, Dennis Johns, James Karpus, Gordon and Muriel Kerr, Ellen Levy, Jaclin Lewbin, David Messenger, David Mucha, Gerald Mussgnug, Marianne Nied- zlek, Ronald Outen, Frances Pivorunas, Joyce Redemsky (Mrs. Gregg Young), Wayne Schennum, Melvin Shemluck, Frank Slansky, Darelyn Weber (Mrs. Charles Handley), and David Werner. We thank Dr. Wallace Hansen, U.S. Geological Survey, Denver, for critically reading the manuscript; and Dr. Paul Ehrlich, Stanford University, for giving us a copy of the manuscript of Ehrlich et al. (1972). William Knott, Forestry Research Technician, Black Mesa Experimental Forest, has been most informative in the history, floristics, and climatology of Black Mesa. Dr. Raymond Price, former Director of the Rock Mountain Range Experiment Stations, allowed us to study populations in the experimental pastures and sent us reprints of their studies of the Black Mesa. We are grateful for the hospitality of George Klaitch, Manager of the X Lazy F Ranch, and the Colorado Synod of the Presbyterian Church for allowing us living and laboratory space at the ranch. The Forest Service land-use maps (T50N-R5-5-1/2W, T49N-R5-5-1/2W, and T49N-R6W), showing recent and projected timber sales, were supplied by R. M. Case, District Ranger. David Mucha’s photographic assistance on Plates 2 and 3 is gratefully received. Mrs. Mary Grant retyped several revisions of this paper. This study was partially supported by Grant GB-2201 from the National Science Foundation, four grants from the University of Illinois Graduate Research Board, two grants from the Society of the Sigma Xi (1963 and 1968), National Science Foundation Pre- doctoral Fellowships to Wayne Schennum, and to James Karpus, and National Science Foundation Undergraduate Research Participation fellowships to William Davis, Sharon Ginsburg, William Hill, Ger- ald Mussgnug, Marianne Niedzlek, Ronald Outen, Melvin Shem- luck and Joseph Wroblewski. Explanation of plates 2 & 3 Plates 2 and 3 are quadrated by the Universal Transverse Mercator Grid (Zone 13) and in PI. 3 every linear 2.5 km is numbered on the ordinate and the abscissa. The combined number (ordinate and abscissa respectively) produces the code number of each quadrat and its demes (Figs. 1 and 2). The lower case letters distinguish demes within a single quadrat. If a deme occurs on a grid line, the locality is numbered according to the code number 1971] Willey & Willey — Populations of Grasshoppers 349 of the quadrat to the south or west of the line (see quad. No. 32). Mer- cator lines 285 km East and 4265 km North are printed also on PI. 2. The U.S. Geoglogical Topographic Maps were drawn from aerial photo- graphs taken between 3 September and 27 September 1955. The light colored areas are grassland, grey is chaparral and dark is forest. Literature Cited Alexander, G. and J. R. Hilliard, Jr. 1969. Altitudinal and seasonal distribution of Orthoptera in the Rocky Mountains of Northern Colorado. Ecol. Monogr., 39: 385-431. Ehrlich, P. R., D. E. Breedlove, P. F. Brussard, and M. A. Sharp 1972. Weather and the “regulation” of subalpine populations. Ecology, 53(1): in press. Hansen, W. R. 1971. Geologic map of the Black Canyon of the Gunnison River and vicinity, Western Colorado. U.S. Geol. Survey, Misc. Geol. In- vest.: Map 1-584. Paulsen, H. A., Jr. 1969. Forage values on a mountain grassland-aspen range in western Colorado. J. Range Management, 22: 102-107. Remington, C. L. 1968. Suture-zones of hybrid interaction between recently joined biotas. Evol. Biol. 2: 321-428. Richmond, G. M. 1965. Glaciation of the Rocky Mountains. In Wright, H. E. and D. G. Frey (Editors), The Quaternary of the United States (Prince- ton, New Jersey: Princeton University Press, pp. 217-230). Tietjen, H. P., C. H. Halvorson, P. L. Hegdal and A. M. Johnson 1967. 2, 4-D herbicide, vegetation, and pocket gopher relationships — Black Mesa, Colorado. Ecology, 48: 634-643. Willey, R. B. and R. L. Willey 1967. Barriers to gene flow in natural populations of grasshoppers. I. The Black Canyon of the Gunnison River and Arphia con- spersa. Psyche, 74: 42-57. 1969. Visual and acoustical social displays by the grasshopper Arphia conspersa (Orthoptera: Acrididae). Psyche, 76: 280-305. Willey, R. B., G. Mussgnug, J. Wroblewsky and P. Wussow A study of the movement within a population of Arphia con- spersa (Orthoptera: Acrididae). In Preparation. PSYCHE INDEX TO VOLUME 78, 1971 INDEX TO AUTHORS Alcock, J. The Behavior of a Stinkbug, Euschistus conspersus Uhler (Hemiptera: Pentatomidae) . 215 Barr, T. C. Jr. Micratopus Casey in the United States (Coleoptera: Cara- bidae: Bembidiinae) . 32 Barr, T. C., Jr. Trechoblemus in North America, with a Key to North American Genera of Trechinae (Coleoptera: Carabidae). 140 Carpenter, F. M. and E. S. Richardson, Jr. Additional Insects from Penn- sylvanian Insects from Illinois. 267 Chickering , A. M. The Genus Oonops (Araneae, Oonopidae) in Panama and the West Indies. Part 2. 203 Evans, H. E. The Larva of Heliocausus Iarroides (Hymenoptera, Spheci- dae). 1 66 Erickson, J. M. The Displacement of Native Ant Species by the Introduced Argentine Ant, Iridomyrmex humilis Mayr. 257 Haskins, C. P. and Paul A. Zahl. The Reproductive Pattern of Dinoponera grandis Roger (Hymenoptera, Ponerinae) with Notes on the Ethology of the Species. 1 Hlavac, T. F. Differentiation of the Carabid Anetnna Cleaner. 51 Jackson, R. R. Fine Structure of the Thread Connections in the Orb Web of Araneus diadematus. 12 Kelsey, L. P. A New Scenopinidae (Diptera) from Bermuda. 49 Kukalovd-P eck, J. The Structure of Dunharia (Palaeodictyoptera ) . 306 Levi, H. IV. The Orb-Weaver Genera Singa and Hypsosinga in America (Araneae: Araneidae). 229 Moxey, C. F. Notes on the Phasmatodea of the West Indies: Two New Genera. 67 Powell, J. A. and J. M. Burns. Colonization of the Northeastern United States by Two Palearctic Moths (Lepidoptera : Tortricidae) . 38 Reiskind, J. The South American Castianeirinae. I. The Genus Psellocoptus (Araneae: Clubionidae) . 193 351 Robinsan, M. H. and H. Mirick The_ Predatory Behavior of the Golden- Web Spider Nephila clavipes (Araneae: Araneidae). 123 Roth, L. M. The Male Genitalia of Blattaria. VI. Blabetidae: Oxyhaloinae. 84 Roth, L. M. The Male Genitalia of Blattaria. VII. Galiblatta, Dryado- blatta, Poroblatta, Colapteroblatta, Nauclides, Notolampra, Litopeltis, and Cariacausia (Blaberidae: Epilamprinae) . 180 Roth, L. M. The Male Genitalia of Blattaria. VIII. Panchlora, Ancho- blatta, Biolleya, Pelloblatta, and Achroblatta. (Blaberidae: Panchlori- nae). 296 Rovner, J. S. Mechanisms Controlling Copulatory Behavior in Wolf Spiders (Araneae: Lycosidae). 150 Sheldon, J. K. and E. G. MacLeod. Studies on the Biology of the Chryso- pidae. II. The Feeding Behavior of the Adult of Chyrysopa carnea (Neuroptera) . 107 Talbot, M. Flights of the Ant Formica dakotensis Emery. 169 W endelken, P. and R. H. Barth. The Mating Behavior of Parcoblatta fulvesccns (Saussure & Zehntner) (Blattaria, Blattellidae) . 319 fVilley, R. B. and R. L. fVilley. Barriers to Gene Flow in Natural Popula- tions of Grasshoppers. II. Maintenance of Narrow Hybrid-Zones be- tween Morphs of Arphia conspersa on Black Mesa, Colorado. 330 352 INDEX TO All new genera, new species and new Achroblatta, 296 Additional Insects in Pennsylvania Concretions from Illinois, 267 AGAMENON IPHIMEDEIA, 75 Anchoblatta, 296 Araneus diadematus , 12 Arphia conspersa, 330 Barriers to Gene Flow in Natural Populations of Grasshoppers. II. Maintenance of Narrow Hydrid- zones Between Morphs of Arphia conspersa on Black Mesa, Colo- rado, 330 Behavior of a Stinkbug, Euschitus conspersus Uhler (Hemiptera: Pentatomidae, 215 Biolleya, 296 Carabid antenna cleaner, 51 Chrysopa carnea, 107 Clepsis unifasciana, 38 Colonization of the Northeastern United States by the Two Palae- arctic Moths, 38 Croesia forskaleana, 38 Differentiation of the Carabid An- tenna Cleaner, 51 Dinoponera grandis, 1 Displacement of Native Ants Species by the Introduced Argentine Ant, Iridomyrmex humilis Mayr, 257 Dunbaria, 306 Euschistus consperus , 215 Fine Structure of Thread Connec- tions in the Orb Web of Araneus, 12 SUBJECTS names are printed in capital type. Flights of Formica dakotensis, 169 Formica dakotensis, 169 Heliocausus larroides, 166 HERDINA MIRIFICUS, 291 HERDINIDAE, 287 Hypsosinga alberta, 254 Larva of Heliocausus, 166 LYCODEMAS ADOLESCENS, 268 Male Genitalia of Blattaria, 84, 180, 319 Mating Behavior of Parcoblatta ful- vescens (Saussure & Zehntner) (Blattaria, Blattellidae) , 319 Mechanisms Controlling Copulatory Behavior in Wolf Spiders, 150 Microtopus Casey in the United States, 32 Nephila clavipes, 123 New Scenopinidae from Bermuda, 49 Notes on the Phasmatodea of the West Indies, 67 NOTORACHIS WOLFFORUM, 273 Oligotypus makowskii, 286 Oonops balanus, 204 Oonops castellus, 206 Oonops GERTSCHI, 211 Oonops in Panama and West Indies, 203 Oonops ronoxus, 212 Orb-weaver Genera Singa and Hypsosinga in America (Araneae: Araneidae), 229 353 Panchlora, 296 Parcoblatta, 319 Pelloblatta, 296 Predatory Behavior of Nephila clavipes, 123 Psellocoptus bucklii, 200 Pseliocoptus prodontus, 201 Reproductive Pattern of Dinoponera, 1 Scenopinus bermudaensis, 49 Singa eugeni, 236 South American Castianeirinae, 193 Spilaptera Americana, 281 Structure of dunbaria (Palaeodicty- optera), 306 Studies on the Biology of the Chrys- opidae, 107 TARAXIPPUS PALIURUS, 70 T rechoblemus westcotti, 142 354 CAMBRIDGE ENTOMOLOGICAL CLUB A regular meeting of the Club is held on the second Tuesday of each month October through May at 7:30 p.m. in Room B-455, Biological Laboratories, Divinity Ave., Cambridge. Entomologists visiting the vicinity are cordially invited to attend. The illustration on the front cover of this issue of Psyche is a reproduction of the drawing by James W. Chapman of the Smaller Elm Bark Beetle, Scolytus multistriatus. This was included in his account of the discovery of this insect in the elm trees of the Harvard Yard, in October, 1909 — the first report of this European insect in the United States (Psyche, 1910, Vol. 17, pi. 4). BACK VOLUMES OF PSYCHE The Johnson Reprint Corporation, 111 Fifth Avenue, New York, N. Y. 10003, has been designated the exclusive agent for Psyche, volumes 1 through 62. Requests for information and orders for such volumes should be sent directly to Johnson Reprint Corporation. Copies of issues in volumes 63-78 are obtainable from the editorial offices of Psyche. Volumes 63-78 are $6.00 each. F. M. Carpenter Editorial Office, Psyche, 16 Divinity Avenue, Cambridge, Mass. 02138. FOR SALE Classification of Insects, by C. T. Brues, A. L. Melander and F. M. Carpenter. Published in March, 1954, as volume 108 of the Bulletin of the Museum of Comparative Zoology, with 917 pages and 1219 figures. It consists of keys to the living and extinct families of insects, and to the living families of other terrestrial arthropods; and includes 270 pages of bibliographic references and an index of 76 pages. Price $9.00 (cloth bound and postpaid). 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